The Ecology of Papua: Part One [6, 1 ed.] 0794603939, 9780794603939

Table of contents :
Intro
Contents
Abbreviations
Contributors
Acknowledgements
Foreword
Preface
Letter from the Papuan People's Assembly
Section I: Introduction to Papua
1.1. Introduction to Papua
1.2. Biological Exploration of New Guinea
1.3. The Socio-cultural Plurality of Papuan Society
1.4. Prehistoric Human Presence in Papua and Adjacent Areas
Section II: The Physical Environment
2.1. Tectonic Geology of Papua
2.2. Soils of Papua
2.3. Climate of Papua
2.4. Papuan Terrestrial Biogeography,
with Special Reference to Birds
2.5. Freshwater Biogeography of Papua
2.6. Paleontology of Papua
2.7. Paleoecology and Paleoenvironments of Papua
Section III: The Flora
3.1. Introduction to the Flora of Papua
3.2. Lichen Biodiversity in New Guinea
3.3. Bryophytes of Papua New Guinea: Their
Diversity, Ecology, and Conservation
3.4. Ferns and Lycophytes of Papua
3.5. Gymnosperms of Papua
3.6. Angiosperms
Section IV: The Fauna
4.1. Introduction to the Fauna of Papua
4.2. Marine Invertebrates of Papua
4.3. Insects of Papua
4.4. Cicada Endemism in Papua
4.5. Marine Wood-Boring Invertebrates of New
Guinea and Its Surrounding Waters
4.6. Herpetofauna of Papua
4.7. The Monitor Lizards of Papua
4.8. Fishes of Papua
4.9. Birds of Papua
4.10. A Taxonomic and Geographic Overview of the Mammals of Papua

Citation preview

EcologyOfPapua Vol.1

7/11/07

9:58 AM

Page 1

THE ECOLOGY OF INDONESIA SERIES Volume VI

The Ecology of Papua Part One

advisory board members Gerald R. Allen Allen Allison Chris Ballard Bruce M. Beehler James B. Cannon Yance de Fretes Geoffrey S. Hope Robert J. Johns J. R. Mansoben Scott E. Miller Dan A. Polhemus Wayne N. Takeuchi Tony Whitten

................. 16157$

$$FM

02-08-07 10:20:38

PS

PAGE ii

the ecology of indonesia series Volume VI: The Ecology of Papua, Parts One and Two Other Titles in the Series Volume I: The Ecology of Sumatra Volume II: The Ecology of Java and Bali Volume III: The Ecology of Kalimantan Volume V: The Ecology of Nusa Tenggara and Maluku Volume VII: The Ecology of the Indonesian Seas, Part One Volume VIII: The Ecology of the Indonesian Seas, Part Two

................. 16157$

$$FM

02-08-07 10:20:38

PS

PAGE iii

................. 16157$

$$FM

02-08-07 10:21:06

PS

THE ECOLOGY OF INDONESIA SERIES Volume VI

The Ecology of Papua Part One Andrew J.Marshall Bruce M.Beehler

For our families: Brenda, Philip, and Will; Carol, Grace, Andrew, and Cary Published by Periplus Editions (HK) Ltd. with editorial offices at 61 Tai Seng Avenue #02-12, Singapore 534167 First edition 2007 © Conservation International Foundation, 2007 All rights reserved. ISBN: 978-1-4629-0679-6 (ebook) Publisher: EricOey Typesetting and graphics: Coghill Composition, Richmond, Virginia, USA Copyediting: Anne McGuire Design: Ann Twombly Distributors North America, Latin America, and Europe: Tuttle Publishing 364 Innovation Drive North Clarendon, VT 05759-9436 U.S.A. Tel: 1 (802) 773-8930; Fax: 1 (802) 773-6993 [email protected] / www.tuttlepublishing.com Japan: Tuttle Publishing Yaekari Building, 3rd Floor 5-4-12 Osaki; Shinagawa-ku, Tokyo 1410032 Tel: (81) 3 5437-0171; Fax: (81) 3 5437-0755 [email protected] Asia Pacific: Berkeley Books Pte. Ltd. 61 Tai Seng Avenue #02-12, Singapore 534167 Tel: (65) 6280 1330; Fax: (65) 6280 6290 [email protected] / www.periplus.com Indonesia: PT Java Books Indonesia Kawasan Industri Pulogadung JI. Rawa Gelam IV No. 9, Jakarta 13930 Tel: (62) 21 4682-1088; Fax: (62) 21 461-0206 [email protected]

Pr inted in Hong Kong 10 09 08 07

5 4 3 2 1

Publication of this book would not have been possible without the generous support of BP and the Gordon and Betty Moore Foundation.

Contents Abbreviations Used in These Volumes xi Contributors xix Acknowledgments xxiii Foreword xxvii edward o. wilson Preface xxix Letter from the Papuan People’s Assembly xxxi SECTION ONE. INTRODUCTION TO PAPUA 1.1. Introduction to Papua 3 bruce m. beehler 1.2. Biological Exploration of New Guinea 14 david g. frodin 1.3. The Socio-cultural Plurality of Papuan Society 108 j. r. mansoben 1.4. Prehistoric Human Presence in Papua and Adjacent Areas 121 juliette pasveer SECTION TWO. THE PHYSICAL ENVIRONMENT 2.1. Tectonic Geology of Papua 137 dan a. polhemus 2.2. Soils of Papua 165 geoffrey s. hope and alfred e. hartemink 2.3. Climate of Papua 177 michael l. prentice and geoffrey s. hope 2.4. Papuan Terrestrial Biogeography, with Special Reference to Birds 196 bruce m. beehler 2.5. Freshwater Biogeography of Papua 207 dan a. polhemus and gerald r. allen 2.6. Paleontology of Papua 246 geoffrey s. hope and ken p. aplin 2.7. Paleoecology and Paleoenvironments of Papua 255 geoffrey s. hope SECTION THREE. THE FLORA 3.1. Introduction to the Flora of Papua 269 wayne n. takeuchi 3.2. Lichen Biodiversity in New Guinea 303 harrie sipman and andre´ aptroot vii

................. 16157$

CNTS

04-13-07 13:52:28

PS

PAGE vii

viii / contents

3.3. Bryophytes of Papua New Guinea: Their Diversity, Ecology, and Conservation 320 benito c. tan, sinikka piippo, and daniel h. norris 3.4. Ferns and Lycophytes of Papua 335 barbara s. parris 3.5. Gymnosperms of Papua 344 jianhua li 3.6. Angiosperms 349 Annonaceae of Papua 349 paul j. a. keßler and johan b. mols Apocynaceae of Papua 355 david j. middleton Arecaceae of Papua 359 william j. baker and john dransfield The Asclepiad Flora of New Guinea 371 paul i. forster Costaceae of Papua 379 mark f. newman Elaeocarpaceae of Papua 381 mark j. e. coode Ericaceae of Papua 389 lyn a. craven Euphorbiaceae of Papua 394 peter c. van welzen Melastomataceae of Papua 399 susanne s. renner Moraceae of Papua 404 george d. weiblen Myristicaceae of Papua 408 willem j. j. o. de wilde Myrsinaceae of Papua 416 john j. pipoly iii Myrtaceae of Papua 429 lyn a. craven Orchidaceae of Papua 435 andre´ schuiteman and ed f. de vogel Sapindaceae of Papua 457 peter c. van welzen Sapotaceae of Papua 462 willem vink Zingiberaceae of Papua 473 mark f. newman

................. 16157$

CNTS

02-08-07 10:20:42

PS

PAGE viii

co ntents / ix SECTION FOUR. THE FAUNA 4.1. Introduction to the Fauna of Papua 479 allen allison 4.2. Marine Invertebrates of Papua 495 fred e. wells 4.3. Insects of Papua 515 scott e. miller 4.4. Cicada Endemism in Papua 532 hans duffels and arnold j. de boer 4.5. Marine Wood-Boring Invertebrates of New Guinea and Its Surrounding Waters 539 simon m. cragg 4.6. Herpetofauna of Papua 564 allen allison 4.7. The Monitor Lizards of Papua 617 kai m. philipp and devi p. philipp 4.8. Fishes of Papua 637 gerald r. allen 4.9. Birds of Papua 654 andrew mack and jack dumbacher 4.10. A Taxonomic and Geographic Overview of the Mammals of Papua 689 kristofer m. helgen

................. 16157$

CNTS

04-13-07 13:52:28

PS

PAGE ix

................. 16157$

CNTS

02-08-07 10:20:43

PS

PAGE x

Abbreviations Used in This Volume A ABL ABRI ACIAR AM AMNH ANPWS ANU asl B

BAPLAN BAPPEDA BAPPEDALDA

BAPPENAS BIN BISH BKSDA BM BMNH BO bp BP

Arnold Arboretum, Harvard University, Cambridge, Massachusetts, USA (Herbarium) Adviesbureau voor Bryologie en Lichenologie, Soest, The Netherlands (Herbarium) Armed Forces of the Republic of Indonesia (Angkatan Bersenjata Republik Indonesia) Australian Centre for International Agriculture Research, Australia Australian Museum, Sydney, New South Wales, Australia American Museum of Natural History, New York, New York, USA Australian National Parks and Wildlife Service Australian National University, Canberra, Australian Capital Territory, Australia above sea level (elevation) Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universita¨t, Berlin, Germany (Herbarium) Forestry Planning Agency, Indonesia (Badan Planologi Kehutanan) Provincial Development Planning Bureau, Indonesia (Badan Perencanaan Pembangunan Daerah) Provincial Environmental Impact Management Agency, Indonesia (Badan Pengelolaan Pengendalian Dampak Lingkungan Daerah) National Development Planning Agency, Indonesia (Badan Perencanaan Pembangunan Nasional) National Intelligence Board, Indonesia (Badan Intelijen Negara) Bernice P. Bishop Museum, Honolulu, Hawai‘i, USA Nature Conservation Bureau, Indonesia (Balai Konservasi Sumber Daya Alam) Natural History Museum, London, UK (Herbarium) British Museum of Natural History (Herbarium). Now BM: Natural History Museum, London, UK (Herbarium) Herbarium Bogoriense, Bogor, Indonesia (Herbarium) [years] before present British Petroleum, now referred to as BP xi

................. 16157$

ABBR

02-08-07 10:20:46

PS

PAGE xi

xii / abbreviations u sed i n this v olume

BP3D

BPBM BPID BPPD BPS BRI BRIT brl BTN BYU ca CANB CAS CI CIFOR CITES CNC CPS CPSW CSIRO CTS DAK DAS DASF DAU dbh DEC DEFRA DFMR DKI DKP DPI DPRD

Agency for Planning and Coordination of Regional Development (Badan Perencanaan dan Pengendalian Pembangunan Daerah) Bishop Museum, Honolulu, Hawai‘i, USA Regional Promotion and Investment Board (Badan Promosi Investasi Daerah) Regional Development and Productivity Board (Badan Pengembangan Produktivitas Daerah) Central Bureau of Statistics, Indonesia (Badan Pusat Statistik) Queensland Herbarium, Brisbane, Australia (Herbarium) Botanical Research Institute of Texas, Fort Worth, Texas, USA (Herbarium) barrels National Park Bureau, Indonesia (Balai Taman Nasional) Brigham Young University, Provo, Utah, USA approximately (circa) Centre for Plant Biodiversity Research, Canberra, Australia (Herbarium) California Academy of Sciences, San Francisco, California, USA Conservation International, Washington, D.C., USA Center for International Forestry Research Convention on International Trade in Endangered Species of Wild Flora and Fauna Canadian National Collection, Ottawa, Ontario, Canada conservation priority setting Conservation Priority-Setting Workshop (of CI) Commonwealth Scientific and Industrial Research Organisation, Australia case tracking system Special Allocation Fund (Dana Alokasi Khusus) Department of Agriculture and Stock, PNG Department of Agriculture, Stock and Fisheries, PNG General Allocation Fund (Dana Alokasi Umum) diameter at breast height Department of Environment and Conservation, PNG Department for Environment, Food and Rural Affairs, UK Department of Fisheries and Marine Resources, PNG Jakarta, Special Capital Province (Daerah Khusus Ibukota) Department of Marine Affairs and Fisheries (Departemen Kelautan dan Perikanan) Department of Primary Industries, PNG Provincial Council (Dewan Perwakilan Rakyat Daerah)

................. 16157$

ABBR

02-08-07 10:20:46

PS

PAGE xii

a b b rev i a tio ns used i n thi s v olu me / xiii DSIR E EBA EEC EIA ELA ENSO EPI ESR FAO FCI FH

FI FKPTP FM FOE FR FUNDWI FWI G GDP GFW GH GIS GNP GNrP GOI Golkar GTZ H ha HDI HE HPH HPT HTI IBA

Department of Scientific and Industrial Research, Lincoln, Canterbury, New Zealand Royal Botanic Garden, Edinburgh, Scotland (Herbarium) Endemic Bird Areas (BirdLife International) European Economic Community Environmental Investigation Agency, UK equilibrium-line altitude El Nin˜o–Southern Oscillation Extended Program on Immunization (WHO) electron spin resonance (archeological dating technique) Food and Agriculture Organization (UNDP) Forest Civil Investigators, Indonesia Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, Cambridge, Massachusetts, USA (Herbarium) Museo Botanico, University of Florence, Florence, Italy (Herbarium) Forum for Conservation and Development in Papua (Forum untuk Konservasi dan Pembangunan di Tanah Papua) Flora Malesiana Friends of the Earth, San Francisco, California, USA forest ranger Fund of the United Nations for the Development of West Irian Forest Watch Indonesia Geneva, Switzerland (Herbarium) gross domestic product Global Forest Watch Gray Herbarium of Harvard University, Cambridge, Massachusetts, USA geographic information systems gross national product gross national-regional product Government of Indonesia Party of the Functional Groups (Partai Golongan Karya) German Technical Cooperation (Gesellschaft fu¨r Technische Zusammenarbeit) Helsinki, Finland (Herbarium) hectares human development index Halmahera Eddy logging concession license or licensed logging concession (Hak Pengusahaan Hutan) limited production forest (Hutan Produksi Terbatas) industrial timber estate (Hutan Tanaman Indistri) Important Bird Areas (BirdLife International)

................. 16157$

ABBR

02-08-07 10:20:46

PS

PAGE xiii

xiv / a b b rev i a tion s used i n thi s vo lu me

IBSAP ICG IHHBK IHPHH IHPHHMHA

IIED INPRES IPCC IPK IPKMA ISSG ITCZ ITTO IUCN IUPHHK JATAM JE K KK KLH Kodam Kopermas KSDA KUHAP L LAE LG LIPI LK LMA

Indonesian Biodiversity Strategy and Action Plan International Crisis Group license to collect non-timber forest products (Ijin Pemungutan Hasil Hutan Bukan Kayu) license to log forests (Ijin Hak Pemungutan Hasil Hutan) license to log forests based on traditional community rights (Ijin Hak Pemungutan Hasil Hutan Masyarakat Hukum Adat) International Institute for Environment and Development presidential decree (Instruksi Presiden) Intergovernmental Panel on Climate Change timber extraction license (Ijin Pemungutan Kayu) license to log traditional community forests (Ijin Pemungutan Kayu Masyarakat Adat) IUCN/SSC Invasive Species Specialist Group intertropical convergence zone International Tropical Timber Organization International Union for the Conservation of Nature and Natural Resources industrial timber license (Izin Usaha Pemanfaatan Hasil Hutan Kayu) Mining Advocacy Network (Jaringan Advokasi Tambang) Jena, Germany (Herbarium) Kew Royal Botanic Gardens, Kew, Richmond, Surrey, UK (Herbarium) work contract (Kontrak Kerja) Ministry for the Environment, Indonesia (Kementerian Lingkungan Hidup) Regional Police Command (Komando Daerah Militer) Community Cooperative (Koperasi Peranserta Masyarakat) Natural Resources Conservation Department (Konservasi Sumber Daya Alam) Indonesian Code of Criminal Litigation (Kitab UndangUndang Hukum Acara Pidana) Nationaal Herbarium Nederland, Leiden University branch, Leiden, The Netherlands (Herbarium) Papua New Guinea Forest Research Institute, Lae, Papua New Guinea (Herbarium) Universite´ de Lie`ge, Lie`ge, Belgium (Herbarium) Indonesian Institute of Sciences (Lembaga Ilmu Pengetahuan Indonesia) event report (Laporan Kejadian) Traditional Community Associations (Lembaga Masyarakat Adat)

................. 16157$

ABBR

02-08-07 10:20:47

PS

PAGE xiv

a b br ev ia ti ons used in this v ol um e / xv LMS LNG MAN MCZ MEL MI MMAF MNI MoF MoIT MoU MSY MT MULO MVP MVZ mya MZB NGCC NGCUC NGO NHDR NHM NICH NNGPM NSW NTFP NY OPM OSL PA PCF PDI-P Pepera

London Missionary Society liquified natural gas Herbarium Manokwariense, Cenderawasih University, Manokwari, Papua, Indonesia (Herbarium) Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA Melbourne, Australia (Herbarium) Millennium Institute Ministry of Marine Affairs and Fisheries, Indonesia minimum number of individuals Ministry of Forestry, Indonesia Ministry of Industry and Trade, Indonesia memorandum of understanding maximum sustainable yield metric tons advanced primary school minimum viable population Museum of Vertebrate Zoology, University of California, Berkeley, California, USA million years ago Museum Zoologense Bogeriense, Bogor, Indonesia New Guinea Coastal Current New Guinea Coastal Undercurrent nongovernmental organization National Human Development Report (UNDP) Natural History Museum, London, UK; formerly British Museum of Natural History (BMNH) Japan Hattori Botanical Laboratory, Nichinan, Japan (Herbarium) Netherlands New Guinea Oil Company (Nederlandsche Nieuw-Guinea Petroleum Maatschappij) Sydney, New South Wales, Australia (Herbarium) non-timber forest product New York Botanical Garden, The Bronx, New York, USA (Herbarium) Free Papua Organization (Organisasi Papua Merdeka) optically stimulated luminescence (archeological dating technique) protected area Papua Conservation Fund Indonesian Democratic Party of Struggle (Partai Demokrasi Indonesia Perjuangan) Act of Free Choice (Penentuan Pendapat Rakyat)

................. 16157$

ABBR

02-08-07 10:20:47

PS

PAGE xv

xvi / a b b rev i a tion s used i n thi s vo lu me

PERPU PHKA PHPA

PKK PNG PNGRIS POLRI PP ppm PPNS PSP PSW Psu QIMR RACE RAP REPELITA RePPProT RI RMAP RNH ROI RSA SEC SK SKSHH SPP SPPP SSC SST TAC

Government Regulation in Lieu of Law (Peraturan Pemerintah Pengganti Undang-Undang) Directorate of Conservation and Protected Areas, Indonesia (Perlindungan Hutan dan Konservasi Alam); formerly PHPA Directorate of Forest Protection and Nature Conservation, Indonesia (Perlindungan Hutan dan Pelestarian Alam); now PHKA Applied Family Welfare Program (Pembinaan Kesejahteraan Keluarga) Papua New Guinea Papua New Guinea Resource Information System Indonesian State Police (Polisi Negara Republic Indonesia) government regulation (Peraturan Pemerintah) parts per million Civil Service Investigation Office (Penyidik Pegawai Negeri Sipil) Priority-setting Program (of CI) Priority-setting Workshop (of CI) practical salinity units Queensland Institute of Medical Research, Queensland, Australia Rapid Assessment for Conservation and Economy (of CI) Rapid Assessment Program (of CI) Five-year Development Plan (Rencana Pembangunan Lima Tahun) Regional Physical Planning Program for Transmigration, ROI Republic of Indonesia (also ROI) Resource Management in Asia-Pacific Nationaal Natuurhistorisch Museum, Leiden, The Netherlands; formerly Rijksmuseum van Natuurlijke Historie Republic of Indonesia (also RI) Rancho Santa Ana Botanic Garden Herbarium, Claremont, California, USA (Herbarium) south equatorial current decree (Surat Keputusan) certificate that logs were legally obtained (Surat Keterangan Sahnya Hasil Hutan) investigation warrant (Surat Perintah Penyidikan), also known as SP2 or SPRINT letter of termination of investigation (Surat Perintah Penghentian Penyidikan), also known as SP3 IUCN Species Survival Commission sea surface temperature total allowable catch

................. 16157$

ABBR

02-08-07 10:20:48

PS

PAGE xvi

abbreviations used i n this v olume / xvii TL TMDU TNC TNI TNS TNWP Trikora UC UNCEN UNDP UNEP UNESCO UNFCCC UNIPA UniTech UNTEA UPNG UPT USNM UU WALHI WCMC WCS WEI WHO WMA WPWP WRI WRSL WSPCW WWF YALI YPMD

thermo-luminescence (archeological dating technique) Tokyo Medical and Dental University, Bunkyo-ku, Tokyo, Japan The Nature Conservancy Indonesian National Army (Tentara Nasional Indonesia) National Science Museum, Tsukuba, Japan (Herbarium) Tri-National Wetlands Program People’s Threefold Command (Tri Komando Rakyat) University of California Herbarium, Berkeley, California, USA (Herbarium) Cenderawasih University (Universitas Cenderawasih), Jayapura, Papua, Indonesia United Nations Development Program United Nations Environmental Program United Nations Educational, Scientific and Cultural Organization United Nations Framework Convention on Climate Change State University of Papua (Universitas Negeri Papua), Manokwari, Papua, Indonesia University of Technology, Lae, Morobe Province, PNG United Nations Temporary Executive Authority University of Papua New Guinea, Port Moresby, PNG (Herbarium) Technical Implementation Units, Indonesia (Unit Pelaksana Teknis) United States National Museum, Smithsonian Institution, Washington, D.C., USA law (Undang-Undang) Indonesian Forum for the Environment (Wahana Lingkungan Hidup Indonesia) World Conservation Monitoring Centre (UNEP) Wildlife Conservation Society Wau Ecology Institute, Wau, Morobe Province, PNG; formerly Bishop Museum Field Station World Health Organization Wildlife Management Areas, PNG Western Pacific Warm Pool World Resources Institute Wroclaw University, Wroclaw, Poland (Herbarium) Western South Pacific Central Water World Wide Fund for Nature; World Wildlife Fund in USA The Papua Environment Foundation (Yayasan Lingkungan Hidup Papua) Irian Jaya Rural Community Development Foundation (Yayasan Pengembangan Masyarakat Desa)

................. 16157$

ABBR

02-08-07 10:20:48

PS

PAGE xvii

................. 16157$

ABBR

02-08-07 10:20:48

PS

PAGE xviii

Contributors Gerald R. Allen. Department of Aquatic Zoology, Western Australian Museum, 49 Kew Street, Welshpool, WA 6106, Australia Allen Allison. Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, USA. E-mail: [email protected] Daniel M. Alongi. Australian Institute of Marine Science, PMB 3, Townsville MC, Queensland 4810, Australia Dessy Anggraeni. Conservation International Indonesia, Jl. Pejaten Barat 16 A Kemang, Jakarta 12550, Indonesia Ken P. Aplin. Division of Wildlife Research, CSIRO, Canberra 0200, Australia Andre´ Aptroot. Centraalbureau voor Schimmelcultures, PO Box 85167, NL-3508 AD Utrecht, The Netherlands William J. Baker. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom Renee Bartolo. GecOz Pty Ltd, Geospatial Consultants Australia, PO Box 42636, Casuarina, NT 0811, Australia Bruce M. Beehler. Melanesia Program, Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA Manuel Boissie`re. CIRAD, Campus de Baillarguet, 34398 Montpellier Cedex 5, France, and CIFOR, PO Box 6596, JKPWB Jakarta 10065, Indonesia Michele Bowe. World Wide Fund for Nature (WWF), PMB Madang, Madang Province, Papua New Guinea John Burke Burnett. Indo-Pacific Conservation Alliance and Pacific Science Association, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, USA James B. Cannon. Center for Conservation and Government, Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA Rob Coles. CRC Reef Research Centre/Department Primary Industries and Fisheries, Northern Fisheries Centre, PO Box 5396, Cairns, Queensland 4870, Australia Mark J. E. Coode. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom Simon M. Cragg. Institute of Marine Sciences, School of Biological Sciences, University of Portsmouth, Ferry Road, Portsmouth PO4 9LY, United Kingdom Lyn A. Craven. Australian National Herbarium, CPBR, CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia. E-mail: [email protected] Arnold J. de Boer. Zoological Museum, Department of Entomology, University of Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands Yance de Fretes. Conservation International Indonesia, Jl. Pejaten Barat 16 A Kemang, Jakarta 12550, Indonesia xix

................. 16157$

CTRB

02-08-07 10:20:48

PS

PAGE xix

xx / contributors

Louis Deharveng. UMR5202 du CNRS, Origine, Structure et Evolution de la Biodiversite´, Muse´um National d’Histoire Naturelle, CP 50, 45 rue Buffon, 75005 Paris, France Ed F. de Vogel. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands Willem J. J. O. de Wilde. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands John Dransfield. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom Hans Duffels. Zoological Museum, Department of Entomology, University of Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands Jack Dumbacher. Department of Birds and Mammals, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, USA Mark V. Erdmann. Center for Applied Biodiversity Science, Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA Paul Erftemeijer. WL 兩 Delft Hydraulics, PO Box 177, 2600 MH Delft, The Netherlands Douglas Fenner. Department of Marine and Wildlife, American Samoa, PO Box 3730, Pago Pago, American Samoa 96799 Paul I. Forster. Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt. Coot-tha Road, Toowong, Queensland 4066, Australia Scott Frazier. Conservation International, Papua Program. Currently at 20 NW 400 Road, Warrensburg, MO 64093, USA. E-mail: [email protected] David G. Frodin. Chelsea Physic Garden, London; Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom Alfred E. Hartemink. ISRIC—World Soil Information, PO Box 353, 6700 AJ Wageningen, The Netherlands Kristofer M. Helgen. School of Earth and Environmental Sciences, University of Adelaide, Adelaide, SA 5005; and South Australian Museum, North Terrace, Adelaide, SA 5000, Australia. Current e-mail: [email protected] Geoffrey S. Hope. Archaeology and Natural History, Australian National University, Canberra 0200, Australia Robert J. Johns. Botanical Research Institute, Fort Worth, TX 76102, USA Neville J. Kemp. Indo-Pacific Conservation Alliance, c/o Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, USA Paul J. A. Keßler. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands. E-mail: [email protected] Keliopas Krey. Department of Biology, University of Papua, Manokwari, Papua, Indonesia Philippe Leclerc. 43 boulevard de la Vanne, 94230 Cachan, France Jianhua Li. Arnold Arboretum of Harvard University, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, USA

................. 16157$

CTRB

02-08-07 10:20:49

PS

PAGE xx

contributors / xxi Andrew Mack. Papua New Guinea Country Program, Wildlife Conservation Society, PO Box 277, Goroka, EHP, Papua New Guinea J. R. Mansoben. Lembaga Penelitian, Cenderawasih University, Jayapura, Papua, Indonesia Andrew J. Marshall. Department of Anthropology and Graduate Group in Ecology, University of California, One Shields Avenue, Davis, CA 95616, USA Len McKenzie. CRC Reef Research Centre/Department Primary Industries and Fisheries, Northern Fisheries Centre, PO Box 5396, Cairns, Queensland 4870, Australia David J. Middleton. Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland Scott E. Miller. National Zoological Park and National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20008-2598, USA Johan B. Mols. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands Mark F. Newman. Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland Daniel H. Norris. Department of Integrative Biology and Jepson Herbaria, University of California, Berkeley, CA 97420-2465, USA Barbara S. Parris. Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands 0470, New Zealand Juliette Pasveer. Archaeology and Natural History, Australian National University, Canberra 0200, Australia Devi P. Philipp. Zoologische Staatssammlung, Sektion Herpetology, Mu¨nchhausenstr. 21, D-81247 Munich, Germany Kai M. Philipp. Zoologische Staatssammlung, Sektion Herpetology, Mu¨nchhausenstr. 21, D-81247 Munich, Germany Sinikka Piippo. Botanical Museum, Finnish Museum of Natural History, PO Box 7, FIN-00014 University of Helsinki, Finland John J. Pipoly III. University of Florida—Institute of Food and Agricultural Sciences/Broward County Extension, 3245 College Avenue, Davie, FL 333147719, USA Dan A. Polhemus. Department of Entomology, MRC 105, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, USA Michael Prentice. Indiana Geological Survey, Department of Geological Sciences, Indiana University, 611 N. Walnut Grove, Bloomington, IN 47405, USA Pratito Puradyatmika. Environmental Department, P.T. Freeport Indonesia, Mimika, Papua, Indonesia Yohanes Purwanto. LIPI, Laboratorium Etnobotani, Puslitbang Biologi, Jl. Ir. H. Juanda 22 Bogor, Indonesia Susanne S. Renner. Institute of Systematic Botany, Ludwig Maximilian University, Menzinger Str. 67, D-80638 Munich, Germany Stephen Richards. Vertebrate Department, South Australian Museum, North Terrace, Adelaide, S.A. 5000, Australia

................. 16157$

CTRB

02-08-07 10:20:49

PS

PAGE xxi

xxii / c o n t r i b u t o r s

Andre´ Schuiteman. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands Garry A. Shea. P.T. Hatfindo Prima, Bogor, Indonesia. Harrie Sipman. Botanischer Garten und Botanisches Museum, Freien Universita¨t, Ko¨nigin-Luise-Str. 6–8, D-14191 Berlin, Germany Neil Stronach. Fota Wildlife Park, Carrigtohill, Co. Cork, Ireland Suer Suryadi. Pusat Informasi Lingkungan Indonesia-NGO MOVEMENT Jl. Tumenggung Wiradireja No. 216, Cimahpar, Bogor 16155, West Java, Indonesia Wayne N. Takeuchi. Herbaria and Arnold Arboretum of Harvard University, c/o PNG National Forest Authority, Lae, Papua New Guinea Benito C. Tan. Department of Biological Sciences, National University of Singapore, 119260, Singapore Jaap Timmer. Radboud University, Nijmegen, The Netherlands. E-mail: [email protected] Burhan Tjaturadi. Conservation International—Papua Program, Jl. Bhayangkara I No. 5, Jayapura, Papua Province, Indonesia Peter C. van Welzen. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA Leiden, The Netherlands Willem Vink. Nationaal Herbarium Nederland, Universiteit Leiden, PO Box 9514, 2300 RA Leiden, The Netherlands George D. Weiblen. Department of Plant Biology and Bell Museum of Natural History, University of Minnesota, 250 Biological Science, 1445 Gortner Avenue, Saint Paul, MN 55108, USA Fred E. Wells. Western Australian Museum, Perth 6000, Western Australia. Currently at Department of Fisheries, Western Australia, Level 3, The Atrium, 168 St. Georges Terrace, Perth 6000, Western Australia Tony Whitten. Senior Biodiversity Specialist, Environment and Social Development Sector, East Asia and Pacific Region, The World Bank, 1818 H Street NW, Washington, D.C. 20433, USA, and Conservation Biology Group, Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, United Kingdom Agustinus Wijayanto. Conservation International Indonesia, Pejaten Barat No. 16A, Kemang, Jakarta Selatan, Indonesia

................. 16157$

CTRB

02-08-07 10:20:49

PS

PAGE xxii

Acknowledgments h e dev e l op m e n t of The Ecology of Papua has a long and complex history. The genesis of the Ecology of Indonesia series dates from 1981, when the Sumatra volume was written under the Government of Indonesia/United Nations Development Program Project INS/78/056 entitled Education and Training in Environment and Resources, executed by the World Bank with a subcontract to Dalhousie University. This was followed by a project entitled Environmental Management and Development in Indonesia (EMDI), financed by the Canadian International Development Agency (CIDA). This effort led to the concept of a national series, including a volume on Papua. Unfortunately, the original plan to prepare the Papua (then Irian Jaya) volume in the early 1990s faced a variety of problems, and it was never completed. Over the long gestation period of this work, during which time the other seven volumes were completed (and two volumes already revised), there have been a series of editors for the Papua volume. Because of a number of insurmountable hurdles, the book project never really moved ahead until 2004, at which point BP (British Petroleum), through its Tangguh Project initiative, agreed to provide a grant to support the book in its latest incarnation as an edited volume. We here recognize BP’s Tangguh partners and their support of the project through BP: KG Berau Petroleum Ltd., Nippon Oil Exploration (Berau) Ltd., MI Berau BV, BP Berau Ltd., BP Wiriagar Ltd., BP Muturi Holding BV, KG Wiriagar Petroleum Ltd., CNOOC Wiriagar Overseas Ltd., Indonesia Natural Gas Resources Muturi, Inc., and CNOOC Muturi Ltd. The BP grant, as well as a grant from CI to Harvard University, permitted the contracting of AJM to serve as managing editor for the project (2004–2006). We thank Karla Boreri Dutton, Lidia Ahmad, Jalal, Erwin Maryoto, and the entire BP Tangguh Project Environmental Team for being instrumental in fostering this partnership with BP. The Gordon and Betty Moore Foundation, through its generous funding of CI’s Melanesia Center for Biodiversity Conservation, provided the impetus for completing this demanding project. We are deeply grateful for its support. With this support from the Moore Foundation and BP, the work began in earnest, resulting in written contributions from eighty-six authors, and with additional technical assistance from Conservation International–Indonesia, especially Dr. Yance de Fretes, a long-term student of the biodiversity of Papua. We thank Tony Whitten for his genius in pulling together the concept for this important book series. We thank also Kathy MacKinnon, who also was central to the series. We thank Ron Petocz for all the background bibliographic research he conducted on Papua, which today remains a remarkable achievement. The advisory team for the book, which included Whitten, as well as Gerald Allen, Allen

T

xxiii

................. 16157$

$ACK

02-08-07 10:20:54

PS

PAGE xxiii

xxiv / acknowledgments

Allison, Chris Ballard, BMB, Jim Cannon, Yance de Fretes, Geoff Hope, Robert Johns, J. R. Mansoben, Scott Miller, Dan Polhemus, and Wayne Takeuchi, provided invaluable thinking on the composition and vision for the work. As a group we decided to make it an expert-chapters book rather than a synthesis. We hope readers approve of this choice. We are greatly indebted to the nearly ninety text contributors, without whose voluntary writing efforts we would have no book. When the final deadline came, every author did submit the required piece, and many of them are superlative contributions to science. Bravo to them! We also thank those who contributed photographs. We especially thank Gerald Allen, J. Burke Burnett, Michael P. Moore, and Stephen J. Richards, who provided a large share of the color photographs. We offer thanks to the staff of Conservation International in Jakarta and Jayapura who helped us in a variety of ways with this immense project. Thanks to Dr. Jatna Supriatna, CI–Indonesia’s Regional Director, for his leadership on this initiative. We also especially note the assistance of Budi Iraningrum with respect to her work on literature research for the project, Hendi Sumantri with maps for the book, Tommy Wakum for gathering information from government agencies in Papua, as well as Scott Frazier. CI’s Center for Environmental Leadership in Business made possible the relationship with BP, and for this we thank Glenn Prickett, Assheton Carter, and Marielle Carter. We thank the Harvard University Herbaria, especially Bob Cook and Frances Maguire, for helping to arrange a postdoctoral appointment for AJM during the period that this book was pulled together. We also thank Rose Balan, Donna Barrett, Anne Marie Countie, Deidre Fogg, Ingrid McDonough, Karen Pinto, Chris Preheim, and Lisa Toste for administrative and technical support. AJM would also like to express thanks and appreciation to his friends and colleagues at the Herbaria, especially Peter Ashton, Jen Baltzer, Stuart Davies, Jessica Dolan, Wendy Duan, Amy Dunham, Ken Feeley, Kanchi Ghandi, Henry Kesner, Walter Kittridge, Genevieve Lewis-Gentry, Dave Lohman, Laura Lukas, Melinda Peters, Sabrina Russo, and Emily Wood. Our publisher has been very supportive of this remaining volume of the series, and for this we thank them, especially Ed Walters and Christine LeBlond, who patiently guided us along the way. We salute the CEO of Periplus, Eric Oey, for his commitment to the series. We are indebted to the Indonesian State Minister of the Environment for providing the Preface for the book, and the Papuan Peoples’ Assembly for their supporting letter. We thank Professor Edward O. Wilson for providing the Foreword. In addition, we recognize J. Burke Burnett for his dedication to the completion of this project. Michelle Brown also deserves mention as an important supporter of the initiative. The project could not have happened without the support and dedication of the Government of Indonesia. In particular, we thank Dr. Dedy Darnedi, director of the Biology Research and Development Center, Indonesian Institute of Science. We also thank the Universitas Cenderawasih in Abepura and the State University

................. 16157$

$ACK

02-08-07 10:20:54

PS

PAGE xxiv

acknowledgments / xxv of Papua in Manokwari for their support. Thanks also to the Papua Conservation Fund for supporting this project. Anne McGuire was this project’s tireless editor, reader, and indexer. For this major contribution, we offer our utmost gratitude to her. In a like manner, Ann Twombly provided the excellent book design, and patiently saw this unwieldy project through to the typesetter and, finally, the printer. Thanks for her wonderful work! Finally, we thank our families for their support, and we hope the students of Papua over the next decade will find this book helpful in their efforts to better know and better conserve all that makes Papua unique—the forests and waters and wildlife and traditional societies. 19 October 2006 Andrew J. Marshall Davis, California

Bruce M. Beehler Washington, D.C.

................. 16157$

$ACK

02-08-07 10:20:54

PS

PAGE xxv

................. 16157$

$ACK

02-08-07 10:20:55

PS

PAGE xxvi

Foreword edward o. wilson a p ua h a s la s t e d into the twenty-first century as largely a blank space on the map, and we will do well to treasure it for that. Here for the last time in history, as human modernity closes irreversibly over the planet, we may take comfort that there still exists a land ‘‘beyond the frontier’’ such as Papua. We can still feel the aura that drew explorers of past centuries and provided an exit from their otherwise controlled and predictable lives. We can share what the great adventurer Richard Burton expressed in 1856: ‘‘Of the gladdest moments in human life, methinks, is a departure upon a distant journey into unknown lands.’’ There, he rejoiced, ‘‘Afresh dawns the morn of life.’’ And exploring Papua is no trek through hourless days across some vast desert. Rather, it offers entry, as the present volume makes clear, into an intact world of ancient cultures and wondrous life forms. I have never visited Papua, but I enjoyed all the emotions just expressed while traveling in 1955 through a large part of Papua New Guinea, to the east. I was a young entomologist then, myrmecologist to be exact, and the first specialist on ants to visit the great island. With no predecessors except a few casual collectors of these little insects, I had only a vague idea of what to expect. Everywhere I searched I found new species. I studied previously unrecorded social behaviors, and I frantically scribbled notes on all I saw (there were no pocket-sized tape recorders in those days). Humble though ants may be, and as modest my own efforts, I considered myself a true explorer in unknown terrain. With no previous myrmecologist’s footprints to give me pause, I felt somehow I belonged to this land and had some responsibility for it. I like to believe that others privileged to pioneer in their respective specialties have felt the same way. New Guinea, including Papua, is a challenge and a paradise for anthropologists and biogeographers. Its complex mountainous terrain has divided its human populations, during 40,000 or more years of occupation, into the most diverse array of cultures and languages of any comparable area in the world. Long before people arrived, the island’s equatorial location and geology combined to make it one of the several most biologically rich regions of Earth, both on the land and in the coral reefs of its marine coasts. For millions of years new species have flowed into it from nearby tropical Asia to the west and from Australia to the south. Many of the immigrants succeeded in penetrating the mountainous interior; and there, like Homo sapiens among the more recent immigrants, they tended to spread out and diversify. Some of the productions of the indigenous fauna and flora were trapped by their adaptation to strictly local conditions. Others, ecologically more flexible, expanded their ranges to penetrate northern Australia, as well as islands and archipelagoes in the remainder of Melanesia.

P

xxvii

................. 16157$

FRWD

02-08-07 10:20:57

PS

PAGE xxvii

xxviii / foreword

The island’s large size, its constant climate favorable to vegetative growth, rugged topography, and nearness to the continental source areas of Asia and Australia have made New Guinea a hypergenerator of human and biological diversity. New Guinea, including the remote marches of Papua, will not remain secluded for long, however. Soon its indigenous people, together with a growing immigrant population, will take over as the explorers and developers. Those of us on the outside able to conduct the early studies of the island nonetheless have a responsibility to make the transition in future generations as secure and beneficent as possible. We will serve them and the whole world to great benefit if we include in this program of assistance the effort to conserve as much as possible of the great island’s extraordinary cultural and natural heritage.

................. 16157$

FRWD

02-08-07 10:20:57

PS

PAGE xxviii

Preface a p ua i s t he la r ge s t i sl a n d in Indonesia (area of 41.48 million ha), with the smallest population compared to other islands of Indonesia. More than 2.6 million people live in Papua, and almost 75% of the population lives in rural areas. Since the early 1990s, Papua has experienced the highest population growth rate of all Indonesian provinces, which stands at over 3% annually. This is partly a result of high birth rate and influx migration from other regions in Indonesia. Papua is endowed with an amazing range of diverse and unusual ecosystems, including glaciers, alpine meadows, cloud forests, lowland forests, savannahs, mangrove forests, coral reefs, and seagrass beds. Well known for its vital tropical rainforest, with the tallest tropical trees and vast biodiversity, Papua plays a pivotal role in Indonesia’s biodiversity, contributing to the country’s status as one of the biologically richest countries in the world and hosting around 50% of all of Indonesia’s biodiversity. Papua is home to 15,000–20,000 plants (55% endemic), 602 birds (52% endemic), 125 mammal (58% endemic), and 223 reptiles (35% endemic). Many of these species are endemic to the island, including birds of paradise, tree kangaroos, rainbow fishes, birdwing butterflies, various orchids, and thousands of other plants and animals. In February 2006, a team of scientists exploring the Foja Mountains discovered numerous new species of birds, butterflies, amphibians, and plants, including a species of rhododendron that may have the largest bloom of the genus. However, biodiversity in Papua is facing very serious problems, such as biodiversity loss and ecological degradation. Ecological threats include logging—induced deforestation, forest conversion into agricultural plantation (especially oil palm), small holder agricultural conversion, the introduction and potential spread of non-native alien species such as the Crab-eating Macaque, which preys on and competes with indigenous species, as well as water pollution from oil and mining operations. I would like to convey my appreciation to the team, especially to the authors, who have provided support and brought together such detailed and fruitful information. I would also like to thank all institution, researchers, universities, nongovernmental organizations, local communities, and all other parties who got involved and participated in the development of this book. I hope this book will serve as a source of information and will give invaluable contributions to the improvement of awareness and knowledge on biodiversity. Finally, I hope the important essence of biodiversity is taken into account in every decision-making process, paving the way toward sustainable development in Papua.

P

Jakarta, July 2006 Ir. Rachmat Witoclar State Minister for Environment xxix

................. 16157$

PREF

02-08-07 10:21:00

PS

PAGE xxix

................. 16157$

PREF

02-08-07 10:21:00

PS

PAGE xxx

Letter from the Papuan People’s Assembly w o uld l i ke to express my sincere thanks and appreciation to Conservation International (CI) for its hard work and devotion in making The Ecology of Papua available. The knowledge presented in these volumes, the last to be released in the Ecology of Indonesia series, represents a significant contribution to efforts made by various local, national, and international institutions and agencies to conserve the biological diversity of this part of the world for future generations. This book will, we hope, also expand readers’ horizons and further their understanding of Papua’s extraordinary diversity and its wide variety of natural ecosystems. These natural treasures need to be properly preserved and managed to prevent their depletion, especially from unwise and inappropriate use of our unique resources. Furthermore, it is my hope that the information on the ecology and biodiversity of Papua presented in this book will significantly contribute to establishing and strengthening awareness and responsibility among various individuals and groups in Papua and in Indonesia to participate in a worldwide movement to conserve this part of the world’s heritage to guarantee a sustainable global future for Papuans and humans in general. Many people and individuals, as well as institutions in Indonesia and abroad under the lead of CI, have contributed significantly and in various ways toward the creation of this book. On behalf of the Papua People’s Assembly (Majelis Rakyat Papua–MRP), then, I would like to thank Conservation International, which has taken the lead throughout each stage of this process—from the collection and compilation of information to the publication process that finally made this book available. This achievement exemplifies CI’s sincere devotion and strong commitment to conserving Papua’s biodiversity, which it has demonstrated throughout its presence in Papua. I would also like to thank the University of Cenderawasih and Harvard University for their significant contributions in making this book available. My deepest gratitude also goes to all chapter authors, as without their contributions this book would never have been possible. In conclusion, allow me on behalf of the Papua People’s Assembly to offer our support for future efforts made by CI in its mission to enhance conservation efforts locally and nationally as well as globally, and through its various programs to conserve the biodiversity of Papua as part of our planet’s last heritages. This is vital for the well-being, prosperity, and sustainable existence of human beings on this planet—particularly Papuans—the indigenous people of this land of Papua.

I

Dr. Agus Alue Alua, M. Th. Chair, Papua People’s Assembly

xxxi

................. 16157$

LETR

02-08-07 10:21:07

PS

PAGE xxxi

................. 16157$

LETR

02-08-07 10:21:07

PS

PAGE xxxii

section one 

Introduction to Papua

................. 16157$

PRT1

04-13-07 13:50:03

PS

PAGE 1

................. 16157$

PRT1

02-08-07 10:21:49

PS

PAGE 2

1.1. Introduction to Papua bruce m. beehler a p ua , t h e w e s t er n h a lf of the great subcontinental island of New Guinea, encompasses 416,129 km2 and supports the largest tract of old growth tropical forest wilderness remaining in the Asia-Pacific region. Dominated by the huge Central Cordillera that generates abundant rainfall, the rivers of Papua drain northward into a vast interior basin (the Mamberamo/Meervlakte) and south into a triangular alluvial platform that broadens as it reaches eastward to the border with Papua New Guinea. At its westernmost, Papua is dominated by a welter of small mountain ranges (accreted terranes), peninsulas (Vogelkop [Bird’s Head], Wandammen, Fakfak, Kumawa), and island groups (Raja Ampats, Cenderawasih Bay Islands). In many respects, Papua resembles its eastern counterpart, mainland Papua New Guinea, but its mountains are higher (reaching to the snow line), its swamps are larger (e.g., Mamberamo, the Asmat), its population is smaller (ca 2.2 million vs. ca 5 million), and the exploitation of its vast forests less extensive at the time of this writing. As with Papua New Guinea, Papua is home to many traditional cultures (250 by one estimate; Petocz 1989). Many of these are forestdwelling societies, who have provided remarkably prudent stewardship of their forest resources. Thus Papua’s forest wilderness and diverse marine ecosystems are human-managed natural systems that give the impression of being pristine. For environmentalists, conservationists, and research biologists, Papua is a rich mother lode of natural and cultural history to be documented studied, shared, and preserved.

P

Wonders of Papua By any standards Papua is special and shrouded in mystery. For nearly a halfcentury (1962–2000) it was essentially inaccessible to all but a few international field researchers (see Hope et al. 1976) and thus a terra incognita. As each year went by, other blank spots on the globe were filled in by intrepid adventurers and naturalists, making Papua more and more enticing to outside naturalists. Those smitten with Papua could only read early accounts and examine the pre-1962 holdings of museums and research institutions to get an idea of what lay behind the Papuan (then West Irian or Irian Jayan) veil of the unknown. We did know that Papua was home to the tropical Pacific’s only glaciers. We did know that Papua was the home to hundreds, no, thousands of undescribed species of plants and animals, not to mention the lesser life forms. Jared Diamond rediscovered the Golden-fronted Bowerbird in the Foja Mountains in 1980. Tim Flannery described a new mountain-dwelling tree kangaroo in 1994. Gerald Allen collected his first rainbow fish in Papua in 1980 and described his most recent new Papuan species Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

3

................. 16157$

$CH1

02-08-07 10:22:01

PS

PAGE 3

4 / bruce m . b e e h l e r

in 1998. Clearly there is so much for us to learn about this little-studied land. Adventurers were claiming ‘‘first contacts’’ with forest-dwelling peoples as recently as 1990—this added to the several hundred named ethnic groups inhabiting Papua, each with its own language, culture, art, and cosmology.

Geographic and Political Nomenclature Let us begin our overview of Papua with some discussion of geographic and political nomenclature (see map on end sheets to this book). New Guinea is the term we use to describe the whole island, this largest tropical island, some 2,700 kilometers long by 900 kilometers wide. The eastern half of the island is today the mainland section of Papua New Guinea, which achieved independence from Australia in 1975. The western half of the island is today known informally as Papua (‘‘West Papua’’ in some circles). Papua became the official name of western (Indonesian) New Guinea, Indonesia’s easternmost province in 2000. In 2004, Papua Province was ‘‘illegally’’ but formally bisected; the easternmost and central sections retain the name Papua, and the westernmost section is Irian Jaya Barat (a planned Central Irian Jaya has been put on hold because of a court ruling). Western New Guinea has held various names over the last hundred years. During the days of Dutch colonial administration this area was named Dutch New Guinea, part of the Dutch East Indies. Upon Indonesian accession of this last fragment of Dutch colonialism, the region was named West Irian (Irian Barat). Shortly thereafter it was given the name Irian Jaya (‘‘glorious Irian’’), and more recently Papua. This last is confusing mainly because the southeastern portion of mainland Papua New Guinea was once officially named Papua, when overseen by colonial Australia. Finally, political activists call western New Guinea ‘‘West Papua’’—the name that the local assembly had chosen for the planned independent nation that was to arise in 1962 through a United Nations mandate.

Physiography, Geography, and Geology Papua is a complex piece of the planet, partly because of its convoluted tectonic history, discussed in some detail in Chapter 2.1. In brief, the Papuan component of the Australian tectonic plate has been rafting northward, building a prodigious central cordillera as well as sweeping up island arcs in the north and northwest. This plate continues to drift northward and northern coastal ranges are presumably still rising. Mountains define Papuan geography, no doubt. Two east-west ranges dominate—the Central Cordillera (Merauke Range, ‘‘Maoke’’ is a misnomer; this includes a western component, Sudirman Range, and an eastern component, Jayawijaya Range) and the north coastal ranges that extend westward into Cenderawasih Bay as rugged Yapen Island. The Central Cordillera has been created by the compression of the Australian plate with the Pacific plate, with massive uplift over the last several million years. The highest points of the Sudirman and Jayawijaya

................. 16157$

$CH1

02-08-07 10:22:01

PS

PAGE 4

Introduction to Papua / 5

ranges are oceanic sediments. This cordillera rises to more than 3,000 meters for its entire length in Papua, creating a challenge for Indonesian road builders wishing to link up the northern and southern catchments. The cordilleran watershed dips rather gradually on its northern face and abruptly on the south side. Heavy rainfall striking the southern scarp has deeply dissected this southern face, creating scores of sediment-laden and unstable rivers that dump out onto a rocky alluvial plain in the south that is almost 200 km wide in the east and only 40 km wide in the far west (west of Timika). The highest peaks of Papua are scattered about the main cordillera. Highest of all is Mt Jayakusuma or Mt Jaya (4,884 m) once known as Mount Carstensz or Carstensz Toppen, dominating the western terminus of the Merauke Range. Nearby Ngga Pilimsit or Mount Idenburg stands at 4,717 m. In central and eastern segments of the cordillera stand Mount Trikora (formerly Mount Wilhelmina) at 4,730 m and Mount Mandala at 4,640 m. Small, rapidly melting glaciers cap Jaya and Pilimsit. The accreted island arcs in the north can be seen today as isolated coastal ranges: the Cyclops, Foja, and Van Rees Mountains (north of the Tariku and Taritatu [formerly Idenburg] rivers), mountainous Yapen Island, the Wandammen, Arfak (2,940 m), and Tamrau mountains (2,824 m) of the Vogelkop Peninsula, as well as the Raja Ampat Islands west of the Vogelkop. Strange tectonic contacts apparently have also produced the Kumawa and Fakfak mountains south of the Vogelkop on the Bomberai and Onin peninsulas. The Bird’s Neck region, which connects the Vogelkop with the main body of Papua, is karstic, with fjordlands, white sand barrens, and lakes. Papua is scored by a range of major rivers both north and south, east and west. In the north, the Mamberamo system drains the interior Mamberamo Basin and virtually the entire northern watershed of Papua’s central range. The main channel of the north-flowing Mamberamo cuts between the Foja Mountains (on the east) and Van Rees Mountains (on the west) on its way to the sea. This ramrod straight, swiftly-flowing stream is one of the most remarkable on this great island, even though it is only 150 km in length. At the head of the Mamberamo, the river drains the great interior basin swamplands that are infested by meander belts and oxbow lakes. The Taritatu (formerly Idenburg) River drains the eastern half of the basin and the central mountains to the south, its tributaries reaching to the Papua New Guinea border and nearly to Jayapura. Its western branch, the Tariku (or Rouffaer) River, drains the smaller western side of the basin, and quickly divides into the main flow of the Rouffaer (on the north) and the Van Daalen (to the south). The Van Daalen drains the north slope of the Central Cordillera, and thus is a much more substantial flow. Papua’s other great rivers drain the ragged southern scarp of the central range in the eastern half of Papua. Among these, the Digul is the greatest, followed by the Catalina, which in the mountains becomes the famous Baliem that drains the Grand Valley of the Baliem, discovered in the late 1930s by explorer-pilot Richard Archbold. Scores of lesser rivers sweep heavy gravels southward toward the muddy

................. 16157$

$CH1

02-08-07 10:22:02

PS

PAGE 5

6 / bruce m . b e e h l e r

Arafura Sea. These turbid and unstable rivers tumble out of the mountains, with torrential flows in the mountain gorges, and heavily braided channels in the flats that spread out from the bottom of the ranges. As one moves westward, one finds river after river, each shorter than the preceding, until the central mountains pinch off the alluvial plain at the bottom of the Bird’s Neck.

lakes Papua has a few prominent lakes. Lake Sentani, near the Papuan capital Jayapura, was apparently created by tectonic movement related to the uplift of the coastal Cyclops Mountains just to the north. The lower Mamberamo features Lake Rombebai, the largest lake in Papua, as well as smaller Lake Bira. These are swampy backwater lakes. At the western end of the central cordillera we find the Paniai Lakes in an interior highland basin. Lake Yamur, on the Bird’s Neck, is home to a freshwater shark. Finally, highlands lakes (Anggi Gigi and Anggi Gita) are found in the Arfak Mountains of the Vogelkop.

swamps, mangroves, and savannas The vast lakes plain of the Mamberamo Basin is dominated by seasonally inundated swamplands of various types. There are great coastal swamplands along much of the southern coast, from the Casuarina coast in the southeast to the swamplands south of Timika, far to the west. Indonesia’s largest mangrove ecosystem is nestled in the head of Bintuni Bay, which separates the Vogelkop (Bird’s Head) Peninsula from the more southerly Bomberai Peninsula. Elsewhere in Papua, swamps can be found in many alluvial localities where drainage is impeded, around lowland rivers, and in and around Dolok (Yos Sudarso) Island in the far south. In the far southeast, by the Papua New Guinea border, is a swath of savanna that ranges westward to Dolok Island—part of the great Trans-Fly savannas that have the look of Australia rather than New Guinea. This is a highly seasonal low-rainfall zone that toggles from an inundation season to a burning season.

coasts Papua’s abundant coastline is not uniform. In the northeast, one finds hilly country reaching the coast, which features a mix of white sand beaches and rocky shorelines. Long stretches of beach dominate in the north, backed by coastal hills. The eastern shore of Cenderawasih (formerly Geelvink) Bay features swamps and mangroves, whereas the western shore is more rugged and hilly. The north side of the Vogelkop is rugged, whereas the south side is low and swampy. Much of the southern and southeastern coastline are low and silty, with dark sand beaches backed by casuarinas, with swamplands further inland. The most spectacular coastlines are found on the south side of the Bird’s Neck, between Arguni Bay and Etna Bay. Here one finds tropical karstic fjordlands that feature coastal mountains rising to more than 1,000 meters, steep cliffs, deep embayments, and scenery galore.

................. 16157$

$CH1

02-08-07 10:22:02

PS

PAGE 6

Introduction to Papua / 7

islands Papua has more than a thousand fringing islands, from tiny to quite large. The Raja Ampat Islands range off the western coast of the Vogelkop Peninsula, and include Waigeo (3,155 km2), Salawati (1,632 km2), Misool (2,041 km2), Batanta (453 km2), and Kofiau (150 km2), among others. This remarkable archipelago supports the world’s richest coral reefs and considerable endemic forest biodiversity (e.g., Wilson’s Bird of Paradise, Red Bird of Paradise, Waigeo Brush-turkey). The islands of Cenderawasih Bay include two isolated oceanic islands with distinct island faunas (Biak/Supiori, 2,497 km2, and Numfoor, 311 km2), as well as the mountainous land-bridge island of Yapen (2,227 km2). In addition, there are the Padaido Islands southeast of Biak, and Num Island west of Yapen, and a number of small coastal islands in the south and west portions of the Bay. Small islands also dot the north coast and fringe the Fakfak and Triton Bay region. Papua’s largest island is Dolok (11,192 km2), which is a vast mudbank outwash from the silt-laden rivers of the southeast coast. It is often forgotten because of its unprepossessing nature and isolation, and its minimal distance from the mainland.

Ecological Setting New Guinea is the northern quadrant of the Australian tectonic plate; thus this island is geologically one with the Australian continent. And yet in spite of geological linkages, there are considerable environmental differences. In particular, Australia today is dry and temperate, whereas New Guinea is tropical and perhumid. These two fundamental distinctions can explain much of the differences between these sister biotas, north and south. Climatologically, Papua is remarkable mainly for its cloudiness. It is perhaps one of the cloudiest places on earth. Spanning latitudes from the equator to 12 degrees south latitude, Papua’s equatorial climate is seasonally dominated by the Northwest Monsoon and the Southeast Trade Winds. In most parts of Papua, the effects of the Northwest Monsoon dominate in the period from November to March, bringing rain and unsettled weather. The Southeast Trade Winds tend to bring cool and dry weather, and predominate from April until September. That said, Papua has many microclimates. Rainfall regimes range from low in the southeast (less than 2,000 mm/year) to extremely high on the southern scarp of the Central Cordillera (more than 5,000 mm/year). The highest rainfall on record for Papua is from Tembagapura town, which receives 7,500 mm/year on average. In the wetter areas, the typical seasons are reversed, and the most rain falls in the April–October period. In fact, the wettest sites receive rain from both the monsoon and the trades, and they tend to be found in the mountains along the southern front of the Central Cordillera. Moreover, annual accumulation in the very wettest areas tends to show great variability. This variability can exceed the mean annual accumulations recorded for typical medium-rainfall sites. Seasonally, temperature varies little. Elevation is the key to temperature in equatorial zones. This ‘‘lapse rate’’ is equivalent to 5C per 1,000 m elevation. Thus, at

................. 16157$

$CH1

02-08-07 10:22:02

PS

PAGE 7

8 / bruce m . b e e h l e r

sea level, in the forests near Timika, one will encounter an unpleasant combination of high humidity and warm temperature day and night on all but the coolest days of the austral winter. By contrast, at 4,000 m in the Sudirman Range one must expect regular night frosts during the dry season, when the skies are clear. Above 4,500 m periodic snowfalls are common. And glaciers cap the highest peaks of the Jaya Mountains (formerly the Carstensz Range). These glaciers expanded outward and downward during the Pleistocene cooling, melted altogether by 6,000 years bp, and returned during the recent cooling, only to begin retreating again in the last century. The elevation-temperature equation is a defining environmental phenomenon in mountainous Papua. This allows essentially distinct biotas to inhabit adjacent patches of land, separated only by elevation. It certainly explains much of the species-richness of Papua (beta diversity). Rain shadows exist in some interior valleys (such as the Baliem), on the Bomberai Peninsula, and in the Trans-Fly of the far southeast. Rainfall is also slightly attenuated along the northern coast, from the mouth of the Mamberamo east to Jayapura. Much of the interior receives well in excess of 3,000 mm/year. Papua is a land in flux. Significant chronic disturbance is produced by ongoing mountain-building in contest with rainfall-driven erosional processes, as well as by periodic vulcanism, human-caused and naturally occurring fire regimes, plus El Nin˜o droughts. Over the long history of human occupation, swidden agriculture has disturbed large swaths of habitat, most of which is now regenerated forest. Thus historical disturbance is a dominant factor dictating the distribution and pattern of today’s vegetation. Much of what appears to be ‘‘virgin rainforest’’ is, in fact, the product of recent and not-so-recent patch disturbance. This is abundantly evident when conducting plot-based plant surveys in the forest. Thus any attempt to characterize forest types is a rough generalization, and at best a qualitative assessment with minimal predictive power at the taxonomic scale.

forest types Closed forest is the default vegetation type over virtually the entirety of Papua except perhaps in the southeast (although the fire regime that produces savannas there may be anthropogenic). Papua’s forests are highly species rich, with minimal stand dominance by particular tree species, and with remarkable history-driven variation from site to site, even within single catchments. One-hectare stands of forest typically support between 70 and 200 species of trees larger than 10 cm diameter breast height (dbh). It is thus difficult to characterize the forest types of Papua taxonomically. Instead, forest types are delineated by elevation, rainfall, and structure. In general, New Guinea’s forests can be termed ‘‘tropical humid forests.’’ Tree species of the following families are important components of this tree flora: Podocarpaceae, Fagaceae, Moraceae, Lauraceae, Meliaceae, Myristicaceae, Sapindaceae, Euphorbiaceae, Combretaceae, Sapotaceae, Annonaceae, Clusiaceae, and Rubiaceae, among others (Oatham and Beehler 1997). In the lowlands, one finds tall alluvial forests in well-drained catchment basins,

................. 16157$

$CH1

02-08-07 10:22:02

PS

PAGE 8

Introduction to Papua / 9

as well as various types of periodically inundated swamp forests in the more poorly drained areas. The finest alluvial forests are grand, indeed, with emergent species reaching 60 meters (e.g., Octomeles sumatrana), and canopy height topping 45 meters. The canopy of this alluvial lowland forest is often irregular and broken, except where there has been uniform regeneration after some disturbance. Typical canopy tree genera of the wooded swamps include Barringtonia, Terminalia, Alstonia, Diospyros, Carallia, Syzygium, and Campnosperma. Palm swamps, dominated by sago, pandanus, or nipa are commonplace in the vast deltaic areas of the major rivers (e.g., Digul). These grade into herbaceous swamplands where inundation is the prevalent condition. Coastally one finds small strips of mangrove or large and extensive mangroves forests, depending upon conditions. These comprise species of Sonneratia, Xylocarpus, Brugiera, Rhizophora, and Avicennia. Mangrove formations are dominant in the south and southeast, between the southern Vogelkop and Bomberai Peninsula, and along the Waropen coast (northeastern Cenderawasih Bay). In the far southeast one encounters closed monsoon forest that grades southward into Melaleuca woodland and Eucalyptus savanna. Much of Papua is hilly, and here forests are on well-drained soils and tend to be less grand, with smaller-boled trees of lesser height. In the low hills on the southern side of the Central Cordillera above Timika one finds a very poor ‘‘white sand’’ and heath forest that is both structurally bizarre and taxonomically distinct. Above 1,000 meters one encounters submontane forests that in places have a strong representation of oaks (Castanopsis acuminatissima, Lithocarpus spp.) and several genera of Lauraceae. A cloud line settles on the mountains at varying elevations, depending on local conditions. This produces cloud forest conditions, which are typified by the abundance of moss on tree trunks as well as an effusion of epiphytes. This cloud line most typically can be found between 1,500 and 2,500 meters elevation. Midmontane forests are more species-poor and can be dominated by the Antarctic beech Nothofagus as well as several genera of gymnosperms from the family Podocarpaceae (Podocarpus, Dacrycarpus, Dacrydium, Phyllocladus). Above 3,000 meters, one encounters an elfin woodland that is low in stature (15 m), and small-boled (10–30 cm) and dense, with heavy mossing and with tangled moss-laden root mats on the ground in the place of soil. Climbing higher into the mountains, this leads to areas where patches of dense thicket-like dwarf forest is interdigitated with open boggy grasslands in the more poorly-drained and frost-prone areas. In these areas one can find prominent stands of large Dacrycarpus compactus as well as the more conifer-like Papuacedrus papuanus. On the summit areas above 4,000 meters one encounters a mix of tussock grasslands, rocky areas, low ericaceous thickets, and a variety of tropical alpine herbaceous vegetation. Botanically, Papua is remarkable, estimated to house more than 15,000 species of vascular plants, notably some 2000 species of orchids, more than 100 rhododendrons, one species of the great and ancient Araucaria conifers—Papua’s tallest tree, as well as the magnificent and valuable kauri pine (Agathis labillardierei). Dipterocarp trees are relatively uncommon, but appear in abundance in certain

................. 16157$

$CH1

02-08-07 10:22:03

PS

PAGE 9

10 / bruce m. b e e h l e r

patches, the result of some natural disturbance regime. Other important timber trees include Intsia bijuga (‘‘merbau’’), Pometia pinnata (‘‘matoa’’), Pterocarpus indicus (‘‘rosewood’’), and Dracontomelon (‘‘black walnut’’), among others.

Fauna

vertebrates Birds dominate the Papuan vertebrate fauna, with more than 600 species recorded. This includes more than 25 species of birds of paradise, three species of cassowaries, and some two dozen each of parrots, pigeons, raptors, and kingfishers. The mammals are less in evidence, mainly because of chronic hunting and their nocturnal habits. Fruit bats, insectivorous bats, tree kangaroos, possums, and rats are the best represented among the 180 or so species. Amphibians include more than 150 species of frogs, many still unknown to science. Reptiles include two crocodiles, 61 snakes, and 141 lizards. The fishes comprise ca 150 freshwater species and more than 2,250 marine taxa (about 1,500 of which inhabit coral reef ecosystems). Of special note are the 36 species of rainbow fish that inhabit Papua. This is an incomplete list, undoubtedly, and new taxa were described as recently as 1998.

terrestrial invertebrates The forest invertebrate fauna is diverse beyond imagination, defying our ability to enumerate it. There are probably in excess of 100,000 species of insects alone, only a fraction of these having been cataloged. Most prominent are the huge and beautiful birdwing butterflies, the giant phasmid stick insects, several lineages of giant beetle (longicorn, dynastine, etc.), and the world’s largest moth. One can also find freshwater crabs, a range of edible freshwater shrimp and crayfish, and an abundance of blood-sucking leeches.

marine life and coral reefs The marine reef environments found in Cenderawasih Bay and the Raja Ampat Islands are among the very richest on earth in terms of species diversity. One finds extraordinary numbers of hard corals, mollusks, and reef fishes. These environments are also very productive, and form an important sustainable resource for local communities. The region also supports a significant pelagic fishery, with key migratory species (such as various tuna).

Human Cultures

cultural setting Although the island of New Guinea is rather young in geological terms, its peoples are of apparently ancient stocks, and there is evidence that humans has been present on the island at least 40,000 years, perhaps longer. Not surprisingly, the details of the earlier habitation on the island are scanty, and it is possible that humans have occupied New Guinea for as long as 60,000 years. The whole island of New

................. 16157$

$CH1

02-08-07 10:22:03

PS

PAGE 10

Introduction to Papua / 11

Guinea supports more than 1,200 language groups. No other comparable land mass supports more languages. This could be taken as an indication of the longevity of human occupation of New Guinea. The Papuan half of the island supports about 250 languages (dwarfed by PNG’s 800 languages). We can offer no explanation as to why the west supports so many fewer languages, but physiographic and biogeographic diversity may offer a partial explanation (or it may be nothing more than sampling error—a nonconformity in classification methodology by scientists working in Asia vs. the Pacific). Many of Papua’s language groups are small and insular, with fewer than 1,000 speakers. A few other languages (e.g., Dani, Asmat) are spoken by many. These dominant languages seem to indicate cultural dominance as well. As with Papua New Guinea, the language diversity parallels diversity in local culture and thus Papua is culturally very diverse and heterogeneous. This is one reason there has been only limited local development in Papua. Small, diverse, egalitarian societies do not have the human capacity and structure needed for complex social and economic structures to develop, as has been explained eloquently by Jared Diamond in his book Guns, Germs, and Steel (1999). The absence of stratified societies and the lack of key domesticated livestock and grain crops has certainly contributed to the generally minor development of local economies in Papua. In one point of contrast, important sweet potato cultures in the fertile valleys of the central highlands have developed since the arrival of the sweet potato on the island—perhaps as little as 500 years ago. The major traditional population centers are found in the interior uplands (Baliem and Ilaga valleys, Paniai Lakes, and Arfak Mountains). Most societies are forest- or coastal-dwelling, with primary dependence upon sweet potatoes and pigs (interior) or fish and yams (coastal). It seems all New Guineans are accomplished gardeners as well as accomplished warriors. In most instances, the warlike traditions have been suppressed over the last century, mainly through the teachings of Christian missionaries.

history of western engagement and political history Papua was undoubtedly first contacted by Islamic traders from the west in search of spices and other exotic trade goods. The undocumented first contacts between the traders and the coastal Papuans perhaps first took place more than a thousand years ago. But initial trade was probably local—between Papuan people and those of Maluku just to the west and south. Major trade probably did not begin until after 1000 bce. The first Europeans to sail the coastline of Papua were Portuguese, in the 1500s, and they were followed by the whole cast of exploring nations (Spanish, Dutch, then English). These explorers were seeking trade routes as well as products to trade. This exploring era lasted from the 1500s to the early 1800s. It was followed by a period of regular trade (beˆche-de-mer or trepang, bird of paradise skins, turtle shell, massoi bark, etc.), which, in turn was followed by initial settlements (trade driven), then missionary activity. Early naturalist/explorers included Alfred Russel Wallace, who visited the Vogelkop (Bird’s Head) and Raja Ampat Islands in the 1840s, and Odoardi Beccari and Luigi d’Albertis, who visited

................. 16157$

$CH1

02-08-07 10:22:03

PS

PAGE 11

12 / bruce m. b e e h l e r

the Arfak Mountains in the 1870s. The Dutch made a preliminary claim to New Guinea west of the current border of 141 east longitude in 1826, but this infamous border was not formalized with the colonial powers of Britain until 1895 (in the south) and with Germany in 1910 (in the north). What followed was a rather weak attempt to establish government outstations, some rather stronger efforts to explore the interior (1900–1930), and to surmount Papua’s forbidding high peaks. Remarkably, the highest peak of Papua, Mt Jaya, was not successfully ascended until 1962 by Austrian Heinrich Harrer. Dutch, British, and American biological expeditions were conducted into the remote interior in the 1930s. Most famous was the Snow Mountains Expedition led by Richard Archbold, who discovered the populous Baliem Valley in 1938 during his aerial reconnaissance flights that allowed the expedition to ascend successfully high into the interior mountains. World War II brought this era of exploration to a close. After the War, independence issues dominated Indonesia and this led to the eventual annexation of Papua into the young Indonesian state in 1962. Indonesia has aggressively developed Papua through a bout of transmigration of landless poor from western Indonesia, through significant government and military oversight (which have included considerable conflict, tension, and bloodshed between Papuan ethnics and western Indonesians), and through natural resource exploitation (mining, fishing, logging). One expects this exploitation to expand considerably in the next several decades, and there is a question whether this exploitation will be predatory or, we hope, environmentally and culturally sustainable. Certainly that issue is a theme that runs through this book.

This Book and Its Goals This book, following the model of the preceding eight volumes of the Ecology of Indonesia series, seeks to provide a clear, comprehensive, yet concise account of the environment of this easternmost region of the vast archipelagic nation of Indonesia. The text is written with a university student in mind, but there is authoritative material that will be of interest to the serious academic researcher as well. We have departed from the plan of the original series in that we have sought out the world’s experts to contribute chapters on their specialties. In doing so, we have collected the very latest thinking on each subject. Through judicious editing, we have made certain that this cutting-edge material is accessible to the reader. We have attempted to avoid use of specialized and jargon terminology, or at least carefully defined these terms for the reader. Our goal is to have compiled a broad and comprehensive accounting of the natural history of Papua.

Literature Cited Diamond, J. (1999). Guns, Germs, and Steel: The Fates of Human Societies. W. W. Norton and Company, New York.

................. 16157$

$CH1

02-08-07 10:22:03

PS

PAGE 12

Introduction to Papua / 13 Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.). 1976. The Equatorial Glaciers of New Guinea. A.A. Balkema, Rotterdam. Petocz, R. 1989. Conservation and Development in Irian Jaya. Brill, Leiden. Oatham, M., and B. Beehler. 1997. Richness, taxonomic composition, and species patchiness in three lowland forest plots in Papua New Guinea. Pp. 649–668 in Dallmeier, F., and J. Comisky (eds.) Forest Biodiversity Research, Monitoring and Modeling: A Conceptual Background and Old World Case Studies. Parthenon Publishing, Casterton, UK.

................. 16157$

$CH1

02-08-07 10:22:03

PS

PAGE 13

1.2. Biological Exploration of New Guinea david g. frodin i o lo g i c al e x p lo r a t io n in New Guinea and its surrounding islands has a relatively long post-Columbian history, but until the 1760s it was casual, with ‘‘curiosa’’ and narrative descriptions the most tangible results. Even the ‘‘great voyages’’ of the subsequent decades paid but fleeting visits, with some— notably the Endeavour—actually rebuffed; the few contemporary attempts at settlement from outside were failures. Only in the last third of the nineteenth century did serious exploration begin, with the last large ‘‘white spaces’’ in the interior highlands ‘‘filled in’’ just as World War II approached. The marvelous birds of paradise, whose center of diversity is in mainland New Guinea, were among the first objects of natural history to attract attention from Europeans, but for long they were known only from their legless skins, obtained in direct or market trade. But the land soon came to be seen as hostile to settlement, even for the Portuguese and Spanish and, after them, the Dutch East India Company, so no extended surveys were made. Indeed, until the latter part of the eighteenth century (thus through most of Linnaeus’s lifetime) New Guinea was effectively ‘‘beyond the frontier,’’ with only its western fringes anywhere near a commercial realm (and so—fortunately for posterity—within the reach of Rumphius at Ambon). Otherwise, acquisition of geographical and natural history knowledge was casual, with Dampier among the few prominent contributors. The ‘‘great voyages’’ of the seven decades preceding 1840 did touch upon several parts of mainland New Guinea and its neighboring islands, with the naturalists of one voyage demonstrating that birds of paradise indeed had legs. Yet, although they established the main geographical outlines of the region, their visits were brief and their collections, though primary, were generally small and from but few localities. Apart from these contributions—not all of them fully reported upon— the only substantial collections until 1870 were those made in the late 1820s on the southwestern coast by Zipelius and Macklot and later—mainly in the Vogelkop (Bird’s Head) peninsula—by Wallace in 1858 and von Rosenberg from then through the 1860s. Not even the formal annexation of western New Guinea by the Dutch Indian government in 1848 provoked significant activity. The opening of the Suez Canal, the development of settlements in Australia, increasing commercial interest in the Pacific Islands, the growth of the plume trade, and scientific curiosity (particularly in the wake of Wallace’s Malay Archipelago) finally led to sustained outside interest in Papuasia and an opening up of its interiors. A veritable ‘‘rush’’ by explorers then ensued, particularly in the wake of the territorial acquisitions by Germany in the northeast and its large island neigh-

B

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

14

................. 16157$

$CH2

02-08-07 10:22:05

PS

PAGE 14

Biological Exploration of New Guinea / 15

bors, and Britain in the southeast—all under the gaze (and even sponsorship) of the now well-developed popular press. By 1914 very considerable progress had been made, with after 1900 greater official interest—but largely in the Dutch and German spheres, surpassing the very effective work by Macgregor, administrator in British New Guinea over the decade leading up to 1898. Sadly, that was succeeded by relative indifference—particularly after 1901 with transfer of control to Australia. After World War I (though slightly later in western New Guinea), the rest of New Guinea also became something of a ‘‘backwater’’—with few official undertakings in natural history. Exploration did, however, continue—though largely under outside sponsorship—leading to further major discoveries, particularly in the 1930s. By the end of that decade, the major outlines of the biota had become known—particularly after the prodigious efforts of the Third Archbold Expedition—and the age of ‘‘primary’’ exploration was over. In contrast to the ‘‘Great War,’’ during World War II New Guinea and its islands were a major theater of conflict, greatly increasing the region’s profile. The stage was now set for three decades of ‘‘secondary’’ exploration, much of it under the auspices of the administering countries (including their ‘‘metropolitan’’ organizations), and the establishment of local collections and research facilities. Through the 1960s, substantial resources were allocated to land, agricultural, forest, and marine surveys in both east and west; in 1959 the Dutch mounted a final, ‘‘major’’ exploring expedition to ‘‘the last white spot on the map,’’ Juliana Top (now Mt Mandala) and the western Star Mountains. There was also much extraofficial exploration and other activity, including the establishment of biological stations, beginning with what is now the Wau Ecology Institute, set up in 1961 by the late J. Linsley Gressitt. After transfer of control of western New Guinea to Indonesia (1963/1969) and, in 1975, the independence of Papua New Guinea (PNG), official efforts fell away— particularly after 1980 in PNG. Biogeography and Ecology of New Guinea (1982; see section on ‘‘Collections’’ below) could thus be said to mark the end of an era. Individual and group exploration and research (under sponsorship or otherwise) has, however, continued over the subsequent quarter-century, now with ecology, conservation, and ‘‘sustainability’’ as guiding themes. In this, nongovernmental organizations (NGOs)—including universities in both west and east New Guinea as well as research stations—have played an increasing role. Though progress— perhaps inevitably—has been fragmented, there have been significant achievements, some of them recorded in the Post-World War II section, below. Future exploratory efforts should focus particularly on poorly known areas (documented in recent conservation assessments; see References section, below). But at the same time work must continue on consolidating, enhancing, preserving, and making more available our knowledge of what we already have to hand—not easy in the face of competition for research resources, changing interests, insecurity, and indeed a fall in new entrants to the sciences.

................. 16157$

$CH2

02-08-07 10:22:05

PS

PAGE 15

16 / david g. f r o d i n

Before the Rush: Early History (1500–1875)

when birds of paradise had no legs (1500–1815) 1500–1760 Settled by humans in the late Quaternary, with two further waves of immigrants respectively in the early and post-historical Recent, New Guinea and its islands— particularly after the Lapita migrations—were initially visited by Malay (and perhaps also Chinese) traders from Dobo (in the Aru Islands) and elsewhere. The earliest recorded explorers were, however, post-Columbian Europeans, sailing from both west and east partly in search of the ‘‘great southern land’’ then thought to be necessary to balance the large masses in the north, particularly Eurasia. Not until the 17th century—and passing into general knowledge only much later— were the northern fringes of the supposed southern landmass shown to be a great island. Although Magellan’s expedition—to which we owe the first European use of the word ‘‘Papua’’ and knowledge of its birds of paradise—sailed near New Ireland in 1521, the first to arrive in the waters off the mainland was the Portuguese Jorge de Meneses in 1527. But, reaching only as far as Biak and the north coast of the Vogelkop Peninsula, he would have had no idea of its extent. He was followed in 1528 by Alvaro de Saavedra and in 1537 by Hernando de Grijalva, neither in turn venturing beyond Yapen and Biak. In 1545 Ynigo Ortiz de Retes, also—like Saavedra—in an attempt to sail to Mexico, reached as far as Manam and the Schouten Islands (off the mouths of the Ramu and Sepik) as well as the western Admiralty Islands, Aua and Wuvulu, before having to turn back to Tidore. On the voyage back he called in near present-day Sarmi and named the land ‘‘Nueva Guinea’’ because the people looked like Africans. Entry from the Americas only came later: in 1567—two years before Mercator’s world map first appeared—the Spaniard Alvaro de Mendan˜a discovered the later-lost Solomon Islands. He attempted a return in 1595 but died at sea, with Pedro Ferna´ndez de Quiro´s eventually taking command of that expedition. In 1606 Quiro´s, with papal and other support, once more sailed to the South Pacific, discovering what is now Vanuatu; but there his expedition fell apart. His associate Luis Vae´z de Torres—aided by the southeasterly trade winds—continued west towards New Guinea, reaching the present Milne Bay Islands near Samarai (where, on Sideia, the company dined on what is now the first record of a Papuasian marsupial) and afterwards sailing along the south coast and traversing the strait now named after him. The first Dutch voyage to the East Indies set sail in 1597 and soon afterwards contacts by Dutchmen with New Guinea began in earnest. Willem Jansz on the Duyfken in 1606 may have been the first; but more important was the voyage of Willem Schouten and Jacob Le Maire in 1616, who viewed much of the north coast along with Manus Island and present-day New Ireland. Both voyages suffered massacres. In 1623 Jan Carstensz sailed along the southwestern coast and was the first to see—in disbelief—the snow- and ice-capped highest peaks of the

................. 16157$

$CH2

02-08-07 10:22:05

PS

PAGE 16

Biological Exploration of New Guinea / 17

mainland which for long bore his name (but now collectively are Mt Jaya). Two decades later, in 1643, Abel Tasman touched upon New Ireland and, for the first time, sailed by the north coast of what is now New Britain, but yet thought them continuous with New Guinea (and New Holland, as Australia was then known). But this would be the last such effort on their part: Jan Compagnie was a commercial enterprise, and of potential profit little was to be seen. Contacts with New Guinea became largely restricted to its western fringes, with which an active trade would be carried on and whence G. E. Rumphius, from 1653 at Ambon in the Jan Compagnie’s service for half a century, received much valuable information, in time incorporated into his famous Thesaurus amboinensis (1705) and Herbarium amboinense (1741–1750, 1755). In 1700 came the epoch-making visit by the Englishman William Dampier—the region’s first ‘‘enlightened’’ explorer. In his ancient ship Roebuck he visited the north coast and discovered Mussau and Emira north of New Ireland, sailed along the north coast of New Ireland and then past his ‘‘St George’s Bay’’ along the south coast of what was still thought to be a large peninsula. After discovering the deep strait between it and New Guinea, he bestowed on what was now an island the name ‘‘Nova Britannia’’ (New Britain) by which it has been known ever since (save as ‘‘Neu-Pommern’’ during German rule). But the ex-buccaneer and explorer was also a natural historian and collector, so he brought back ‘‘curiosa’’ for appreciation and study: the earliest scientific specimens from the region (apart from those from Rumphius surviving in Florence). Publication of Dampier’s A Voyage to New-Holland (1703) stimulated further coastal exploration over the subsequent three decades, particularly in the west (with Dampier himself returning in 1705), and in 1714 the Sultan of Tidore ceded his territories in New Guinea (with the southern Moluccas) to the Dutch. But it was the now-ascendant French and English who were to set an entirely new trend. Taking a cue from Dampier, most voyages from the 1750s onwards involved serious scientific work as well as exploration and contact, and carried naturalists or physician-naturalists.

1760–1815 The first of the ‘‘new’’ expeditions was British. In 1767 Philip Carteret in the Swallow visited parts of the southwestern Pacific, the northwestern Solomons and ‘‘greater’’ New Britain—and found that ‘‘St George’s Bay’’ was a channel. He thus gave the northern island its name of ‘‘Nova Hibernia’’ (New Ireland), so-called ever since (save as ‘‘Neu-Mecklenberg’’ during German rule). Significantly, Carteret discovered some safe anchorages at its southwestern end (including Gower Harbor—soon afterwards named ‘‘Port Praslin’’ by Bougainville and visited by many later expeditions, and in the late nineteenth century the scene of the tragic ‘‘Nouvelle France’’ settlement scheme). Carteret was followed in 1768 by the Frenchman Louis Antoine de Bougainville who, in two ships (La Boudeuse and L’E´toile) and with Philibert Commerson as naturalist, visited among other places parts of the southeastern coast, the Louisiade Archipelago, the northern Solomon

................. 16157$

$CH2

02-08-07 10:22:05

PS

PAGE 17

18 / david g. f r o d i n

Islands (notably those now known as Choiseul, Bougainville, and Buka), and southwestern New Ireland before hastening westwards to Java to relieve his crews. In 1770 James Cook, with J. Banks and D. Solander, definitively verified New Guinea’s distinctness from New Holland (now known as Australia) by sailing through Torres Strait. Beyond that treacherous passage, the Endeavour only landed on the southwest coast for one day, where Banks made some thirty plant collections—of which a list survives—under protection of the ship’s guns and marines. The French now used the new knowledge of the region, particularly that gained by Bougainville and Commerson, for economic gain—their governor in Mauritius, Pierre Poivre, was determined to break the Dutch spice monopoly. With advice from Commerson (who had joined Poivre’s service), Simon Provost in 1769–1770 (as part of an expedition on two ships, L’E´toile du Matin and Vigilant) and then Pierre Sonnerat in 1771–1772 (under Provost), as part of extensive missions in the Moluccas and Philippines, reached Gebe´ (near Gag) in the extreme west of Papuasia; but they touched neither on other New Guinean islands nor the mainland (in spite of the title of Sonnerat’s popular 1776 book, Voyage a` la Nouvelle-Guine´e). Economically, however, the French voyagers were successful; the principal spices came to be established in the Mascarenes and elsewhere, in time contributing to the demise of the Dutch East India Company. Sonnerat’s natural history collections (Paris) are primarily zoological; he also lives on in the epithet for one of that museum’s collections databases. In 1781 a Spaniard, Francisco Antonio Maurelle, discovered more isles of the Bismarcks including the Los Negros near Manus; but still the north coast of New Britain remained poorly known. This would be partially remedied in the next decade. In 1792 and 1793 another French world voyage—charged by Louis XVI with searching for the lost expedition of La Pe´rouse and under the command of A. R. J. de Bruny d’Entrecasteaux—was in New Guinea waters with La Recherche and L’Espe´rance. The two ships called at several points, including for the first time Huon Gulf (named after the Espe´rance’s commander, Huon de Kermadec); other important work was done in the Milne Bay region, southwestern New Ireland, around the Bismarck Sea, and on tiny Rawak off Waigeo. Many well-known and still-current geographical names were at this time introduced. D’Entrecasteaux’s naturalists were J. J. Houtou de La Billiardie`re, Louis Ventenat, L. A. Deschamps, and Claude Riche, with Fe´lix de Lahaie accompanying them as a ‘‘gardenerbotanist.’’ Sadly, the commander died at sea west of Manus on 20 July 1793 and later, in Java, the expedition broke up in confusion over the consequences of the French Revolution (A. Hesmivy d’Auribeau, second in command, was a staunch royalist—but died in 1794 just before capture, while La Billiardie`re led the republican faction). La Billiardie`re’s collections (and those of others) were confiscated by the Dutch and sent to England, but through Banks’s good offices restored to him a few years later (his plants are now in Florence). Unfortunately for Papuasia, he published only on his Australian and New Caledonian plants (the former in 1804– 1807 as Novae Hollandiae plantarum specimen, the latter in 1824–1825 as Sertum austro-caledonicum). Lahaie’s own collections are in Paris (and Geneva).

................. 16157$

$CH2

02-08-07 10:22:06

PS

PAGE 18

Biological Exploration of New Guinea / 19

reality, disappointment, and renewal (1815–1875) 1815–1850 The wars and disruptions of the French Republican and Napoleonic eras were to restrict exploration for the next 20 years or so, but after 1815 a new flowering took place, associated with the growth of mercantile trade and the related development of detailed marine charts—the latter first undertaken on a large scale by Flinders in the Investigator. In New Guinea and its islands the quarter-century from 1815 was dominated by several great French voyages—all with naturalists—which collectively added substantially to natural history knowledge and amassed considerable collections (now in the Natural History Museum in Paris, though many perhaps remain littleknown or even undocumented). The voyages were part of a diplomatic and mercantile initiative, intended to show that after all its humiliations France still mattered—but, save for French Polynesia (and, somewhat later, New Caledonia), they did not lead much to new overseas territories (although the French claim to ‘‘Ade´lie Land’’ in Antarctica dates from the visit there by the last of these expeditions). Inspired by the example of von Humboldt, the collections, elaborated by professional naturalists, formed a basis for many sumptuous publications—these in turn inspiring the undertakings of the late nineteenth and early twentieth centuries. First into New Guinea waters were the Uranie and Physicienne under Louis de Freycinet in 1818–1819. His naturalists were Jean R. C. Quoy and Joseph Gaimard with Charles Gaudichaud-Beaupre´. They called, however, only at Rawak (off Waigeo—earlier visited by d’Entrecasteaux’s expedition; see above)—en route to Guam. While returning to France, the Uranie was lost in the Falklands (also known as Malvinas) Islands, though many collections survived (and others were added). Freycinet was followed in 1822–1824 by the Coquille under Louis Duperrey; accompanying him were Jules Se´bastien Dumont d’Urville, Prosper Garnot, and Rene´ Lesson. That team collected insects, birds, other animals, and plants in the Solomons, Port Praslin (southern New Ireland), Rawak (see above) and Dore´ Bay (Manokwari, in the Vogelkop Peninsula), this last spot a new frontier for science— though first surveyed in 1775 by Thomas Forrest. Lesson in particular there collected and studied birds of paradise, and was the first to learn—three centuries on from when, in 1522, skins of Paradisaea minor had reached Seville with the Vittoria—that they had legs; but his collections (and, particularly, living individuals) then helped to create the more than half-century-long fashion in Europe (and elsewhere) for their feathers—and, in turn, contribute to popular conservation awareness. Later Dumont d’Urville led two more expeditions through the region: the first during 1826–1829 in L’Astrolabe (ex-Coquille), with Quoy, Lesson, and Gaimard as naturalists, calling at New Ireland, the present Astrolabe Bay off Madang, Dore´ Bay, and Waigeo as well as—for the first time after Dampier—sailing along the south coast of New Britain (naming, among other places, Cape Merkus and Jac-

................. 16157$

$CH2

02-08-07 10:22:06

PS

PAGE 19

20 / david g. f r o d i n

quinot Bay, the latter after his second-in-command Charles Hector Jacquinot); and the second in 1838–1839 and 1840 as part of his voyages to the South Pole in L’Astrolabe and (under C. H. Jacquinot) La Ze´le´e with naturalists Jacques Hombron, Honore´ Jacquinot, and Elie J. F. Le Guillou—respectively specializing in zoology, botany, and entomology. Around New Guinea they called at various points: Triton Bay (where the Dutch settlement had then been recently abandoned), the Louisiades (to complete d’Entrecasteaux’s surveys), and also sailed along the southeastern coast (naming the Varirata ridge near present-day Port Moresby as the Astrolabe Range). Other nations, however, were not inactive. In 1820 the Dutch, with the Indies restored to them (and accorded international recognition from 1824), set up under Willem I a ‘‘Natural Sciences Commission’’ (Natuurkundige Commissie). Over the next thirty years they were to make extensive expeditions, inland as well as coastal, in the still poorly-known archipelago—but mortality was high. Among these was, in 1828, a visit to New Guinea. In connection with a projected settlement, A. J. van Delden with the Triton and Iris led a surveying expedition along much of the southwest coast. Accompanying van Delden were Commission members Heinrich C. Macklot, Alexander Zipelius, and Salomon Mu¨ller, the last the first to document the marked zoological differences between the western and eastern parts of the Indies. They were accompanied by two artists, P. van Oort and G. van Raalten. The settlement—known as ‘‘Merkusoord’’ after then then-Governor of the Moluccas and one of the promoters, Pieter Merkus—was established in the lands of the Lobo at Triton Bay (not far east of present-day Kaimana) and protected by a fort, ‘‘du Bus’’ (after the then-Commissar-General of the Indies, Leonard du Bus de Gissignies). All these names have been used in collections and literature and here are set out for convenience. But the settlement did not last long; and of the naturalists and artists only Mu¨ller was to survive early death or (in Macklot’s case) murder. Their collections—the first significant lot from this part of New Guinea and for decades one of the few available—made their way to Leiden in the Netherlands and were variously written up by Temminck, Blume, Mu¨ller, and others. From 1840 the British returned, but—like the Dutch—were now concerned as much with detailed coastal and hydrographic survey as with primary exploration. This continued a tradition begun with the Investigator under Flinders and skillfully developed over the middle decades of the nineteenth century (and since, with modifications). Such surveys—tedious but essential in a new and increasingly global age of commerce and settlement—did, however, continue to provide opportunities for natural history research. Indeed, it was on such a voyage that the young Charles Darwin sailed with Robert Fitzroy in 1831–1836. After 1850, however, surveys of Australasia and the western Pacific were largely conducted from Sydney rather than London. The first of the Royal Navy vessels to sail through New Guinea waters were the Sulphur (with the Starling) under Edward Belcher (who had succeeded F. W. Beechey on what had become an ‘‘interminable voyage’’). Himself strongly inter-

................. 16157$

$CH2

02-08-07 10:22:07

PS

PAGE 20

Biological Exploration of New Guinea / 21

ested in natural history and assisted by R. B. Hinds and A. G. Barclay, respectively as naturalist-surgeon and gardener-botanist, Belcher called in to the Solomons, Port Praslin (southwestern New Ireland), Kairiru off the north coast, and Yapen, collecting some animals and plants (now at London: BMNH, Kew)—though with rather less profit than in the eastern Pacific and the Americas, the voyage’s main objectives. The Sulphur was soon followed by two more focused voyages to the south, reflecting the increasing importance of the future Australia and the passage between it and New Guinea, the treacherous Torres Strait—now becoming a key route between India, Southeast Asia, and New South Wales. The 1842–1846 voyage of the Fly and Bramble under F. P. Blackwood with, as geologist, J. Beete Jukes and a naturalist-artist, John MacGillivray, focused in particular on the Torres Strait and the northern Australian coast, but also (in 1845) examined the western Gulf of Papua and discovered the Fly and Turama rivers, sailing some ways up the Fly. Blackwood’s work was continued by the Rattlesnake under Owen Stanley in 1846–1850, with particular attention to the Louisiades, the future China Strait, and the southeastern coasts (including Yule Island); he was accompanied by MacGillivray and the future evolutionist T. H. Huxley (who in particular collected cnidarians and mollusks). The emphasis on natural history in both voyages was on geology, zoology, and marine biology, with but few land plants collected (all at BMNH). Jukes and MacGillivray, respectively, wrote the narratives of these last two voyages, Owen Stanley sadly having died at Sydney before the return (under C. B. Yule) of the Rattlesnake to Britain via Cape Horn. But, in contrast to the work of the Sulphur, publication of the scientific results would be somewhat piecemeal—times were harder, and the Admiralty was more interested in those of the Southern Ocean voyage of the Erebus and Terror. All this exploration was, however, not followed by much settlement. An attempt had been made by an English party in 1795 at Dore´ Bay, but until the last quarter of the nineteenth century that at Triton Bay mentioned above would be the most serious effort. But it, too, would soon be defeated by disease and a hostile environment. Missions also remained few and far between; that of the Congregatio Mariae at Woodlark Island—being the first in the east, following proclamation of the apostolic vicariate of Melanesia by Pope Pius IX. Though after some years also suffering the fate of Merkusoord, the station was during 1847–1852 a collecting locality for one of the Marist priests, Pe`re J. Xavier H. Montrouzier—the earliest French missionary-naturalist to be active in the New Guinea region. At and around the settlement he collected insects, mollusks, and fish (Paris, partly lost), but no plants. From Woodlark he moved to New Caledonia, settling on the northwesterly Ile Art. In 1855 he published (at Lyon, France) his pioneer Essai sur la faune de l’ıˆle de Woodlark ou Moiou.

1850–1875 With the departure of the Rattlesnake, the age of the major exploring and survey expeditions for the New Guinea region was over. The way, however, had been paved for safe passage of commercial shipping, including the new mixed steam

................. 16157$

$CH2

02-08-07 10:22:07

PS

PAGE 21

22 / david g. f r o d i n

and sail ships. But official interest remained relatively low in this third quarter of the nineteenth century save for its last five years or so when potential annexations loomed. Instead, it was independent, often privately sponsored naturalists— particularly in the west, closer to the developing East Indian shipping network and its connections to Australia, Asia, and Europe—who came to dominate natural history exploration for these years. Some, such as Miklucho-Maclay as well as Beccari and d’Albertis, had official assistance in the form of passage on naval vessels. Extension of mission networks provided other opportunities, notably at Dore´ Bay, the Torres Strait Islands (from 1871; see below), Port Moresby (1874), and the Duke of York Islands (1875); some of the missionaries themselves made collections and sent them ‘‘home.’’ The first was none other than the most famous: Alfred Russel Wallace, who during his sojourn in Malesia made two visits to the New Guinea region. In 1857 he visited the Aru Islands, while in 1858 he spent a few weeks collecting at Dore´ Bay (primarily insects but also birds). Himself largely ill and coast-bound, from there his collectors went into the Arfak Mountains, achieving many new finds. In 1860, he collected (and observed) on Gam Island and Waigeo, while his assistant Charles Allen reached Salawati, Misool, and the Sorong area of the northwestern Vogelkop Peninsula. Their collections are mainly in London (BMNH). Wallace was followed by other naturalists—some of them colorful—who also sought out the northwestern peninsular region and the Raja Ampat Islands to the west. The sword-carrying German Baron C. B. H. von Rosenberg collected birds in several visits over 1858–1870 (Leiden, BMNH), meeting Wallace at Dore´ Bay and overlapping with Allen in Misool; while in 1864 Heinrich Bernstein collected animals at Sorong, Waigeo, and Salawati. But going into the last third of the nineteenth century—particularly with the opening of the Suez Canal and the publication of The Malay Archipelago by Wallace, both in 1869, as well as the spread of the world steamship network—outside contacts increased rapidly. In 1871 there came the first European resident in the east (after Montrouzier and the other Marists at Woodlark), the now almost legendary Russian ethnologist-naturalist N. N. de Miklucho-Maclay. He was landed near Bongu on Astrolabe Bay by his country’s Vitiaz—a name now given to the deep strait between New Guinea and New Britain, first traversed by Dampier— and remained there for over a year. He returned to what is now the Rai Coast in 1876–1877, 1878, and 1883, and at other times visited Triton Bay (the former Merkusoord), Gebe´, and (in 1880) the Torres Strait Islands as well as Samarai (near China Strait—the latter by then becoming a trading post taking advantage of the growth of local commerce as well as a key new shipping route established following Moresby’s surveys; see below). The Russian collected some animals, a few plants, and much ethnographic data; in addition he introduced some fruits and other plants—among them papaw (papaya), Carica papaya (‘‘banana bilong Maclay’’). Many of his specimens and data were lost, though some insects were described by Sir William J. Macleay in Sydney. His New Guinea Diaries (1975, translated and edited by C. L. Sentinella) and Travels to New Guinea: diaries, letters,

................. 16157$

$CH2

02-08-07 10:22:08

PS

PAGE 22

Biological Exploration of New Guinea / 23

documents (1982, compiled by D. Tumarkin) along with two biographies (Who travels alone (1944) by F. S. Greenop and The Moon Man (1984) by E. M. Webster) cover his travels and in particular give an interesting picture of the untouched north coast more than a century and a quarter ago. Also in 1871, the London Missionary Society made its first landings in the region, Samuel Macfarlane (later senior missionary) and A. W. Murray reaching the Torres Strait Islands and other points along the south coast. Later, from bases at Cape York and (after 1877) at Maer (Murray) Island in the Strait, the mission under Macfarlane—with the aid of a small ship, the Ellengowan—would establish a number of stations over a wide area. These included, as already indicated, a station at Port Moresby, and, in 1877, one at South Cape (Suau). This gave him many opportunities for exploration, yielding some collections (plants, Melbourne); he also sailed with d’Albertis and Macleay (see below). In the west, the appearance of outside powers in the waters of the Archipelago now spurred the Dutch Indian authorities into some action with respect to their lands, including New Guinea; the ‘‘tempo dulu’’ of the past was about to recede. In August 1871 the steamer Dassoon (under Capt. A. Smits) with the smaller Wilhelmina Frederika and with two Tidorean chiefs (brothers of the Sultan, who retained some residual rights), P. van der Crab on behalf of the Indies government, and, as botanist, J. E. Teysmann (from Bogor, now acting as an agent for the new director of the Botanic Garden, R. H. C. C. Scheffer), sailed from Ternate. Over some three months they called at several points in Papua, reaching east to Humboldt Bay (and also for a distance east of 141, the then-nominal border). Considerable plant collections were made (Bogor, Melbourne, Leiden). They were written up in 1876 by Scheffer—part of the Garden’s first steps towards an independent scientific existence. Of other biota there was obtained but little—a disappointment, if less deathly than for the Triton and Iris in 1828. The fame of Wallace and his book—and even more the fabled birds of paradise, whose feathers were now becoming seriously fashionable—now brought a stream of other visitors, including many naturalists. A. A. Bruijn (from Ternate) collected in 1871–1879, partly for the plume trade (birds, Tring/AMNH and BMNH). In 1872–1873 the Italians Odoardo Beccari (an all-round botanist and later a famous palm specialist) and Luigi M. d’Albertis came to Dore´ Bay, from there climbing into the Arfak Mountains as far as the Hatam district—Beccari there obtaining the first botanical collections from anywhere in the mountains of New Guinea, as well as insects and other zoological materials (Florence, Genoa; d’Albertis had trained at Genoa’s Museo Civico di Storia Naturale under its head curator, Giacomo Doria). Ramoi (south of Sorong), Mt Epa, Andai, and some of the Raja Ampat Islands were also visited (as well as, in 1873, the Aru group). Almost at the same time, there came to the northwest A. B. Meyer from Dresden, specializing in zoology and ethnography; but his extensive itinerary and localities were at least partially falsified or fictitious. In 1874 the great Challenger oceanographic expedition (of 1872–1876) sailed for the first time into New Guinea waters, voyaging westwards via the Torres Strait Islands and visiting, in September, the Aru (and

................. 16157$

$CH2

02-08-07 10:22:09

PS

PAGE 23

24 / david g. f r o d i n

Kai) Islands with H. N. Moseley there making plant and animal collections (Kew, BMNH); the next year the ship would sail along the north coast and visit the Admiralty Islands (see below). In 1875–1876 Beccari returned to the Vogelkop Peninsula, but this time on his own; during a long stay—which included another ascent into the Arfaks—he also visited Yapen and Biak as well as most of the main Raja Ampat Islands (Misool, Batanta, Salawati, Kofiau, and Waigeo, some for the second time). But even to this day his substantial plant collections have never fully been worked up, though he is commemorated in many binomials. His former companion d’Albertis now turned to the east. A Garibaldi veteran and in recent years viewed as one of the most notorious of adventurer-naturalists ever to visit New Guinea, he made in 1875–1877 three trips to present-day Papua New Guinea: the first with Macfarlane (see above) for some distance up the Fly River as well as staying on his own for some time on Yule Island (there meeting with the Chevert under Macleay; see also below); and his more famous second and third trips far up the Fly in the steam launch Neva, in 1876 as far as the foothills of the central ranges—the furthest stab into the dark interior hitherto made by an outsider. Substantial collections of ethnographic material as well as plants and many animals were taken back (Genoa, Florence, Melbourne); his travel book, Alla Nuova Guinea, appeared (1880, in the same year also released in English as New Guinea: what I did and what I saw). A string of species was named after him including a ring-tailed possum, Pseudochirops albertisi and the Buff-tailed Sicklebill, Epimachus albertisi. The increasing level of activities in and around New Guinea (as well as the labor trade of Queensland) not unnaturally attracted renewed British naval attention. As a result, two final major surveying voyages were undertaken from Sydney by the steam/sail ship Basilisk under the command of Capt. John Moresby. His first voyage of six months in 1872–1873 made further detailed coastal surveys along the south and southeast coasts and the nearer Milne Bay Islands, including China Strait, which then received its name. With surer propulsion and maneuverability, he also was able to penetrate the long fringing southern reef in several places and so become the first outsider to see Fairfax Harbor (Port Moresby) on 21 February; the reef passage was (and is) named after his ship. A Congregational mission station was set up beside the bay the next year, paving the way for serious interior exploration and (from 1886) outside settlement. In early 1874 Moresby returned and continued his detailed surveying, now traversing the northeast coast and by mid-May reaching Huon Gulf (before sailing around to the north coast and on to Ambon and, finally, Britain). On both voyages Moresby was accompanied by Peter Comrie as naturalist; he collected insects, mammals, and birds but very few plants, and published but little. As with the Sulphur in the Pacific and, to a lesser extent, the Rattlesnake, a good opportunity for significant scientific contributions in monographic form was not taken up or lost due to official opposition, parsimony, or narrowly-drawn instructions—or passed over in favor of the Challenger results. But, in any case, by 1875—the year the Challenger sailed through the northern parts of the New

................. 16157$

$CH2

02-08-07 10:22:09

PS

PAGE 24

Biological Exploration of New Guinea / 25

Guinea region—a single shipboard naturalist could not handle all the requirements demanded of the different departments of natural history, now quickly deepening and further diversifying.

‘‘Guinea Gold’’: The Scramble for Specimens and Species (1875–1914)

southeastern new guinea (british new guinea, later papua) Pre-proclamation Years (to 1884) The discoveries and surveys of John Moresby appeared in popular form in London in 1876 and quickly were reviewed in the Australian press. Not long before, the Challenger—which had visited the continent during the first half of its voyage— focused a good deal of attention on natural history. This coincided with the first wave of nationalism in Australia, by then relatively prosperous from wool and mining; indeed, for some time the colonies were per capita collectively among the wealthiest polities in the world. Settlement was already looking beyond Queensland, and there also was a fear of covetous outside powers with their own interests in trading and labor recruiting. The great business houses of Towns (commemorated in Townsville), Burns Philp (founded 1875, also at Townsville), and others were coming into existence, with Burns Philp early becoming interested in Pacific trade. Miklucho-Maclay’s presence in Sydney in the 1870s (and his marriage there to a socially prominent young woman) as well as his writings also attracted metropolitan attention to New Guinea. All this was embodied in Sir William Macleay’s private natural history and marine-biological voyage in 1875 in the old sailing ship Chevert; its departure from Sydney on 18 May was a festive occasion. Natural history was at the height of its social popularity and Macleay was not only a notable scientist but also a wealthy parliamentarian and socialite—his grandfather Alexander having come out from Britain as Colonial Secretary fifty years before. A number of ‘‘scientific men’’ accompanied him on the voyage. Following an interest expressed by Ferdinand von Mueller in Melbourne, J. Reedy, a gardener from the Macarthur estate outside Camden, was also taken on to handle plant introductions and collecting (Melbourne; we owe to him the first collections of eucalypts from southern New Guinea). Regrettably, however, due to conflicts aboard ship as well as adverse weather (the course of the expedition largely coinciding with the southeastern monsoon) and other factors, geographical discoveries were few although the marine zoological collections would be considerable (Macleay Museum/AM). At Somerset near Cape York came a meeting with Macfarlane, who accompanied them in the Strait and along the nearby mainland coasts and mouth of the Fly. At Yule Island, d’Albertis’ simultaneous presence also had an effect. The expedition returned to Cape York by early September and the Chevert then sailed back south to Sydney; but over time only few publications resulted. Succeeding expeditions from Australia would generally be more modest undertakings.

................. 16157$

$CH2

02-08-07 10:22:10

PS

PAGE 25

26 / david g. f r o d i n

Steamship services advanced rapidly through the 1870s to and from Australia and along the Queensland coast; by 1877 traffic through the treacherous but now relatively well-surveyed Torres Strait was such that the Queensland Government established Thursday Island as a northern station (with pilotage) in place of Cape York. This enhanced access to the Torres Strait Islands and the adjacent New Guinea mainland. Two years later the surface border was proclaimed, annexing all the Straits islands—including some very close to the mainland—to Queensland, a protocol which remains (although the seabed border is now slightly to the south). Boom-town Cooktown became a favored transfer point, especially for Samarai and the northern areas. Thus the southeastern parts of New Guinea became relatively accessible and as frequented as the Vogelkop Peninsula, albeit more by British and Australian collectors (although Italians too were notable in the early years, as we shall see). As already noted, the establishment of a London Missionary Society (Congregational) station at Port Moresby—initially under the leadership of W. G. Lawes— was crucial for the beginnings of inland exploration and potential settlement. First into the field were Kendall Broadbent (1873–1879; collections, Pittsburgh) and Andrew Goldie (mainly from 1876–1882, before becoming largely a storekeeper and so remaining until 1890 when he sold out to Burns Philp). The Sogeri Plateau was reached first by O. C. Stone in 1875 and then by Goldie in 1877. Goldie was in 1875–1877 joined by Morton from the Australian Museum (with in late 1877 a visit to the Louisiades, not long before the first discoveries there of gold) and in 1878 by the German collector Carl Hunstein. Over the next half-decade Hunstein, as Goldie’s assistant, traveled extensively in the difficult ranges beyond the Sogeri Plateau (east and northeast of Port Moresby) and therein discovered several new birds of paradise, so establishing his reputation. Goldie himself obtained many animals (Sydney) and plants (Melbourne). Although outside interest soon fell (as settlement prospects proved disappointing), in the years leading up to 1884 collecting by the residents continued, for plants (Melbourne) with much encouragement from von Mueller (who from their collections published many new species and records). In addition to Goldie and Hunstein, major contributors were Lawes, another missionary, James Chalmers (from 1877), and their assistants (particularly Jakoba). Over the years 1879–1885 all made sometimes lengthy tours, Chalmers for a time from a base at Suau (South Cape, Milne Bay) but later from Port Moresby. In 1882 Goldie went once more to the east, visiting the D’Entrecasteaux Islands; in 1884 Chalmers accompanied the annexation squadron (and with Cyprian Bridge ascended the Cloudy Mountains); and in 1885 he sailed with Scratchley (see below) on an extensive familiarization cruise, also making some further inland explorations—in part as a ‘‘mediator,’’ he being held in high regard— before proceeding on furlough in 1886. The attempted annexation in 1883 of all eastern New Guinea by Queensland (under then-Premier McIlwraith) proved a sensation—and in short order was disowned by Britain. W. E. Armit, a tough former policeman (and naturalist) then on assignment for a Melbourne newspaper, came in mid-year with three Dentons,

................. 16157$

$CH2

02-08-07 10:22:10

PS

PAGE 26

Biological Exploration of New Guinea / 27

all from Massachusetts, U.S.: father William (who died in the field) and sons S. W. and S. F. They traveled across the Astrolabe Range through Sogeri to the Moroka (Meroka) district (northeast of Iawarere, in the upper Musgrave River basin, an area earlier visited by Hunstein and to this day still remote, without road access); Armit collected some plants (Melbourne). The next season, Armit continued collecting in the Kabadi district (inland from Yule Island), penetrating into the foothills (and discovering, among other plants, Buddleja asiatica), and also in the Milne Bay region; and likewise in 1884 E. G. Edelfelt visited the Astrolabe Range and environs (plants, Melbourne) while J. Strachan, C. Stewart, and George Belford (the last still early in a long New Guinea career, although he had already accompanied Armit and the Dentons)—again sponsored by leading newspapers in Australia—made a ‘‘gentlemen’s’’ trip to the Trans-Fly mainland north of Torres Strait (as well as offshore islands including Saibai), returning therein in 1885 and 1886 (plants, Melbourne).

British New Guinea (1884–1898) While Britain in 1883 may have rebuffed annexation, in the following year it could no longer be avoided. At the famous ‘‘intergovernmental conference’’ of 1884–1885 in Berlin, western Melanesia was included along with Africa as part of a world program to delimit metropolitan ‘‘spheres of influence.’’ Developing commercial and, finally, official interest thus propelled Germany into raising their flag in November 1884 over northeastern New Guinea (renamed Kaiser-Wilhelmsland), the newly named Bismarck Archipelago (and Sea), and the western Solomons (with much of Micronesia added in 1885). Partly to placate the Australian colonies, Britain almost simultaneously followed suit, Commodore Erskine on 6 November reading out the proclamation of British New Guinea—initially as a protectorate. The first administrator of the new territory was Sir Peter Scratchley, who arrived in August 1885 but—after an energetic start—served but a few months, passing away near the end of the year. He did, however, lend considerable support (including provision of passage with him to New Guinea) to the first major exploring expedition, that of H. O. Forbes. Forbes and his team were active northeast of Port Moresby from their arrival until May the following year, with an attempt on Mt Victoria in the main Owen Stanley Range their primary goal. Though this was not successful, he spent (with one short break) a full wet season in the foothills between the Sogeri Plateau and the upper Iawarere River, collecting many animals and plants (BMNH, Melbourne, etc.). Twice he was joined by Chalmers and once by Scratchley. Unfortunately due to arguments—particularly at BMNH—many of Forbes’ plants remained unstudied until the early 1920s although the monocotyledons were worked up by H. N. Ridley within a few months of their receipt in London, the results appearing in print in 1886. The 1880s also saw the continued formation of geographical societies—many, if not most, of them also advocates for colonial expansion and development. Some of the branches of the Geographical Society of Australasia were co-supporters

................. 16157$

$CH2

02-08-07 10:22:11

PS

PAGE 27

28 / david g. f r o d i n

(along with the Royal Geographical Society in London) of Forbes, but also in 1885 the Sydney branch organized its own expedition to the Fly and Strickland rivers. Its steam launch Bonito carried 12 scientists led by the zoologist Haacke; W. W. Froggatt collected insects (AM) while W. Ba¨uerlen collected several hundred plants (Melbourne). They were joined for a time by Carl Hartmann, a Queensland horticulturalist. In 1887 the Melbourne branch sponsored W. R. Cuthbertson and W. A. Sayer in their successful ascent of Mt Obree in the Owen Stanley Range— northeast of the Kemp Welch Basin in the present Rigo District—in the Owen Stanley Range (plants, Melbourne). There, Cuthbertson and Sayer overlapped with Hartmann, then on his second expedition—on the invitation of John Douglas, Scratchley’s successor as administrator—which unfortunately led to his death from fever (plants, Melbourne). Hartmann at this time also attempted to establish the first botanical garden in eastern New Guinea (at Konedobu, Port Moresby, later the site of a police barracks and, in time, other government offices). In 1888 British New Guinea became a Crown Colony, and the energetic William Macgregor—previously in service in Fiji—was appointed as administrator (later Lt-Governor). Macgregor vigorously fostered natural history work but with a history of incidents during contacts with villagers made geographical exploration more a government monopoly. Under his direction—and sometimes his own efforts—most of the major mountains in the southeastern peninsula were climbed and as well deep interior penetration made there, up the Fly River, and along the hitherto poorly-known Gulf of Papua and present Oro Province coasts. Apart from Macgregor himself (Mt Victoria, 1889), party leaders included Belford, now in official service (Mt Yule, 1890), R. E. Guise (grandfather of Sir John Guise, first Governor-General of Papua New Guinea; Goropu Mountains including Mt Suckling, 1891), Armit (Mt Dayman, 1894) and A. C. English and Amadeo Giulianetti (Mt Albert Edward and Mt Scratchley, 1896–1897). Animals went along with ethnographic and other official collections to Brisbane (Queensland Museum, where C. W. de Vis studied the birds), while plants went initially to Melbourne (until von Mueller’s death in 1896) and then to Kew (for study by W. B. Hemsley), with some mosses going to V. F. Brotherus in Helsinki as well as (later) to H. N. Dixon (BMNH) and the liverworts to F. Stephani in Geneva. Through these considerable efforts the mountain flora and fauna became much better known. On Mt Victoria it was first demonstrated that a true ‘‘alpine’’ flora existed. Previous efforts in the Vogelkop and the dry parts of the southeast led to hypotheses over Malesian affinities (Beccari) and Australian affinities (von Mueller) in the flora. In his 1889 study of the Mt Victoria flora, Mueller argued for a connection with temperate Australasia for montane plants and both therein and the Northern Hemisphere for high-montane and ‘‘alpine’’ plants, with links to the south better represented. Two years later—partly on the basis of his own extensive investigations in Malesia and New Guinea—the Berlin botanist and orientalist Otto Warburg, then an associate of Adolf Engler, presented a synthesis (further discussed below) for New Guinea phytogeography. But the majority of pronouncements on Papuan biogeography—from the time of Salomon Mu¨ller on-

................. 16157$

$CH2

02-08-07 10:22:12

PS

PAGE 28

Biological Exploration of New Guinea / 29

wards—have been based on analyses of higher vertebrates (especially marsupials). Only in the latter part of the twentieth century did the picture with respect to plants become clearer. Of independent naturalists in British New Guinea in the Macgregor years, Italians as well as British (and Australians) were the most conspicuous. The Italians, encouraged as before by Giacomo Doria (at Genoa), continued the tradition begun by d’Albertis and Beccari. In 1889 Lamberto Loria began the first of two expeditions to the Port Moresby region and other parts of Papua. After d’Albertis, he was the main early entomological collector in the region. Assisted by Giulianetti (see above), he spent long periods in the Sogeri and Meroka regions (to 1,300 m in the latter), the Rigo and Mekeo districts further distant from Port Moresby, as well as the coast of the gulf of Papua and Milne Bay islands (insects, crustaceans, vertebrates, etc., to Genoa; some plants to Florence). During this time, S. Giuseppe separately collected birds in the Gulf of Papua (Genoa). After Loria’s final departure, Giulianetti—as also Armit (in 1894) and, earlier, Belford—became a public servant, working for MacGregor as Government Agent for collection of natural history specimens, including birds (Brisbane) and plants (Kew) and, as already indicated, partly leading a major mountain expedition (during which he found the magnificent (and edible) highland screw-palm, Pandanus jiulianettii). Another officer (and fairly prolific writer), C. A. W. Monckton, collected at Kokoda and elsewhere, and made a remarkable crossing of the Owen Stanley Range into the Lakekamu Basin (BMNH; see also below). From Australia, three further botanists came in the latter years of the Macgregor administration. Two were under official auspices: W. V. Fitzgerald (later active in Western Australia) in 1895, collecting particularly in Oro and Milne Bay Provinces (plants, Melbourne); and in 1898—at the close of Macgregor’s second term—F. M. Bailey, Queensland Government Botanist, undertook a tour with Lord Lamington (the then-Governor of the colony, whose surname today graces a notorious volcano in Oro Province as well as a popular small, chocolate-filled square cake topped with coconut flakes), also collecting and later reporting on his plants (Brisbane). Until 1915 Bailey would act as ‘‘consultant botanist’’ to British New Guinea (and Papua), laying the foundations of one of the key holdings in Australia of New Guinean plants (now fully databased and accessible online). The founder of the Anglican Church in New Guinea, the Rev. Copland King, collected from the time of his arrival, principally ferns (Manila, Sydney); these were described by E. B. Copeland. British non-official plant collectors were during this time represented by the ‘‘plant hunters’’ David Burke (1887–1888, for the Veitch nursery firm in Chelsea, greater London) and Wilhelm Micholitz (1894 and again in 1898, for the Sander firm at St Albans). In the latter year he was at South Cape and in the Milne Bay Islands (finding, among other orchids, Dendrobium atroviolaceum, and also gathering insects while in the Louisiades). However, with greater private support zoological collectors now became more numerous, with Walter Rothschild of Tring—who had begun building up his museum—now emerging as a major spon-

................. 16157$

$CH2

02-08-07 10:22:12

PS

PAGE 29

30 / david g. f r o d i n

sor. In 1888 Basil Thompson collected birds and mammals in the Milne Bay Islands (partly York Museum); in 1890 he was followed by James Cockerell with the Rev. George Brown (from then-German New Guinea; see also below), both engaged in bringing Methodism in the southeastern islands. Another missionary, the Rev. R. J. Andrew, collected insects on Misima, by then a mining site (BMNH). From Germany there came a sponsored collector, Emil Weiske; he obtained Coleoptera, Lepidoptera, and other fauna (including birds and mammals) in 1897–1898 (Dresden and other museums). Localities visited were largely in the present Central Province—much as those of Loria a few years before: the Astrolabe Range, Sogeri, Meroka, Elema, Kabadi, Vaitala, Kerema, Brown River, Kemp Welch River, Paimomo River, Aroa River, Vanapa River, Rigo/Hula, and elsewhere. But by this time German attention was otherwise largely becoming directed towards their possessions (see below).

British New Guinea (1898–1906); Papua (1906–1914) With the departure of Macgregor for a governorship in Lagos, Nigeria, government interest in natural history greatly receded—although his successors, Lt. Gov. G. R. Le Hunte and Administrator F. R. Barton made miscellaneous collections of animals and plants (Brisbane, but also in BMNH). The latter, during his extensive travels, obtained amongst other biota a red-flowered rata vine, Metrosideros ovata (now M. regelii) in the Owen Stanleys and, in 1904–1905 on Woodlark Island, found fossils of dugong, turtle, and gavial. Some collecting was also undertaken by Armit and Giulianetti, the latter accompanying Le Hunte and Barton in the Mekeo region in 1901; but late that year Giulianetti was murdered, Armit having died of blackwater fever some ten months earlier. Of Macgregor’s staff only Monckton continued on (with an ascent in 1906 to the top of Mt Albert Edward); in 1907, however, he resigned—uncomfortable with a ‘‘new era.’’ And Belford’s last assignment was in 1906 with the Royal Commission under Col. J. A. K. Mackay— which included a march right across the Owen Stanleys through Kokoda; but little was (or perhaps could be) collected, apart from a few orchids. Under the new Lieutenant Governor, Hubert Murray, only scant official attention would be paid to the biota. It was bird (and butterfly) collecting that until World War I now took first place in the territory, with Rothschild the dominant sponsor. For a long period, his principal collector was Albert S. Meek. From 1894 to 1916 Meek collected, spending 1895–1901 in the Milne Bay Islands and the eastern part of the mainland. Many assistants worked for him, among them his younger brother (W. G.), Albert and George Eichhorn, brothers-in-law W. B. Barnard and Harry Barnard, and Mr. Gullivers. After Meek’s retirement, the Eichhorns continued work in this region until 1923 in this region (including Owgarra, Bragi, and Mambare River); in 1925, they would collect in New Britain, also obtaining miscellaneous insects. (Most of the bird collections of the Tring Museum were sold in 1932 to AMNH to pay debts, with the museum itself passing into the care of the BMNH in 1937 under Rothschild’s will.) For the British Museum (Natural History), A. S. Anthony was

................. 16157$

$CH2

02-08-07 10:22:12

PS

PAGE 30

Biological Exploration of New Guinea / 31

also active in the region; and very early in the twentieth century (1902–1903) the Pratts—father A. E. and one son, Henry—worked extensively at lower elevations northwest and southeast of Port Moresby and in addition visited parts of the Goilala region, following the gradually developing Sacred Heart Mission interior road (through Mafulu) to the upper Vanapa Valley (at Ononge) before finally returning. They concentrated on birds and Lepidoptera (Tring/BMNH and AMNH); and in 1906 the senior Pratt published a popular book, Two Years Among Cannibals. Botanical collections in this period were rare, partly due to losses in transit, and, with one exception, unofficial. The Pratts in 1902–1903 (see above) had also obtained plants but a goodly part, if not all, were lost. Mary (Mrs. H. P.) Schlencker from Queensland collected extensively from 1905 into the 1920s (but mainly before World War I) in the Rigo district (Boku) and elsewhere while associated with the London Missionary Society (LMS); the majority were described or recorded by Bailey. In 1910–1911 Miles Staniforth C. Smith, director of Mines and Agriculture and long-time Administrator, with a party collected ferns and mosses on his ill-fated trip up the Kikori River to Mt Murray; unfortunately all the specimens were lost on the river descent. Afterwards Smith—perhaps soured by this experience—never again took an interest in biotic exploration, though he did support geological work (on account of mineral and petroleum exploration) and established a small museum in Port Moresby (the forerunner of the present National Museum). He also had come into prolonged conflict with Murray; only in 1918—with Smith away on war service—did the Lieutenant-Governor feel himself able to support a new botanical expedition (see below). By this time the Victorian popular interest in natural history had subsided, as curiosity became satisfied and new leisure pursuits were taken up—although a byproduct was the rise of conservation movements. There were also changes in horticultural fashions, notably the marked decline in the cultivation under glass of tropical plants; while in the sciences, description was giving way to ‘‘higher’’ laboratory-based analysis, along with increasing specialization—a ‘‘linearist’’ development which led to considerable neglect of the Australian biota, let alone that of Papua, for several decades. The first half of the 1890s was moreover for Australia a period of severe and prolonged economic depression; recession also struck in other parts of the world. Official support for such work was thus liable to a lack of funds as well as changing interests and priorities. Only wealthy individuals (or well-founded museums or herbaria) could afford to continue extensive sponsorship and collection formation; and what was obtained tended to be the more spectacular or saleable items. The plume trade, which continued until the end of World War I, also remained an important source of new material. Thus, by 1914 birds and larger insects in southeastern Papua were comparatively well known; but, with far fewer useful points of access, this was simply not true for the interior parts of the Gulf, Delta, and Western Divisions, while botanical knowledge was very patchy and disorganized, and now well behind German New Guinea. Even

................. 16157$

$CH2

02-08-07 10:22:13

PS

PAGE 31

32 / david g. f r o d i n

today, biotic coverage of much of the southern fall of the main ranges remains thin, with more intensive sampling relatively localized.

northeastern new guinea and the bismarck archipelago Before the Germans (to 1885) Large-scale exploration in mainland northeastern New Guinea (from 1884–1914 styled Kaiser-Wilhelmsland) developed later than in British New Guinea or in the Vogelkop Peninsula, but during the decade from 1875 to 1885 the islands to the northeast saw rather more activity. Apart from Miklucho-Maclay (see above), prior to 1885 the sole mainland ‘‘pioneer’’ was F. H. Otto Finsch. This German naturalist, anthropologist, and covert agent undertook two major tours, investigating all the coasts of eastern New Guinea (and the northeastern islands)—and very likely influencing the choice of lands for future German enterprise. In 1880–1882, as part of a world cruise funded by the German Humboldt Foundation, he carried out his coastal surveys, while in 1884–1885—on behalf of the just-formed NeuGuinea Compagnie—with the Samoa he explored in more detail the northeastern and island regions. Finsch made many natural history collections—particularly in 1881 around Astrolabe Bay, already well known from the work of Maclay—and also worked in the Milne Bay region. In 1885 he sailed fifty kilometers up the Sepik, naming the river for Empress Augusta (Kaiserin-Augusta Fluss). Later his interests turned more towards ethnography, already discernable in his book Samoafahrten (1888). His collections (Bremen, Brunswick, Leiden) were partly destroyed in World War II. The tree genus Finschia, a macadamia relative also with edible nuts, is named after him. The northeastern island region was at the opening of the period visited by two great marine expeditions, both in 1875. The first was the Challenger (see above), returning to New Guinea waters after calls elsewhere in Malesia (and ‘‘rest and relaxation’’ in Hong Kong). Under G. Nares as commander, C. W. Thomson as scientific leader, and with Moseley continuing as naturalist, they worked during March in the Admiralty Islands (Nares Harbor in northwestern Manus remains so-named as a token of their presence) before sailing into other waters. The second was the German Gazelle (after which the large northeastern peninsula of New Britain was, and is still, named). With the future Bismarck Archipelago as one of the Gazelle’s main objectives, the vessel and its crew and scientists (with Freiherr G. E. G. von Schleinitz—later the first administrator of the German territories—as commander and F. C. Naumann as surgeon-naturalist), after calling in western New Guinea (see above) anchored during the third quarter of the year at several points: New Hanover, New Ireland (west coast and Carteret Harbor), New Britain (Blanche Bay—then not long known—with an ascent of the volcanic Mt Kombiu or ‘‘The Mother’’), and, in the Solomons, Bougainville (collections, Berlin). In the French ‘‘grand’’ tradition, both expeditions gathered their results into substantial

................. 16157$

$CH2

02-08-07 10:22:13

PS

PAGE 32

Biological Exploration of New Guinea / 33

sets of volumes, those of the Gazelle appearing in five volumes in 1889–1890 although preliminary papers had appeared elsewhere, while the Challenger series was about the most extensive ever published. Both works contain significant materials of primary record for the biota of the region. About the same time as the Gazelle came the first missionaries, traders, and explorers. In mid-August of 1875 George Brown (see also above) arrived at Port Hunter in the Duke of York Islands, there opening the first Methodist mission; he remained for almost a decade. He himself collected some plants, chiefly ferns (Melbourne), but—significantly—was accompanied from Samoa and Fiji (on the mission ship John Wesley) by a German-Australian photographer and naturalist, Carl Walter (who also was collecting for Anatole von Hu¨gel, a wealthy Austrian traveler and naturalist). He worked for about three weeks in the islands and around Blanche Bay (plants, Melbourne and Cambridge, England, albeit with a number of losses in the field); afterwards he returned to and remained in Australia where he continued to collect for von Mueller. In 1877–1878 G. Turner, a collector and gardener from Sydney, traveled to the northeastern archipelago—partly on behalf of the Royal Botanic Gardens, Sydney. From Port Hunter (sometimes with Brown) he visited New Ireland, Spacious (now Wide) Bay (New Britain), and elsewhere. At Spacious Bay he discovered the majestic tree, kamarere (Eucalyptus deglupta)—always a beautiful sight in New Britain’s forests, but a seeming oddity. But this was a time of punitive expeditions, and not a lot apart from more common herbaceous plants could be obtained (Melbourne). In 1881 E. Betche—initially trained as a horticulturalist—traveled from the central Pacific through the northeastern archipelago to Australia. There his base was on Mioko, another island in the Duke of Yorks, where a German trading company had become established in the 1870s. For a week or so in July he collected there and, like his predecessors, around Blanche Bay (plants, Melbourne). Later in the year he joined the Royal Botanic Gardens in Sydney, remaining there for the rest of his career as a collector (and co-author of two standard works on the flora of New South Wales). In 1882 R. Parkinson arrived as a settler, there joining his sister-in-law Emma Forsayth, and at the end of the year establishing the first plantation on Blanche Bay at Ralum (Malapau, near present-day Kokopo, the latter under German rule known as Herbertsho¨he). Over the next quarter-century he traveled extensively and made considerable collections (plants, Melbourne, Sydney and elsewhere); unfortunately, their data are scanty. Parkinson is best known for his book, Dreissig Jahre in der Su¨dsee (1907; English translation, 1999)—a mostly geographical and ethnographic work, but with some remarks on animals, plants, and vegetation (with those generally used by humans in Chapter 11) and a useful survey of exploration in the archipelago (Chapter 12). As already noted, Finsch also visited the archipelago (and also much of Micronesia on his first voyage), but as on the mainland obtained only few natural history collections. (His Micronesian plants were, however, much more extensive; they are now in Co´rdoba, Argentina.)

................. 16157$

$CH2

02-08-07 10:22:14

PS

PAGE 33

34 / david g. f r o d i n

German Rule: The Neu-Guinea-Compagnie (to 1899) The increasing activities by traders, settlers, and recruiters in the northeastern mainland and the archipelago occasioned a fair number of ‘‘incidents’’ which often led to punitive actions. The islands in particular became a center of German activities and in time the German government felt it necessary, in spite of British objections, to proclaim their sphere of interest—whose limits were perhaps at least partly suggested by Finsch—as a territory. This was proclaimed in November 1884 by Capt. Schering, and in the following year von Schleinitz (see above) became administrator (Landeshauptmann), with headquarters at Finschhafen (transferred in 1891 for health reasons to Stephansort (now Bogajim, Astrolabe Bay, PNG), and in 1892 to Friedrich-Wilhelmshafen (now Madang) on the present Schering Peninsula, where there was—and is—a good port). The islands were first administered from Kerawara, then Herbertsho¨he (from 1890, but after 1899 also the capital for all German New Guinea), and—after 1905—Rabaul (which until 1941 remained the German, and afterwards Australian, seat of administration). Under Bismarck’s philosophy of devolved government—a form of rule by ‘‘public-private partnership’’ (PPP)—for the Second Reich’s new territories, German New Guinea (expanded in 1885 to include the Marshall Islands and in 1899 also taking in the Carolines, Palau, and the Marianas) was entrusted to a chartered company, the Neu-Guinea-Compagnie—who also were authorized to undertake commercial operations. Formed in 1884, its founding chairman was Adolph von Hansemann, a Berlin banker who never visited the territory. While until after 1899 not a financial success—the company managing in a relatively short time to use up all their initial capital, and then some—Hansemann at least was interested in exploration and the sciences, and indeed over the fifteen years of their administration much of lasting value was accomplished. One of the first men into the field under the new regime was Carl Hunstein. In 1884 he left British New Guinea with Finsch and until 1888 worked in various parts of northeastern New Guinea, partly with R. Mentzel (and others) on geographical and forest reconnaissance work and partly on the Neu-GuineaCompagnie exploring and scientific expedition of 1886–1887 (see below). Early in 1888, while with an official and writer, Stephan von Kotze, in the outliers of the Rawlinson Range behind Finschhafen, he discovered the beautiful (and endemic) southern pine, Araucaria hunsteinii; but not long afterwards—on 13 March—he was drowned by a tidal wave while in the Kilenge district at the western end of New Britain. As before, he specialized in birds, many now bought by Finsch. With Mentzel, Hunstein in April 1886 visited the mouth of the KaiserinAugusta (now Sepik) River. They thus blazed the way for the major Neu-GuineaCompagnie expedition of 1886–1887, the first of two significant, relatively successful forays into the interior under their auspices—although most collections were of lowland animals, insects, and plants. These apart, with a predisposition to caution—perhaps from experiences with local populations in the Bismarck Archipelago—interior penetration by individuals was limited. Until the mid-1890s the only inland mountain areas reached were Sattelberg in the Huon (Kai) Peninsula and

................. 16157$

$CH2

02-08-07 10:22:14

PS

PAGE 34

Biological Exploration of New Guinea / 35

the western Finisterres south of Astrolabe Bay, both by Neu-Guinea-Compagnie official F. C. Hellwig (see below). The 1886–1887 expedition—lasting for nearly a year and half from 19 April 1886—was led by von Schleinitz, partly on the Neu-Guinea-Compagnie steamer Ottilie, and accompanied by C. Schrader (geography and bryophytes, as well as scientific coordinator), U. M. Hollrung (vascular plants), and Hunstein with F. Grabowsky (birds and insects). Over the course of the expedition they worked around and inland of Finschhafen and visited Samoahafen (Salamaua, in the Huon Gulf), Astrolabe Bay around Constantinhafen (Melamu), the coast north and northwest of Friederich-Wilhelmshafen (now Madang), including Hatzfeldthafen, and made two forays up the Sepik: a shorter one in 1886 followed the next year by a deep plunge—the first great voyage thereupon, with camps at I. AugustaStation (Zenap near the mouth of the May (Iwa) River far in the interior) and II. Augusta-Station (Malu near present-day Ambunti). The large haul of collections went to Berlin (plants partly destroyed, but duplicates at Melbourne, Kew, and elsewhere); the botanical results appeared in 1889 in a special supplement to the Neu-Guinea-Compagnie Nachrichten, or series of official reports. Hansemann’s interest in natural history ensured that a fair number of individuals—some employees of the Neu-Guinea-Compagnie—took up collecting. Among the first was the Pole J. S. Kubary, who in 1887–1891 from a Neu-GuineaCompagnie base at Constantinhafen—the first official post in that area—collected for the Godefroy Museum in Hamburg (partly now in the Senckenberg Museum, Frankfurt; some plants went to Berlin). Kubary also made land acquisitions around Astrolabe Bay which later led to much local strife, with no ‘‘settlement’’ until 1932, and a bone of contention even long afterwards. In 1892–1895 Kubary returned to New Guinea for a second contract with the Neu-Guinea-Compagnie. In 1888 R. Rohde, another Neu-Guinea-Compagnie employee, collected animals and insects at Finschhafen and Stephansort (the latter a plantation first established in August 1888 on a portion of the Kubary lands particularly for raising fine ‘‘Deli’’ (Sumatran) tobacco; planting began in 1889). A change in Neu-GuineaCompagnie external shipping routes from Cooktown to Surabaya (in Java)—then followed, as well as the transfer of government headquarters from Finschhafen; henceforth Astrolabe Bay became a center for Neu-Guinea-Compagnie activities on the mainland. From 1886 the pioneer Lutheran missionary Johannes Flierl collected animals on the Huon Peninsula, beginning at Simbang near Finschhafen; in this he was followed by his son and grandson. In botany Flierl was complemented by another Lutheran priest, G. Bamler who collected in the Peninsula and particularly on the Tami Islands (collections to Berlin). In 1891 they were followed by H. Fruhstorfer, who collected Lepidoptera at Finschhafen and elsewhere. Non-missionary botanists were also soon active. Apart from Hollrung (see above), important collections were made by Hellwig (1888–1889) and C. Weinland (1889–1891), the latter also a Finschhafen-based Neu-Guinea-Compagnie official (and Hellwig’s professional successor), and, over six weeks in 1889, by

................. 16157$

$CH2

02-08-07 10:22:14

PS

PAGE 35

36 / david g. f r o d i n

Warburg (see above) (Berlin; now largely destroyed, but duplicates in Wroclaw (Breslau), Kew and elsewhere). Hellwig—with whom Warburg made some trips— was the first to reach mountain localities. The first foray was to Sattelberg west of Finschhafen (ca 900 m, where ‘‘oaks’’ (Lithocarpus) were found for the first time and soon afterwards a mission ‘‘hill station’’ would be established). The second foray—accompanied by another metropolitan visitor (a well-known writer and newspaperman from Cologne, Hugo Zo¨ller)—was to the western Finisterres, from where the Bismarck range and its high peaks, Mts Otto, Wilhelm, Herbert, and Marien were all seen (and named) and the first rhododendrons collected (all in sect. Vireya). One of the more stunning (and relatively widespread) of these— which may reach as low as 100 m—remains known as R. zoelleri. As we have seen, the Neu-Guinea-Compagnie was strongly active in agricultural development, initially in the hope of attracting German smallholders, later turning to plantations. Over 1887–1891 and 1893–1894 (and at other times on his own account) its horticulturalist, L. Ka¨rnbach, a pupil of the Berlin Botanical Garden, traveled widely in the territory. In addition to much information on useful plants, gathered together particularly in a paper of 1893 and (after his loss at sea in 1897) in M. Krieger’s 1899 book Neu-Guinea, Ka¨rnbach also obtained many marine algae—beginning the investigation of these plants since ably carried on from time to time (a first consolidation of knowledge was made in 1900 in Lauterbach’s Flora; see below). In 1890–1891 C. A. Lauterbach, a botanist-geographer of private means with an estate now within the western part of the present Polish city of Wroclaw, made the first of three trips to New Guinea. In addition to the Gazelle Peninsula, he collected along the Huon Gulf from Finschhafen toward the Bukaua district and ‘‘Burgberg’’ (Lo Wamung at Lae), on the Sattelberg, and around Astrolabe Bay and was the first to penetrate the Gogol River south of Friederich-Wilhelmshafen (now Madang). He was to return to New Guinea for some time in 1896 and again in 1899–1900 (see below). Until the end of his life in 1937 he remained strongly interested in New Guinea, both in a business role (he was an early director of the Neu-Guinea Compagnie as a purely commercial concern after mid-1899) and a personal role (supporting the publication of the later botanical results of New Guinea explorations and himself preparing accounts of many plant families). Lauterbach’s collections ranged across the whole plant kingdom (private herbarium now in Wroclaw, a fortunate survivor of major upheavals there in 1944– 1945; duplicates in Berlin, Kew, and elsewhere). This is reflected in his important 1900 work (with the Berlin botanist K. Schumann), Die Flora der deutschen Schutzgebiete in der Su¨dsee; a supplement (Nachtra¨ge) followed in 1905. Long standard, for non-primary forest plants in the lowlands in particular, it provides fairly considerable coverage. In later years, in addition to his accounts of plant families Lauterbach was to add considerably to knowledge of the phytogeography and vegetation of New Guinea, including in 1928–1930 ‘‘cross-sections’’ of different formations in which he drew upon his personal knowledge as well as the results

................. 16157$

$CH2

02-08-07 10:22:15

PS

PAGE 36

Biological Exploration of New Guinea / 37

of others. He also contributed to general works on New Guinea as well as German territories as a whole. Warburg also summed up his experiences (and plant records), notably in his already-mentioned paper of 1891. Here, he argued that while there were several distinct elements in the flora of New Guinea, its predominant relationship was with western Malesia. Yet, for him no ‘‘line’’ in the island region west of New Guinea could be recognized as a major discontinuity comparable with the several for faunal (mainly higher vertebrate) discontinuity proposed, among others, by Wallace, Weber, and Lydekker. The Berlin botanist L. Diels was, however, later to propose a major floral line at Torres Strait—subsequently adopted by van Steenis as a demarcation for Flora Malesiana. Evidence aired at a symposium in Australia in 1972 and since has now somewhat weakened this argument. Warburg also proposed the term ‘‘Papuasia’’ to refer to the whole region from New Guinea to the Solomon Islands, an area known to zoogeographers as the ‘‘Papuan Subregion’’ (though Mayr’s ‘‘Papuan Region’’ of 1941 only includes New Guinea and its more or less fringing islands, excluding the Bismarcks and Solomons). The aforementioned changes to shipping and transfer of activities to Astrolabe Bay—where under a subsidiary, the Astrolabe-Compagnie, tobacco cultivation was during the 1890s vigorously expanded (though at some human cost)—brought in many collectors who were also active elsewhere in Malesia, among them a number of non-Germans. Scheduled shipping services, along with hotels and guesthouses, were now making it easier for visitors to travel (and tour) around the ports. Among the first was Bernard Hagen, engaged from Sumatra by the AstrolabeCompagnie as an expert in ‘‘Deli’’ tobacco. From 1894–1895, in addition to his duties, he undertook ethnographic studies and collected mainly insects and birds (Berlin). In addition to Astrolabe Bay localities such as Stephansort and Erima, he also collected at Berlinhafen (now Aitape)—then a ‘‘new’’ locality (but where, on the offshore island of Seleo, Ka¨rnbach (see above) had established for himself a coconut plantation). Departing for health reasons, Hagen later worked up his observations into a popular book, Unter den Papuas (1899). Hagen was followed in 1895 by one of the early dilettante-travelers, O. Ehlers, who came with an idea to cross the mainland cordillera from north to south—something not previously attempted. A police-master, W. Piering, was grudgingly seconded by the administration; but, starting from near Samoahafen, some weeks into their trip (and out of food) both were lost with collections and notes in a river. Only their two local assistants got through to the Lakekamu River (in British New Guinea). From then on, the authorities—faced with much strife elsewhere (ultimately one of the factors leading to the withdrawal of the Neu-Guinea-Compagnie from government)— were, as already indicated, more cautious with regard to the mainland interior, with the result that its penetration was relatively slow and only for experienced persons with strong support. British-supported collectors also began to make their appearance, reflecting wealthy backing. In 1890 Carl Wahnes began his long association with New Guinea (until 1909) as a collector for the British Museum, Tring, and other insti-

................. 16157$

$CH2

02-08-07 10:22:15

PS

PAGE 37

38 / david g. f r o d i n

tutions (e.g., Odonata at the University of Michigan). During several sojourns he collected insects, birds, and other animals on the Huon Peninsula (Finschhafen, Simbang, Sattelberg, Rawlinson Range), at Bongu on Astrolabe Bay (MikluchoMaclay’s old base), and on some offshore islands (Tarawai, Isle Deslacs) and in New Britain. Wahnes’s collections led to numerous publications, including some by Rothschild and the other Tring zoologists, C. Jordan and E. Hartert. In 1893– 1894 the ‘‘eccentric’’ English sea captains Cotton and H. Cayley Webster (the latter returning in 1897)—with recommendations from Hansemann as well as Rothschild—collected birds and butterflies (Tring/BMNH). Some specialist papers resulted and Cayley Webster soon wrote a popular work, Through New Guinea and the Cannibal Countries (1898)—among the earliest in English in an extended tradition of such books. Austria-Hungary was once more represented, mainly in the long sojourns of the Hungarians Samuel Fenichel (1893–1895) and Lajos Biro´ (1895–1901). The two worked extensively around Astrolabe Bay, at Berlinhafen and its offshore islands, and in the eastern Huon Peninsula (Simbang, Bingala, Sattelberg) as well as on the Gazelle Peninsula (see also below)—though mostly at lower elevations. Their primary emphasis was on insects but many other animals were collected (Budapest, but birds, reptiles, amphibians, mollusks and Diptera—excluding Pupipara—were destroyed during the 1956 uprisings; parts of Fenichel’s collections are now at the College of Nagyenyed in Transylvania, in modern Romania). Plants were also collected (Berlin and elsewhere). Biro´ also obtained many lichens (or lichenized fungi), which were included by Schumann and Lauterbach in their 1900 Flora and much later (1956) by O. Szatala in a first checklist of all those in New Guinea. Biro´ was the first real scientist-zoologist to spend much time in New Guinea. Later he became something of a ‘‘hero’’ in Hungary, writing two books of travels popular with young people (Szent-Ivany, personal communication to J. L. Gressitt). Sadly, Fenichel died at age twenty-five in Stephansort of cerebral malaria, for which Stephansort had acquired a bad reputation (and would later be visited by Koch; see below). There is a memorial to both Biro´ and Fenichel at the University of Papua New Guinea. The early 1890s were, as we have seen, a difficult time for the Neu-GuineaCompagnie. Circumstances improved after 1895, however, and official thoughts once more turned to interior exploration—now with the prospect that minerals (particularly gold) might be found. The next big foray was the 1896 Gogol-Ramu Expedition, organized by the Neu-Guinea Compagnie with some support from the Reich government and frugally led by Lauterbach with O. Kersting as policemaster and E. Tappenbeck as quartermaster. After a trip to Mt Oertzen (between the Gogol and Stephansort), their party passed up through the Naru Valley running southwest from the lower Gogol River, traversed (via the Ssigauu uplands) the dividing range (west of the modern road), and entered the then-unknown Ramu (now Jagei) Valley (northwest of present-day Usino). From there they traveled by canoe to near what is now Annaberg (downstream from Aiome), returning

................. 16157$

$CH2

02-08-07 10:22:15

PS

PAGE 38

Biological Exploration of New Guinea / 39

by the same route. Side trips were taken, including the relatively dry Bismarck Range foothills. The expedition overall was a success, with good returns in geographical knowledge as well as collections of plants, birds, and animals (Berlin, Wroclaw, and elsewhere). The plants would soon be incorporated into his Flora (see above). The Ramu expeditions were continued for the Neu-Guinea Compagnie by Tappenbeck with two associates, H. Rodatz and H. Klink. In 1898 the inland Ramu River was linked to its mouth (Ottilie-Fluss), while for two months during ‘‘winter’’ low water Rodatz and Klink lived in a temporary camp on the lower Ramu River. In 1899—after the change of administration—traces of gold on the upper Ramu River (near Usino) attracted the Neu-Guinea Compagnie’s interest. A sternwheel riverboat was obtained and further trips made. During the dry season, Rodatz and Klink worked from a base at Arumene (near Aiome), reaching the foothills of present-day Mt Aiome in the western Bismarcks, and in November 1899 Lauterbach returned for a month’s trip along the river, leaving early in 1900 (plants, Berlin and Wroclaw). Rodatz and Klink remained in the area at a base camp near Usino, later visited by Schlechter (late 1901), and subsequently they gave many years of service as district officers, Klink eventually at Morobe, from which he explored the mid-Waria Valley (around present-day Garaina). There he found the stands of Araucaria for which he is remembered (at first described as a distinct species, A. klinkii, but later—not entirely critically—united with A. hunsteinii ). The presence of Parkinson in the Bismarck Archipelago helped to attract additional scientists to that region. One who remained for almost a year (1896–1897) on Blanche Bay was F. O. Dahl, who with the assistance of Parkinson and the now-wealthy Emma Forsayth (now Kolbe) set up a small ‘‘station’’ at Ralum and collected extensively around the area (including the Duke of York Islands and the Baining Mountains, the latter from a new mission station, Vunamarita) both on land and sea (Berlin). Dahl’s stay would be very productive, with many published results. These included a regional florula (1898) by K. Schumann and a stream of zoological papers. Dahl’s marine collections, along with those of his contemporary, A. Willey from Cambridge, England, marked the beginning of serious study of the rich undersea biota in the Bismarck Archipelago (apart from the relatively short visit of the Challenger). Dahl was later a curator at the Museum fu¨r Naturkunde, Berlin. Willey enjoyed two long sojourns in eastern New Guinea, the first in 1895 in the Archipelago before proceeding to British New Guinea (and elsewhere), returning there in 1897 before sailing back to Europe. Willey also published extensively (sometimes with others)—notably a six-part collection entitled Zoological results . . . collected during the years 1895, 1896, 1897 (1898–1902). The last noteworthy visitor under the Neu-Guinea Compagnie—arriving just before the transfer of administration—was the Swede E. O. A. Nyman. From field studies in Java he came in 1899 to New Guinea for nearly nine months (March– November). Often ill, he collected birds, plants, and lichens around Astrolabe Bay (including the low Hansemann massif near Friedrich-Wilhelmshafen), Finschha-

................. 16157$

$CH2

02-08-07 10:22:16

PS

PAGE 39

40 / david g. f r o d i n

fen, the Sattelberg area (partly for health reasons), and northeastern New Britain (plants and lichens in Uppsala). Sadly, Nyman was never able to work up his collections, dying in Munich in 1900 before reaching Sweden. Some of Nyman’s collections were accounted for in Schumann and Lauterbach’s Flora, while at Uppsala in the 1940s R. Santesson described his epiphyllic lichens; but many others have never been documented, including some higher plants seen by the writer in 2003.

German Imperial Rule: The Era of Benningsen and Hahl (1899–1914) The developments in the Ramu Valley (now Jagei Valley)—relating in part to mineral exploration—were strongly promoted by R. von Benningsen, first Imperial Governor of German New Guinea. While scientific results under Neu-Guinea Compagnie rule had been impressive, economic results were not so, and most of its capital had been expended. After long negotiations, the Neu-Guinea Compagnie was relieved of its administrative responsibilities as of 1 April 1899 and von Benningsen assumed control. The new Governor himself was an amateur entomologist of note, specializing in Coleoptera (beetles). Von Benningsen collected as opportunities permitted in New Britain, the Duke of York Islands (called Neu-Lauenberg at this time), and elsewhere in the Bismarck Archipelago as well as in the mountains behind Stephansort especially the Kani Range and upper Minjim River (collections mostly in Dresden). As Kotze (see above) wrote in his popular (and amusing) book about his time in New Guinea, Aus Papuas Kulturmorgen (1905), Coleoptera were popular with Germans as Lepidoptera were for the British. In this respect Benningsen set a good example; many other amateurs also collected beetles which likewise found their way to the Royal Dresden Museum and its noted coleopterist, K. M. Helle. Public health (and, in some places, depopulation) were not unnaturally serious concerns of the new government, and with the 1898 discovery of the mechanism of malaria transmission new possibilities for its control opened up. In 1899–1900 the famous microbiologist Robert Koch came from Java to the territory for seven months (from December 1899). Traveling partly with Benningsen and Biro´, he visited Berlinhafen, Astrolabe Bay and its villages and settlements (including the plantations at Stephansort, now Bogajim), Finschhafen, Huon Gulf (including the Tami Islands), the Siassi Islands, parts of northwestern New Britain and the Witu Islands, the mouth of the Ramu, and finally the Gazelle Peninsula and elsewhere in the Bismarck Archipelago including the islands northeast of New Ireland. Along with his malariological and parasitological work he managed to collect some animals and plants. Koch’s studies were followed up by Dr O. Dempwolff (with a trader, F. E. Hellwig); during 1902–1904 (Hellwig on his own in 1903) they spent considerable time in Wuvulu, Aua, and other western Admiralty Islands—places not visited by the microbiologist. One result was the only monograph on the first two of these islands (notable for their Micronesian people), Wuvulu und Aua (1908) by

................. 16157$

$CH2

02-08-07 10:22:16

PS

PAGE 40

Biological Exploration of New Guinea / 41

P. Hambruch in Hamburg (home to the former Godeffroy Museum, in the nineteenth century an active patron of Pacific biological and cultural exploration). In it is a clear demonstration of the contemporary rise of professional and popular interest in the region’s ethnography, already the subject of a major 1898 expedition from Cambridge University to the Torres Strait and also prominent in Richard Parkinson’s Dreissig Jahre of the previous year. At this time a considerable, sometimes detrimental trade in artifacts developed along with the rising traffic in bird plumes, while at the same time fewer large private collections of natural history objects were being formed. Among the last private collectors to visit German New Guinea was B. Mencke, who with the yacht Eberhard—purchased in Monaco—and two zoologists, G. Duncker and O. Heinroth, collected in 1900– 1901 in various coastal localities including the Arawe Islands (southern New Britain). Mencke’s expedition, however, came to grief in the Mussau group; Mencke was seriously wounded in a fight (dying 2 April 1901) and Heinroth, after some further work in New Ireland, departed in June. Tangible results were but few. Benningsen—who had departed the territory in mid-1901 for leave—was in 1902 succeeded as Governor by Albert Hahl. Familiar with the Bismarck Archipelago from prior service, he concentrated more on developments (and pacification, following some notorious incidents) there. However, as circumstances permitted (particularly after the mid-1900s) he also vigorously promoted mainland exploration and scientific work, much as had Macgregor in British New Guinea (although less so in person). Like his predecessor, Hahl could also rely on goodly numbers of officers and others interested in natural history to make contributions, though the important expeditions, both on the mainland and in the Bismarcks, usually involved visiting, usually professional scientists. Shortly before Hahl’s appointment (but while he was Vice-Governor), the territory was visited by its next major metropolitan visitor, F. R. R. (Rudolf) Schlechter. He was sent out by the Kolonial-Wirtschaftliche Komitee in Berlin to find latex-producing trees and vines useful for gutta-percha and rubber—Germany wishing to develop its ‘‘own’’ sources of these substances. The first of his two expeditions (1901–1902) was exploratory, the second (1906–1910) developmental. On his first tour (in the territory lasting ten months from mid-October 1901) he visited the Bismarck Archipelago (including southern New Ireland) and, on the mainland, Berlinhafen (and behind it, the Torricelli Range) as well as its offshore islands, and also crossed the Gogol-Ramu dividing range (along Benningsen’s new ‘‘road’’) to the Ramu gold camp, from there proceeding into the Bismarck Range as far as the present Bundi district. Schlechter’s recommendation to the administration for a botanical garden was followed up by Hahl in 1906 upon the establishment of Rabaul; in time this became a ‘‘beauty spot’’ (though not really a center for botanical exploration), lasting until 1942. In 1907–1909 the now-noted scientist (who had at other times also collected in southern and central Africa) returned to New Guinea. For this more extended stay he set up a base camp on Astrolabe Bay at Bulu (near Bongu). From here Schlechter did intensive fieldwork on the inland coastal/Ramu divide

................. 16157$

$CH2

02-08-07 10:22:17

PS

PAGE 41

42 / david g. f r o d i n

(especially in the steep Kani and Ibo ranges) and in the western Finisterres, and also made long trips to the Bismarck Range (with Hahl for an attempt on Mt Saugueti (also known as Mt Otto) in 1908)—but just missing discovery of the central highlands. Schlechter also visited the lower Waria in 1908 and 1909 (in conjunction with a border survey), and (with O. Schlaginhaufen and Hahl) the Torricellis in August and September 1909, where Schlaginhaufen collected insects and other animals at Paup (east of Aitape) and elsewhere (Dresden). In addition to his gutta-percha and rubber plant discoveries (thought moderately successful at the time but in the end without lasting economic impact, there being no market after 1914), on his two expeditions Schlechter made large collections of plants including very many orchids (Berlin; partly destroyed, with some duplicates elsewhere) and some gatherings of animals. In this he was aided by two men from New Ireland, Sikin and Takadu. Unfortunately some of Schlechter’s main work areas have never again been re-studied, particularly with the destruction in 1943 of so many of his primary collections (see below). A useful general account of his second expedition, including maps, is Die Guttapercha- und Kautschuk-Expedition nach Kaiser-Wilhelmsland, 1907–09 (1911). Soon after Schlechter’s first visit another leading botanist, the bryologist (and expert photographer) M. Fleischer, came to the territory. In March 1903, after some three years in Java, Fleischer visited the mainland (including Astrolabe Bay and Finschhafen) and the Bismarck Archipelago (including Mioko Island). His collections included orchids, fungi, and pteridophytes, in addition to mosses and liverworts (Berlin and elsewhere, some by sale; his private herbarium is now in the Farlow Herbarium of Harvard University Herbaria). A first synthesis of his and other New Guinea mosses, by W. Schultze-Motel, appeared in 1963. With gradually improving economic conditions during the latter period of Hahl’s rule—part of the ‘‘long boom’’ of the decade or so before World War I—and more opportunities (including further transport developments as well as more government posts) came other collectors. In 1906 H. Schoede collected animals at Berlinhafen and Simpsonhafen. Eugen Werner in 1907 collected insects and plants south of Astrolabe Bay, partly with Schlechter (plants, Wroclaw). Werner also wrote a popular illustrated book, Kaiser-Wilhelmsland (1911). In 1909 E. F. Dahl, Preuss, O. Heinroth (now on his second visit), and C. Ribbe collected in northeastern New Britain (Ribbe also reaching the Duke of York Islands); and P. Nagel visited the Finisterres. In 1908–1910 Prof. R. Neuhauss carried out many months of field research for his three-volume Deutsch Neu-Guinea (1911), much of it in the Huon (Kai) Peninsula but also at many other places, both coastal and inland (not, however, in the Bismarck Archipelago—then under active survey by other expeditions). Dallmannhafen (Wewak) was visited with J. Walis. Insects, animals, and a few plants were collected (Berlin) along with much ethnographic material; many photographs were also made, and his book is thus richly illustrated. While in the Huon Peninsula, Neuhauss traveled with the great Lutheran missionary and traveler C. Keysser. In 1909 Keysser established a station at the head of Huon Gulf close to

................. 16157$

$CH2

02-08-07 10:22:17

PS

PAGE 42

Biological Exploration of New Guinea / 43

‘‘Burgberg’’ and not far from where Lae would develop in the 1920s. Later (1911– 1912) Keysser reached the top of the Saruwaged (Salawaket) Range, Mt Bangeta (‘‘Bolan’’) at nearly 4,000 m, and also traveled through the Bulolo and Wau valleys. In 1916 Keysser was to repeat his Saruwaged trip with H. Detzner during Detzner’s fantastic four-year journey (1914–1918) ‘‘on the run’’ from the Australians. The plants found (Berlin) were ultimately summarized by Diels (1929). After his return to Germany Detzner would, however, become notorious for his semifictitious Vier Jahre unter Kannibalen (1921). Other collectors of this period—some of them with the Lutheran Church— included F. Kunzmann (fish, reptiles, and butterflies), and Lothar von Wiedenfeld in 1909–1910 (birds and insects at Berlinhafen, Sattelberg, and adjacent coast at Simbang and Heldsbach). In 1909 W. Mu¨ller collected insects (Dresden) and L. Maschmeyer animals; around 1913 K. Ma¨ilander collected plants near Morobe (Berlin). In 1913 Ma¨ilander also traveled through the upper Waria and the Wau and upper Watut valleys. P. Nagel collected insects at Komba in the Finisterre Mountains. Other missionaries making collections included Hoffmann, Bergmann (who was active into the inter-war years, and host for a time to M. Clemens; see below), Kunze, and Vetter. Also in the latter part of 1913, K. Gehrmann, botanistin-charge at the Rabaul garden (see above) and veterinary officer M. Braun undertook an extensive study tour in the Gogol-Ramu area. From this, two substantial general reports emerged but few collections (Berlin) are known. The years from 1907 to 1914 were also marked by a further wave of large expeditions, both on land and sea. These included major undertakings such as those of the Planet, Peiho, and Stolle´/Behrmann (see below) as well as two mainland border surveys. The first was the British-German border survey of 1909, for which Fo¨rster led the German team; it was accompanied by Schlechter (see above) on the lower Waria as far as the dividing Maboro ridge (where he found a new slipper orchid species, Paphiopedilum violascens). The second was the German-Dutch border survey of 1909–1910, for which the German team was led by the geographer L. Schultze-Jena. The latter party explored near the border (141 E) as far as the Bewani Range and, afterwards, right up the Sepik River as far as the Peripatus Range (possibly what incorporates Sepik Mt, 1,570 m) in the foothills of the present Star Mountains. They secured extensive geographical results but only scattered animal and plant collections (Berlin; plants largely destroyed). Peripatus has not been biologically surveyed since then. Of the other expeditions, both the first and second focused on the Bismarck Archipelago. The expedition of the Planet, of 1906–1909, was organized by the Reichs-Marineamt, and participating scientists included E. Stephan (as leader, who died during the latter part of the expedition), E. Graeffe, Schlaginhaufen (see above), K. T. Sapper, and A. Kra¨mer. The first stage (1906–1907) took them mainly to the Admiralty Islands (including its western chain), while in the second stage (1907–1909) they focused particularly on New Ireland (and its northeastern islands), Kra¨mer in April 1909 making a first ascent of the Lelet Plateau (visited in 1973 by the writer). Many publications resulted, beginning with Forschungsreise

................. 16157$

$CH2

02-08-07 10:22:17

PS

PAGE 43

44 / david g. f r o d i n

SMS ‘‘Planet’’ 1906/07 (5 vols., 1909) but also encompassing a major survey of New Ireland by Sapper (1910)—a forerunner of the many similar studies sponsored from Australia in the 1950s and 1960s (see below). Considerable collections were made, largely zoological but also algae (by Graeffe, now in Hamburg; they would go towards a regional revision (1928, by O. C. Schmidt) of marine species) as well as some other plants (Kra¨mer; to Berlin), a number of them from the Hermit Islands. After the close of operations in June 1909 Schlaginhaufen proceeded to Berlinhafen and the Torricelli Range. A popular account by H. Vogel appeared in 1911 as Eine Forschungsreise im Bismarck-Archipel. The Planet was followed into New Guinea waters by the Hamburg Academy of Sciences-sponsored ‘‘Su¨dsee-Expedition’’ (1908–1910) with the Peiho (commanded by Capt. Vahsel). On the vessel were a number of scientists led by Dr Fu¨lleborn (commemorated in a harbor on the south coast of New Britain) and including Duncker (see above under Mencke) as zoologist (specializing in fishes) as well as F. E. Hellwig (see above) as ‘‘liaison officer.’’ The expedition visited several localities in the Bismarck Archipelago, particularly along the south coast of New Britain (1908–1909); they then sailed for the mainland and after a call at Langemak Bay (Simbang, a mission station near the old site of Finschhafen) they proceeded in mid-1909 for a voyage up the Sepik. After departure from FriedrichWilhelmshafen (now Madang) they sailed to Micronesia, where the scientists were led by G. Thilenius. Duncker’s zoological collections, as well as a few plants, are in Hamburg. Thilenius edited the series of expedition results, but their overwhelming emphasis is on ethnography; only one ‘‘general’’ study was published including the expedition itinerary (Allgemeines, 1927). Any biological results (e.g., on birds, by G. H. Martens) appeared elsewhere. These not insubstantial undertakings were, however, soon outdone by the great ‘‘Kaiserin-Augustafluss Expedition’’ of 1912–1913—the largest and longest of all those mounted under German rule and, in retrospect, a fitting climax to its thirtyyear run. The expedition was sponsored by the Geographical Society in Berlin and other German organizations and led by Bergassessor Capt. A. Stolle´, who already had some field experience in New Guinea. The six-man scientific team was headed by the geographer Dr W. Behrmann. From March 1912 to September 1913, by ship and launch and on foot, they surveyed most of the Sepik basin including the tributary rivers (May or Iwa, Freida, Leonhard-Schultze, Do¨rfer, To¨pfer or Keram, etc.) as well as a number of outliers and foothills of the central ranges (but not reaching above 2,200 m). Their ‘‘Hauptlager Malu’’—a base camp just upriver from the village of Malu (as already noted, the site in 1887 of a Neu-Guinea Compagnie expedition camp)—is today close by (and effectively part of) the East Sepik town of Ambunti (the name being a local rendering of the site of ‘‘Hauptlager Malu’’ as heard by G. W. L. Townsend, an Australian official who in the 1920s established the station (District Officer, 1968)). Other localities were Seerosensee (Chambri Lakes), Peilungsberg, Zuckerhut, Pionierlager, Ma¨anderberg, Hu¨gellager am Sepik, Sepikberg (1,570 m, see above)—all on or near the Sepik; Hunsteinberg (or Sumset; 1,350 m), Etappenberg (850 m), Lordberg (or Durch-

................. 16157$

$CH2

02-08-07 10:22:18

PS

PAGE 44

Biological Exploration of New Guinea / 45

blick, 1,000 m) and into the Zentralkette (up to 1,800 m) as well as Lager 18, all in the April River drainage; Pfingstberg on the May (Iwa) River; Felsspitze in the Westkette (1,400–1,500 m) west of the May; and, to the southeast (via the To¨pfer) to the Lehm and up to Regensberg (550 m) and Schraderberg (2,070 m). One of the ethnologists, R. Thurnwald, would in 1913 reach the headwaters of the Sepik, only turning back near the present site of Telefomin. Animals were largely collected by W. Behrmann and J. Bu¨rgers, while C. Ledermann secured all the plants (6,600 numbers). Collections went to Berlin and Dresden (animals) and BerlinDahlem (plants). Most animals survived World War II but the plants suffered severe losses; only some duplicates have survived (mainly Leiden, Edinburgh, Singapore, and Wroclaw) and little ‘‘topotype’’ collecting has been done (save on Mt Hunstein, now considered a site of exceptional biological interest). As well as geomorphologic work, Behrmann made excellent maps which served for half a century. But it came to be realized that further effective progress— particularly in the central ranges, still thought to be unpopulated—would have to be aided by some form of aerial support. Airships (Zeppelins) were first proposed (and advocated notably by Neuhauss, who was not confident about using airplanes given the difficult terrain and near absence of suitable landing places at that time); in 1913 ‘‘stamps’’ were even printed. But the use of aircraft belongs to the next era (between World War I and World War II), starting in 1926. Until then, movement continued by water and foot, sometimes (particularly in western New Guinea) with scores if not hundreds of men—all requiring life’s necessities. Publication of the results of the Behrmann expedition was deemed to merit special consideration, although much would appear in existing journals rather than in a special series (some contributions did, however, go into the Dutch series, Nova Guinea). Support came mainly from the Hermann und Eliza HeckmannWentzel-Stiftung. Botanical results—edited by C. Lauterbach (later by L. Diels)— appeared in Botanische Jahrbu¨cher, a leading systematics journal, mostly in a specially titled series Beitra¨ge zur Flora Papuasiens (1912–1942). This remains among the best sets of consolidated work on the plant taxonomy of New Guinea. Zoological results appeared in the bulletin of the Berlin Zoology Museum (Mitt. Zool. Mus. Berlin), the birds in 1923 by Erwin Stresemann, a curator in that museum (and professor in ornithology). In 1922 Behrmann brought out a readable popular account, Im Stromgebiet des Sepik. In summary, while its beginnings were slow, natural history investigation in northeastern New Guinea and the Bismarck Archipelago under German rule rapidly gathered speed and by 1914 had produced a great mass of material and information, even partial digestion of which would take years. The ‘‘data base’’ had now pulled well ahead of Papua, where after 1898 collecting—except in some taxonomic groups—had slackened considerably and would fall further for a time after the advent of Australian rule (see above). Even today—thanks to its ‘‘Mitteleuropean’’ experience—with respect to mainland New Guinea as a whole former Kaiser-Wilhelmsland retains its lead in collections. But while World War I caused some disruption, a rather more severe and lasting blow would be delivered by

................. 16157$

$CH2

02-08-07 10:22:18

PS

PAGE 45

46 / david g. f r o d i n

World War II through significant collection losses in central Europe and elsewhere. Though widely recognized, the scale of these losses and the obstacles they posed for future research and reference, were in the decades after World War II never sufficiently perceived by some in Papua and New Guinea and followed up with appropriate action.

netherlands new guinea In contrast with the eastern territories, serious official interest in western New Guinea—particularly the interior—remained relatively limited until outside pressure forced the authorities into greater action. The late nineteenth century was thus dominated by private (including commercial) activity, with a strong emphasis on the saleable. After 1898, activities were largely dominated by the official and semi-official sector, the authorities also concerned to avoid ‘‘incidents’’; but government sponsorship at least partly ensured the collection of a better representation of the biota, though by 1915 still below what had been by then obtained in the east.

The Nineteenth Century (to 1898) Beccari’s successful explorations in the Vogelkop Peninsula, including his penetration of the Arfak Mountains, and the growth of the commercial bird plume trade led to a considerable increase of collectors in Netherlands New Guinea from the mid-1870s. But most collectors were there on their own account or had outside sponsorship. Several were under contract to Rothschild (Tring) or sold their hauls to the Ternate firms of A. A. Bruijn and C. W. R. van Renesse van Duivenbode. In the tradition of Wallace, the main attractions would be the more ‘‘attractive’’ birds, butterflies, beetles, and perhaps also shells and orchids; soon, as in eastern New Guinea, these would be joined (and partly supplanted) by ethnographic items. Most other plants and animals were of secondary importance. Many discoveries are due to these intrepid men, including mammals (notably Zaglossus bruijnii, the Long-beaked Echidna), various birds of paradise, bowerbirds, and orchids, along with a few other horticulturally useful plants. The richness of the orchid flora in western New Guinea, as in the east, also began to be revealed, especially after 1898. Often, however, the field data gathered by collectors was sketchy or absent; and over the years the paucity of field data has inspired additional searches—but for some taxa speculation remains (see, for example, The Lost Birds of Paradise (1995) by E. Fuller). The Vogelkop and Bomberai Peninsulas, Cenderawasih (Geelvink) Bay and its islands, the Raja Ampat archipelago (Misool, Kofiau, Salawati, Batanta, Waigeo and its satellites, and Gag), and Gebe´ (administratively now in Maluku) were popular destinations, but calls were also made at Humboldt Bay (the present site of Jayapura) and here and there elsewhere; some hunters reached the northern mainland Van Rees (Rouffaer) and Gauttier (Foja) Mountains. But until after 1898 the central ranges were but rarely, if at all, penetrated. Among the zoological (notably bird) collectors of this time were L. Laglaize (primarily birds of paradise but also other birds and butterflies, largely for Bruijn)

................. 16157$

$CH2

02-08-07 10:22:18

PS

PAGE 46

Biological Exploration of New Guinea / 47

in 1876–1884 in westernmost New Guinea and nearby islands (birds, Paris; insects, Brussels); C. Platen in 1883 on Waigeo (birds, Berlin) and, also in 1883, R. Powell on Waigeo and Salawati and in the Vogelkop Peninsula (birds; Tring, AMNH); F. H. H. Guillemard in 1883–1884 (on a cruise in Malesia with the Marchesa) at Waigeo, Batanta, Misool, Yapen, and the Vogelkop Peninsula (mainly birds but also some beetles, butterflies and shells; BMNH and Cambridge University, England); H. Kuhn in 1884 on the Onin Peninsula including Sekar (near Kokas) on the McCluer Gulf (now Bintuni Bay) (birds and other groups); H. Fruhstorfer in 1891 in the Vogelkop Peninsula (Takar, etc.) and elsewhere (including Lepidoptera; Berlin); and W. Doherty in 1892–1893 in many areas of the Vogelkop Peninsula, around Cenderawasih Bay (including Yapen Island) and on Humboldt Bay (birds and insects, the former at AMNH and the latter at BMNH). A collection of various animals from the Charles-Louis Range at the western end of the Central Cordillera—apparently one of the earliest such visits— was made through Renesse van Duivenbode and reached BMNH in 1898. In 1899 a Eurasian professional collector, J. M. Dumas, collected birds at Mt Moari on the Vogelkop Peninsula (Tring/AMNH), beginning an association with New Guinea which would last until 1911. In the latter part of the year the Raja Ampat Islands and a few points on the mainland (one of them near Fakfak) were visited by the Siboga oceanographic expedition (further mentioned below). In the new century J. Waterstradt in 1902–1903 visited Waigeo (birds, Copenhagen and Tring/AMNH), while the Pratts (see above) were at Merauke for a short time in 1902 collecting Lepidoptera but due to local hostilities (see below) sailed on to the Torres Strait and Port Moresby. Terrestrial plant collectors were relatively few, both on the mainland and in the major offshore islands; the latter were largely neglected until the new century (and some for decades beyond). After the visits of Moseley and Beccari (with in 1876 a return by Teysmann to the Raja Ampat group), in 1888 Warburg (see above) called at Sekar (near Kokas) in the McCluer Gulf (now Bintuni Bay) during his tour through Papuasia and eastern Australia (phanerogams and cryptogams, Berlin (largely destroyed) with some also elsewhere). In 1893 the director of the Bogor Botanic Garden, M. Treub (with an assistant, Jaheri), briefly called at the Onin Peninsula (and the Aru Islands); and in 1900 these localities were also visited by the American plant explorer D. Fairchild (with an emphasis on living plants). Of plant-hunters for the trade, W. Micholitz (for Sander and Sons in St Albans, England) around 1890 collected orchids in Batanta and elsewhere (Kew), while D. Burke (for Veitch) in 1891 obtained orchids and other plants in the northern Arfak Mountains (Kew). The principal oceanographic expeditions visiting during and after 1875 were, in that year, the Challenger and Gazelle (for both, see above), and, a quarter-century later, the Siboga. The Challenger sailed along the north coast (including Humboldt Bay) before heading to the Admiralty Islands, Moseley in particular collecting drift objects off the mouth of the Mamberamo. The Gazelle called during June in McCluer Gulf (now Bintuni Bay), Naumann collecting at Sekar (not far north of the

................. 16157$

$CH2

02-08-07 10:22:19

PS

PAGE 47

48 / david g. f r o d i n

future Fakfak). The year-long Malesian cruise of the Siboga, commanded by G. F. Tydeman and with a scientific corps led by the zoologist M. Weber, called in the latter part of 1899 in several of the Raja Ampat islands as well as at a few mainland points, one of them being Ati-ati Onin (near Fakfak, just established as a government post). A wide range of marine (and some terrestrial) organisms was obtained, the plants (including numerous marine algae) collected by Weber’s wife A. Weber-van Bosse (Leiden, all collections). The expedition’s results appeared—like those of the Challenger—in the grand manner over several decades, Weber-van Bosse’s main work, Liste des Algues du Siboga, appearing in four parts (1913–1928).

The Last Frontier I: Major Expeditions and Surveys (1898–1914) The arrival of the Siboga in the western waters of New Guinea followed hard upon a change in official Dutch policy towards their easternmost possessions, still largely unexplored. This was provoked by increasing foreign interest and, not unnaturally, by the development of the neighboring British and German territories. On 1 March 1898 what had been a single administrative unit was divided in two, and for the first time official posts were established: Manokwari for the north, Fakfak for the west and south. In early 1902 a third post, at Merauke in the southeast, was set up for the control of armed Marind-Anim (Tugeri) raiders and headhunters—who were also creating disturbances in British New Guinea, then in some turmoil following the cannibalistic murder in 1901 of the now-aging Chalmers (see above). In later years other posts, among them Serui (on Yapen Island) and Hollandia (now Jayapura; on Humboldt Bay), would follow. In the sciences, too, the Dutch realized that serious attention was required, particularly to the extensive interior east of the Bird’s Neck. The scene was thus set for the series of large-scale wholly or partly governmentsponsored expeditions that largely dominated exploration until 1939 (with a ‘‘finale’’ in 1959, filling the ‘‘last white spot on the map’’). Results of these expeditions were over several decades largely presented in the special serial Nova Guinea (Leiden, 1909–1966), sponsored by the Indies Committee for Scientific Research and other bodies. The first of these major undertakings was the 1903 North New Guinea Expedition, led by C. F. A. Wichmann, then a geology professor at Utrecht University. Much of its zoological material (Amsterdam) was collected by H. A. Lorentz and L. F. de Beaufort as well as by Dumas (see above). No qualified botanist was in the party, but plants—both living and preserved (Bogor), mainly from the first part of the trip—were collected by the Indonesian officials (mantris) Atasrip and Jaheri from Bogor (Jaheri previously having visited New Guinea with Treub and, in 1901 with the Java, Fakfak, the site of Merauke, and Thursday Island in Torres Strait—there collecting Deplanchea tetraphylla) and, where possible, by Dumas after the Bogor officials’ (mantris) departure. A wide range of mainly coastal localities was visited by Wichmann, ranging (on his own, in January) first from Triton Bay along the Bomberai coast (including the offshore Adi, Karas, and Semai islands) to Fakfak (and McCluer Gulf, now Bintuni Bay), and from there (on the

................. 16157$

$CH2

02-08-07 10:22:19

PS

PAGE 48

Biological Exploration of New Guinea / 49

expedition proper, 7 February to August) around Geelvink (now Cenderawasih) Bay and its islands, afterwards rapidly sailing along the north coast to Humboldt Bay and there taking in the Hollandia (now Jayapura)-Sentani area (after which the Indonesian officials (mantris) returned to Java), finally exploring around Tanah Merah Bay and elsewhere before returning west and concluding with an excursion to the Bird’s Neck at the southern end of Geelvink Bay. An extensive general report appeared in 1917 in Nova Guinea; natural history reports were scattered. Specific localities on the north coast and in Geelvink Bay included Waigeo, Manokwari (on Dore´ Bay), Mansinam, Karoon, Kwawi, Andai, Wendesi, Tawarin, Bawe, Sageisara, Moso, Napan, Angadi, Jende (on Ron I.), Timena, Orum, Mios Korwar, Supiori Island (just west of Biak), Ansus on Yapen Island, Moso, Metu Debi, Tjintjan Bay, Matterer Bay, Pokembo, Wakobi, Siari, Kwatisore, and Wa Udu and (along the north coast) Moaif and Maffin Bay. Points of interest in the Hollandia (now Jayapura)-Sentani area included (among others) Jotefa Bay, the Cyclops Mountains (ascending into the range in the first two-thirds of April and visiting Mts Pisero, Sinagai, and Pancana among others), and (west of Sentani) the Timena River, Ibaiso, and Jaga. Slightly more distant were Tanah Merah Bay (at the western end of the Cyclops), the Korime´ River inland from Moeris, and— just past the German border—Oinake´ (near Wutung in present-day Papua New Guinea). South of Hollandia (now Jayapura) they visited the Tami and Sekanto Rivers. In mid-July they returned to Manokwari and the team then explored (via Kwatisore) across the Bird’s Neck, reaching Goreda (on Lake Yamur, not far from Jabi at the very western end of the Central Cordillera). Thence they attempted to push further south before in mid-August but, plagued by mosquitoes, had to abandon their transit and so turn back to the north coast, Manokwari, and ‘‘home.’’ Wichmann’s undertaking was followed in 1904–1905 by the Southwestern New Guinea Expedition, organized under the auspices of the Royal Dutch Geographical Society and led by R. Posthumus Meijes and E. J. de Rochemont. Animals and plants (Bogor, Leiden) were collected by Dr J. W. R. Koch with the aid of Indonesian officials (mantris) from Bogor; Koch, a physician by profession, focused in particular on ethnography. In the second quarter of 1904 (prior to the expedition proper), he made a preliminary reconnaissance from Merauke with the Lombok, taking in the rarely-visited Frederik Hendrik Island (later known as Kolepom Island, now known as Yos Sudarso Island) as well as parts of the southwest coast and its inlets. He then returned to Merauke for some months before arrival of the expedition proper (from September to the following April). The full party (in the Valk) carried out further work along the southwestern coasts, with one side trip to Dobo, Aru Islands, as well as at Merauke and sailed up the lower course of the Digul (partly in sloops). During this time, Koch was left at Etna Bay (west of Uta) for over two months. The main expedition report appeared in 1908, Koch contributing the sections on ethnography and natural history (with assistance from T. Valeton on the plants). During this cruise Mt Wilhelmina (now Mt Trikora) was first sighted and named from the expedition ship.

................. 16157$

$CH2

02-08-07 10:22:19

PS

PAGE 49

50 / david g. f r o d i n

Valeton incorporated available identifications into his Plantae Papuanae (1907), a work which for the south—most of Koch’s collections having come from Merauke—represented a useful addition to the earlier writings of von Mueller. But Koch’s harvest at Etna Bay was relatively small, as was that in the north; the botanical results of both expeditions thus are, with the exception of records from south of the Digul, relatively slight. By the middle of the decade, Dutch detachments had, with the Lombok and other vessels, explored most of the coasts and penetrated the lower courses of some of the rivers. The stage was now set for more serious inland exploration. From 1907 to 1915 there took place one of the greatest organized exploring efforts of a territory ever—the Military Expeditions (Militaire-Exploratie). Over eight years they obtained an extensive albeit sketchy knowledge of the interior lowlands and associated hills. On certain sorties plants and animals were collected, notably in 1909–1912 by the Dane Dr. K. Gjellerup (see below), but also by Dr. R. F. Janowsky, W. K. H. Feuilletau de Bruijn, Lt. L. A. C. M. Doorman, P. M. van Kampen, Dr. J. K. van Gelder, and Dr. A. C. de Kock. Members of the military teams also accompanied the major Dutch and British expeditions to the central ranges (see below), and Gjellerup and van Kampen joined the 1910–1911 GermanDutch border survey (the Dutch contingent being led by J. J. F. C. ten Klooster). While with the Military Expeditions (Militaire-Exploratie), Gjellerup collected plants and animals (Bogor; plants also in Leiden, Kew, and Utrecht) as other duties permitted. In this work he was assisted by two Indonesian officials (mantris) from Bogor, Ajoeb and Sadeli. On travels in the border areas in 1910 and 1911 he three times visited the Tami River basin (reaching among other places Arso and Sawia) and also undertook a trip to the Bougainville Range (near Oinake´). In between were assignments in 1910 with the German border survey team (under Schultze-Jena) to the Bewani Mountains, from there returning down the Tami River and then the upper Sepik River. Due to the capsizing of a vessel, many collections from these latter trips unfortunately were lost. After the Tami River basin Gjellerup then worked along Lake Sentani and in the Cyclops Mountains, in mid-1911 as high as 2,000 m, obtaining the first significant harvest of plants (including, for example, the endemic Schefflera leiophylla) from this geologically interesting but difficult range. This was followed by a reconnaissance of the Maffin (now Tor) River (southeast of Sarmi), reaching into the Gauttier (now Foja) Mountains. After further activity in early 1912 around Hollandia (now Jayapura), he was transferred to Manokwari. From there, with a geologist and mining surveyor, P. F. Hubrecht, via Siari (on the coast) he climbed up to and collected at the Anggi Lakes (April–May 1912)—a year and a half before Gibbs (see below). Here, also, he suffered losses to his collections. Of other Military Expedition (Militaire-Exploratie) officers, Janowsky—during two trips in 1912 and 1913—penetrated far up the Weyland Mountains at the western end of the central ranges (in 1913 reaching the top at 3,720 m) but also patrolled along the eastern side of Geelvink Bay (Moesoiro, Legare River, Sawa River). Both animals and plants were obtained (Bogor) though a large number of

................. 16157$

$CH2

02-08-07 10:22:20

PS

PAGE 50

Biological Exploration of New Guinea / 51

those from 1913 had to be abandoned in the field. Feuilletau de Bruijn in 1914 collected in the Mamberamo basin and the Lake Plain and in 1915 in the Schouten Islands (plants, Bogor). In 1914, on the third Mamberamo expedition of the Military Expeditions (Militaire-Exploratie) under Capt. J. V. L. Opperman, Lt Doorman went up the Mamberamo (calling at the Pionierbivak base camp) and across the Lake Plain. Finally, via the Taritatu (formerly Idenburg) River, they reached the summit (3,550 m) of Mt Kembu (later named Doorman Top), this being some ways east of Moszkowski’s route (see below). Doorman’s collections were but few (orchids, Bogor). Gelder, in addition to his mineral surveying, collected animals in 1910 on the Mamberamo under Capt. A. Franssen Herderschee. Kampen in 1910–1911 collected animals at Hollandia (now Jayapura), Lake Sentani, Zoutbron, and Bronbeck (1910) and on the Arwo River (1911). Finally, in 1910–1911 de Kock made a dramatic stab towards the eastern part of the central ranges, voyaging up along the Eilanden River and ascending Mt Goliath (3,500 m) where for some time de Kock remained (preserved and living plants, Bogor). That remote mountain region has since hardly seen a collector. Also with some parties was J. M. Dumas (see above), collecting mainly birds and insects (Bogor); he would also join Lorentz’s first expedition (see below). The results from these several undertakings appeared in Nova Guinea and elsewhere. In spite of these cumulatively not inconsiderable contributions, detailed biological investigation was, however, not a primary aim for the military teams. Indeed, no professional botanist and only one professional zoologist were under direct command; the majority of the collections that emerged were the work of team physicians. More significant for science were the major metropolitan expeditions that army men accompanied and, in one instance, also led. Most of these had as their main objective the ‘‘snow mountains’’—a Dutch ‘‘dream’’ since their sighting centuries before by Carstensz. In this quest the generally state-supported Dutch were to be ‘‘challenged’’ by largely privately-funded British interests. By agreement, however, the British directed their attention towards Mt Carstensz (now Mt Jaya)—perhaps the greater prize, being the highest mountain between the Himalayas and the Americas. (Only in 1962 would it be successfully climbed, and then by a less difficult route.) The Germans, as we shall see (and perhaps not to be outdone), also made one abortive attempt in their support of Moszkowski— but from the north, at that time a rather more difficult route although the opening up of the central ranges by the Military Expedition (Militaire Exploratie) teams was underway. The other expeditions sensibly approached the central ranges from the south. Of the three Dutch ‘‘South New Guinea Expeditions,’’ only the third finally reached the top of Mt Wilhelmina (now Mt Trikora), although the second got to over 4,000 m, turning back just 170 m below the summit. The first two expeditions were led by H. A. Lorentz and the third by Capt. Franssen Herderschee (earlier leader of a military team on the Mamberamo, as we have seen). All traveled from the so-called Asmat (or Casuarina) Coast in the first instance up the Noord (now Lorentz) River, with a base camp at ‘‘Alkmaar’’ (ca 100 m) at the start of the

................. 16157$

$CH2

02-08-07 10:22:20

PS

PAGE 51

52 / david g. f r o d i n

foothills. Attached to each of these expeditions was an army lieutenant; for the second, it was D. Habbema. The first South New Guinea Expedition (1907–1908) worked extensively around Merauke and in the Digul and Noord (now Lorentz) river basins; the furthest point reached was the Hellwig Mountains (2,320 m) with an intermediate point (the Resi Mountains) at 900 m. The ‘‘Alkmaar’’ base camp was established at this time. Much collecting was at lower elevations, such as at Sabang and the van Weels camp on the Noord (now Lorentz) River. Animals were obtained by Lorentz and de Beaufort (see also above), while plants were collected by Dr. B. Branderhorst (also of the Militaire Exploratie), Dr. G. M. Versteeg (directly seconded from the forces) and the Bogor officials (mantris) Djibdja and Atmodjo (animals, Leiden; plants, Bogor and Utrecht). The second (1909–1910) and the third (1912–1913) South New Guinea Expeditions both worked all the way from the mouth of the Noord (now Lorentz) River to the summit area of Mt Wilhelmina (now Mt Trikora), going via the Alkmaar camp and the nearby (and slightly higher) Kloofbivak camp north along the ridges between the Noord (now Lorentz) and Noordwest Rivers. They passed through Heuvelbivak base camp, the Went Mts, Dromedarisbivak camp, the Hellwig Mts (2,000 m), the Treub Mts, the Wichmann Mts, the Hubrecht Mts, Bellevue, Peripatusbivak camp, Jenjabivak camp, the Kajan Mts, Waterfalbivak camp, Lake Quarles (ca 3,600 m), Dolomieten, Oranjebivak camp, Rotsbivak camp (4,300 m) and, in November 1909, first onto the then-extant ice-cap on Mt Wilhelmina (now Trikora; 4,750 m). Lake Habbema was seen for the first time—a site very important a quarter-century later, for the Third Archbold Expedition (see the section Between World War I and World War II, below). The same route was followed in 1912–1913. Some collecting was also done at Fakfak and Kaimana on the southwestern coastal approaches. Most of the animals were collected by Lorentz (second South New Guinea Expedition) with some by G. M. Versteeg (third South New Guinea Expedition). A considerable number of plants were obtained under Lorentz by E. S. A. M. Ro¨mer, J. W. van Nouhuys, and Habbema, and under Herderschee by A. Pulle, but at the high elevations by Versteeg (Bogor, Leiden). The richness of the flora of the central ranges now began to be revealed, including a first (but for some time unrecognized) collection for New Guinea of southern beech, Nothofagus—a dominant genus in many areas. A rich array of results appeared in Nova Guinea, including large numbers of new orchids. British explorers mounted two expeditions: the British Ornithologists’ Union Expedition of 1909–1910 and the Wollaston Expedition of 1912–1913. Like their Dutch counterparts, each expedition was accompanied by a Dutch army officer. The first British Ornithologists’ Union Expedition was led by W. Goodfellow; animals (but few plants) were collected by A. F. R. Wollaston, G. C. Shortridge, and W. Stalker. Working mostly along the Mimika, Utakwa (also known as Waitakwa) and Setakwa rivers, the expedition met with great difficulties and some loss of life (Stalker being drowned) and in addition failed to get high in the central range. Wollaston, however, soon organized his own expedition to continue the quest.

................. 16157$

$CH2

02-08-07 10:22:21

PS

PAGE 52

Biological Exploration of New Guinea / 53

Assisted by C. Boden Kloss of the Federated Malay States Museums (Kuala Lumpur), he collected animals and quite a few plants (BMNH; plants also at Kew and elsewhere), also making many geographical and topographical observations (regrettably, most of his notes were lost during the descent). The team reached the Mt Carstensz (now Mt Jaya) glacier fields but was finally stopped at the end of January 1913 by stupendous cliffs and a wall of ice; due to bad weather they did not realize they were still some 550 m below the summit. The animals (by several zoologists) and plants (by H. N. Ridley) were published mainly in Transactions of the Linnean Society, and as well collectively reissued as a fine two-volume set. Wollaston’s expedition was recently revisited in a memoir by his son, Nicholas, My Father, Sandy (2003). The challenge of the ‘‘snow mountains’’ would, as already mentioned, attract one German explorer—another physician-naturalist and ethnographer, Max Moszkowski of Breslau (Wroclaw) University. In 1910–1911 he attempted to reach the ‘‘snow mountains’’ via the Mamberamo River. After explorations in its lower regions (including Teba, Sauwi, Samberi, Assewari, and Tama, and in the lagoon areas and estuaries to the east of the Mamberamo), he began his ascent south from a base camp on the Naumoni River, just before the foothills. But soon, at Edi Falls, just before Pionierbivak camp (ca 225⬘S) Moszkowski lost all his equipment and had to return to Manokwari. After his return to Naumoni Moszkowski passed the falls and through the Van Rees Mountains, and entered the Lake Plain. However, by now—not surprisingly—he was becoming overextended. Towards the end of 1910, while some ways up the Van Daalen (or Zuid River), a tributary of the Tariku River (ca 1385⬘E, in the Nassau Ranges) he had to turn back on account of lack of food. In January 1911, again at Edi Falls, while returning he lost many of his collections. A number of plants were, however, brought back (Berlin, Leiden); zoological and entomological (Berlin) as well as ethnographic collections were also made. Moszkowski’s own contributions, apart from his expedition reports, are all in ethnography and anthropology, a not insignificant accomplishment.

The Last Frontier II: Independent Exploration (1898–1914) In addition to the great expeditions, there were a number of individual undertakings in the final years of the post-Napoleonic century. Some related to official activities; but, as before, other explorers worked independently or under outside sponsorship. Many—official as well as non-official—also were keen photographers; advances in equipment and technique resulted in great additions to the photographic record. But offsetting this was a contemporary decline in fine color-plate art, one of the last of the ‘‘grand’’ hand-painted works having been R. B. Sharpe’s Monograph of the Paradiseidae, or Birds of Paradise, and Ptilonorhynchidae, or Bower-Birds (1891–1898), a book with much on Papuasian exploration. Not surprisingly, with better links the western regions and islands continued to attract the most attention. Of particular interest after 1900 would be the newly discovered twin Anggi Lakes in the southern part of the Arfaks. In April 1904, A. van Oosterzee, from its establishment in 1898 administrator at Manokwari and a

................. 16157$

$CH2

02-08-07 10:22:21

PS

PAGE 53

54 / david g. f r o d i n

zealous explorer of his district, sent living plants (Bogor) from these lakes—as L. S. Gibbs has written, the first European to collect there. Yet, on van Oosterzee’s visit, potatoes were already available from local gardens—the result of indirect missionary introductions. In 1906–1907 the American zoologist Thomas Barbour collected amphibians and reptiles around the Anggi Lakes and on Waigeo Island (MCZ). Barbour was followed up to the Anggi Lakes in 1907 and again—for a stay of close to a year—in 1908–1909 by Pratt with his sons F. and C. In between (1907–1908) the Pratts spent much time around Humboldt Bay, including visits to the Cyclops Mts and Lake Sentani (insects, Tring/BMNH; plants, Bogor, Kew). After the Pratts came Gjellerup with Hubrecht in 1912 (see above) and, in 1913, the senior Pratt. The senior Pratt would lend photographs to Gibbs (see below) for her book, her own not having been satisfactory. But perhaps the most important independent figure of the period is L. S. Gibbs—the first independent lady explorer in New Guinea and moreover the first visiting scientist with an explicit interest in tropical mountain ecology and vegetation. In six days in December 1913 she obtained over 330 numbers (BMNH, with some duplicates elsewhere) at the Anggi Lakes (via the coastal village of Wariap), with some 150 more over the rest of her trip (Dore´ Bay, Roon Island, Biak Island, Wakde´ Island, and Humboldt Bay—in all from November 1913 to February 1914). Her results, which also included checklists of her collections with descriptions of novelties, appeared as Dutch N.W. New Guinea (1917)—a botanical landmark, later to be drawn upon by P. W. Richards for his classic Tropical Rain Forest (1952). Gibbs had earlier visited Borneo (Mt Kinabalu) and Fiji as part of a quest towards working out the structure and origins of the Malayo-Pacific mountain flora—much of it featuring an ‘‘extra-Malayan facies,’’ as Beccari (see above), von Mueller, and some others had earlier indicated. In the northern mainland, Walter Goodfellow visited along the coast, collecting birds in 1904–1906 (BMNH). In 1915 C. L. J. Palmer van den Broek—who was very helpful to Gibbs while Resident (in Ternate)—collected in the Cyclops Mountains and Humboldt Bay (Bogor). In the south, Merauke and its biotically distinctive region had now become relatively accessible. Hassan collected various animals on the Utumbui River and on the Gelib River and at Okaba (west of Merauke) in 1909–1910 (Leiden; plants Bogor)—perhaps with Versteeg as part of the second of Lorentz’s expeditions (see above). In 1910 Rothschild’s agent A. S. Meek collected birds and insects at Merauke, along the Digul and Eilanden rivers, ascending Mt Goliath (in the wake of the Dutch expedition), there reaching 2,800 m (birds, Tring/AMNH; insects BMNH). H. Elgner collected insects in 1912 at Fakfak (Senckenberg). A few further collectors visited the western islands, some of them as members of two expeditions from Freiburg, Germany. In 1907–1908 Roux, a Swiss, and H. Merton, a German, collected various animals on the Aru and Kai Islands. Misool was visited in 1911 with O. D. Tauern collecting animals (Leiden). Botanically, however, after 1898 these islands would largely be neglected, remaining so until at least the 1930s.

................. 16157$

$CH2

02-08-07 10:22:22

PS

PAGE 54

Biological Exploration of New Guinea / 55

With the outbreak of World War I in mid-1914, field activity largely came to an end. Travel became restricted and war conditions soon disrupted world shipping. Though the Netherlands and the Indies remained neutral, after 1915 further exploratory work became unfeasible. The final report of the Dutch Military Expeditions (Militaire Exploratie) appeared in 1920—a very useful summary of what had been accomplished, though not rich in biological data. Among its many maps is one in four sheets for the whole territory—the best then available; another (after p. 74) depicts the Military Expedition teams’ routes, as well as the routes of earlier explorers. It would be five years before serious activities resumed in western New Guinea, and then not for long in the old tradition. Costs of everything were higher, and—significantly—much of the primary interest in exploration had been satisfied. A zenith had passed, although with three more major undertakings to come in the 1920s there would be for a time somewhat more continuity in biological exploration in the west of New Guinea as opposed to the Australian-administered eastern territories. In publication, few new ‘‘grand series’’ would emerge. Some earlier runs, such as Nova Guinea (with many illustrated contributions on orchids by J. J. Smith) and the Siboga volumes, continued, but—as already for the eastern part of New Guinea—the presentation of results generally became more diversified as well as modest, usually appearing in specialist professional journals.

Between World War I and World War II (1918–1942) Although German New Guinea had come under Australian military rule very early in World War I, Papuasia otherwise was not a theater of war and most local administration continued uninterrupted, aided by enhanced revenues for commodities. Some collecting could thus be accomplished in the later 1910s: W. Bradtke in the Duke of York Islands, 1917 (plants, Brisbane); John Todd Zimmer in Woodlark Island, Papua, collecting animals and coconut pests (the latter as part of his extension work), 1917–1918 (BMNH); and, in mid-1918, C. T. White in central Papua (see Plants section below). Keysser also was active in the Saruwaged Mts, as has been mentioned. But it was not a time for large expeditions. After November 1918, however, normal biological work could resume. Initially it developed relatively slowly (except for the major expeditions of 1920–1922 across northern and central Dutch New Guinea, mentioned below) but it gained pace in the later 1920s and again, in a more favorable economic climate, in the 1930s. Yet not for some time in the former German territories (under civil administration only from 1921) did exploration approach its earlier level, a development that attracted some adverse comment. In addition, everywhere infrastructure and resources would largely remain too rudimentary for natural history institutions (as Macgregor already had foreseen). Similarly, official interest in animal and plant life was low save with respect to forestry, agriculture, marine, and wildlife resources (in particular with the end of the bird plume trade resulting in losses in local income and revenue). Official publications of the time likewise reflect the era. Hardly a mention of Dutch New Guinea appears, for example, in Handbook

................. 16157$

$CH2

02-08-07 10:22:22

PS

PAGE 55

56 / david g. f r o d i n

of the Netherlands East Indies (1924, Buitenzorg). By the late 1930s two substantial works had appeared, both with some natural history and resources content, the latter also covering exploration. These were the Australian Official Handbook of the Territory of New Guinea (1937, Canberra; reissued 1943) and the more ambitious Dutch (but New Guinea–wide) compendium, W. C. Klein’s three-volume Nieuw-Guine´e (1934–1938; see References section below). During the quarter-century between the wars, developments lessened the need for large expeditions, while at the same time more individuals were working for greater or lesser periods in the field—a number of whom have left accounts of their adventures. Innovations such as radio and the airplane, as well as aerial photography (well covered, for example, by Klein, 1934–1938), were of particular importance. The number of administrative, mission, and other posts also increased. Although the majority of these posts continued to be on or near water, some church groups (notably the Sacred Heart Roman Catholic mission at Yule Island and the Lutherans at Simbang and Sattelberg) had by then developed extensive graded track systems to their interior mountain stations. There were also some—generally less elaborate—systems of government tracks, including the track across the Kokoda Gap. Motor roads, however, remained few, but from the late 1920s this lack was partly offset by airstrips and the rise of commercial aviation, notably in the Mandated Territory. By the 1930s air links with nearby parts of the Indies and with Australia had also been developed. Collectors and specialists were still predominantly European, but now they came to be joined by others from elsewhere, particularly Japan, Australia, and, increasingly, the United States. There was moreover a greater amount of collaboration than in previous decades, although a partial vogue for expedition reports remained (Nova Guinea, Results of the Archbold Expeditions, etc.).

general undertakings The first major expeditions of the era between the two World Wars were the two over 1920–1922 in Netherlands New Guinea aimed at reaching Mt Wilhelmina (now Mt Trikora)—the furthest point reached by the Herderschee expedition of 1912–1913—but across the central range from the north, as earlier attempted by Moszkowski. The last great exploring expeditions in the pre–World War I style, these ‘‘New Guinea Expeditions’’ were led in turn by Dutch Army captains A. J. A. van Overeem (1920–1921) and J. H. G. Kremer (1921–1922). The approach for both was up the winding Mamberamo River through the Van Rees Mts into the Lake Plain, with van Overeem going up the Idenburg, Rouffaer, and Doorman rivers, finally reaching the Swart Valley and the summit of Mt Doorman (the highest peak along the northern fall of the Nassau Range). Camps were set up at Prauwenbivak, Bivak Batu, Pionierbivak, Doormanpadbivak, Mamonbivak, and Kikkerbivak. After reorganization and resupplying, Kremer (with 800 men!) returned to the field and retraced much of the previous route to the Swart Valley; from there some of the party crossed into the western Baliem basin (missing, however, the Grand Valley) and finally reached Lake Habbema (first seen in 1909)

................. 16157$

$CH2

02-08-07 10:22:22

PS

PAGE 56

Biological Exploration of New Guinea / 57

and nearby Mt Wilhelmina (now Mt Trikora)—at the end of the longest largely pedestrian supply line ever organized in New Guinea. But, as Souter wrote in 1963 (see Reference section, below), the van Overeem and Kremer expeditions were the last of their kind. Never again would such battalion-sized parties take to the field; future large expeditions would rely on radio (first tested by van Overeem), fixed-wing aircraft, and (after World War II) helicopters and motor vehicles. Indeed, the size of these expeditions—particularly in 1921–1922—were not commensurate with the scientific results. The onset of a severe economic recession in the Netherlands East Indies also frustrated any new official plans. Collections in the first phase of these two major 1920–1922 expeditions were made by W. C. van Heurn (animals) and H. J. Lam (plants, Bogor, Leiden, Utrecht), the plants written up in Nova Guinea and elsewhere. In Fragmenta Papuana (English version, 1945) Lam also presented a narrative and set of observations from his work which remains an accessible account of the van Overeem undertaking. Circumstances in the second phase of these expeditions were less favorable to biological collecting; the only substantial contribution was the work of the ethnographer-anthropologist Paul Wirz, who in 1922 remained in the Swart Valley while the rest of the party pushed southwards. He also collected animals (Leiden, Amsterdam), obtained both from there and on Mt Doorman. A few plants (Leiden) were, however, obtained by Hubrecht who was with the main party and moreover a veteran of Herderschee’s successful ascent. The van Overeem and Kremer expeditions were followed in western New Guinea by the ‘‘Netherlands-America Expedition’’ of 1926 under M. Stirling, with the former Dutch army officer C. C. F. M. le Roux as topographer-ethnographer and W. M. Docters van Leeuwen from Buitenzorg as botanist (collections, Bogor, Leiden). While more of an explorer (as well as an anthropologist), Stirling (with his pilot R. K. Peck) pioneered the use of aircraft in New Guinea as a transport aid. Their route to the Lake Plain was similar to that of van Overeem and Kremer, and some of their campsites—in fact dating back to the time of the Dutch Military Expeditions (Militaire-Exploratie) of 1907–1915—were reused (Le Roux making some side trips). It was along this stretch that their amphibious plane saw most use. The upper route followed the Rouffaer River along and then into the Nassau Range up to an altitude of 2,600 m (with Explorationbivak the most distant camp). Deterioration of the plane, however, forced its withdrawal and so brought a premature end to the expedition. Few direct results ever appeared; in particular, both in 1932 and again during World War II, key notes and lists relating to the botanical collections were lost. Only gradually have the collections themselves (often with little data) been worked up as particular families have been revised. The major contribution was thus le Roux’s three-volume monograph on Papuan mountain dwellers, De Bergpapoea’s van Nieuw Guinea en hun Woongebied (1938). The last years of the Jazz Age boom saw three biological ‘‘cruises’’ arrive in New Guinea waters: two American, one Belgian. The first was the Whitney South Seas Expedition of 1928–1929, which collected birds in eastern Papua and in the Milne

................. 16157$

$CH2

02-08-07 10:22:23

PS

PAGE 57

58 / david g. f r o d i n

Bay Islands (AMNH); it also was active elsewhere in the Pacific. The second was the visit in 1929 of Crown Prince Leopold of Belgium (later Leopold III) and his wife Princess Astrid, together with their chief scientist Victor E. van Straelen. During a voyage that ranged widely through the Netherlands Indies, they visited several spots in western New Guinea, including the Raja Ampat Islands, Sorong, Manokwari, the Arfak Mountains and Anggi Lakes, Yapen Island, and, in the south, Triton Bay (the site of long-abandoned Merkusoord). Van Straelen’s collections of insects, other animals, plants, and fungi are in Belgium (in Brussels and Meise (BR), respectively), but among the plants and fungi are only algae, mosses, and lichens. A special series from the Natural History Museum in Brussels (of which van Straelen was, in time, director) along the lines of Nova Guinea, Re´sultats scientifiques du Voyage aux Indes Orientales Ne´erlandaises . . . , encompassed algae (1932) as well as the extensive zoological results. Van Straelen also wrote a more popular book, De Reis door den Indischen Archipel van Prins Leopold van Belgie¨. The last ‘‘cruise,’’ also in the first half of 1929, was the Crane expedition on its yacht Illyria. Sponsored by the Field Museum in Chicago but also with some input from Bostonians, it was headed by Cornelius Crane and S. N. Shurcliff with as chief biologist a former Philippine National Museum ichthyologist, A. W. C. T. Herre. During this world cruise calls were made in the Solomon Islands, the Gazelle Peninsula of New Britain, Huon Gulf, the Sepik River, Manokwari, and Waigeo. The team collected marine specimens as well as many freshwater fish, other animals, and some 400 or so plants (FMNH; plants also at NY). As with the Belgian expedition, the results included a popular book, Jungle Islands (1930), as well as scientific reports by Herre and others (mainly in the Field Museum’s zoological series). The 1930s would be marked by five large expeditions, three from the United States and two from the Netherlands (and East Indies). Those from the United States—the so-called Archbold Expeditions—were organized through the American Museum of Natural History, New York, and personally financed and led by a Standard Oil heir, Richard Archbold. An associate in mammalogy at the Museum and an amateur pilot, Archbold was accompanied by a small cadre of experienced scientist-collectors, particularly the zoologist Austin L. Rand and the botanist Leonard J. Brass. These expeditions—each of them over a year long and progressively more ambitious—were organized so as to encompass substantial altitudinal transects in different areas, each thought to be imperfectly known as well as potentially accessible. Amphibious planes were used for the second and third expeditions— though (alas!) there was a serious accident during the second, when in Fairfax Harbor (Port Moresby) a sudden wind flipped over the expedition’s craft while it was at anchor and it sank, forcing a change of plans. The many places visited are well described (and mapped) in general reports by the team in the Bulletin of the American Museum of Natural History (with Brass playing a considerable role in drawing them up). The second expedition was also the subject of a popular book, New Guinea Expedition (1940). Their aerial exploits and achievements have been well described in Souter (1963; see section on References, below) and elsewhere.

................. 16157$

$CH2

02-08-07 10:22:23

PS

PAGE 58

Biological Exploration of New Guinea / 59

At the American Museum of Natural History, the undertaking was granted unitary status as ‘‘Archbold Expeditions’’; through it were coordinated the expeditions’ scientific results. A separate series of reports comparable to Nova Guinea was, however, eschewed in favor of established serials; but individual papers generally bore a subtitle ‘‘Results of the Archbold Expeditions.’’ Zoological papers—many substantial—were mainly published in the Bulletin of the American Museum of Natural History, while the botanical appeared in Journal of the Arnold Arboretum, Brittonia, the later installments of Lauterbach and Diels’s series Beitra¨ge zur Flora von Papuasien, and elsewhere. The First Archbold Expedition (March 1933–March 1934) was, by its own admission, something of a ‘‘trial horse,’’ sticking to relatively established means of land and water access. Accompanying Archbold were Rand (birds and other animals; AMNH) and Brass (plants, NY with duplicates elsewhere). The First Archbold Expedition worked a complete altitudinal transect from the south coast to the top of the Wharton Range in the western Owen Stanley Mountains. Here, access was facilitated by the already-mentioned Sacred Heart track to Ononge (south of modern Woitape), thence overland to Murray Pass. Stations towards Ononge included Kubuna, Dieni, Mafulu (1,200 m), and Mt Tafa (2,400 m); beyond there—towards Mt Albert Edward and the Neon Basin, the ultimate objectives—camps were made at Urunu, Ero Creek, and Gerenda (this last below the summit area). After its return from the mountains the expedition visited Yule Island en route to Port Moresby for Christmas. Its last foray (from January 1934) was by boat to Kikori, Daru, and the Binaturi and Oriomo rivers (including collecting stations at Wuroi and Dogwa), with also a first visit to the low Oriomo Plateau (northwest of Daru). The thirteen-month Second Archbold Expedition to New Guinea (February 1936–March 1937) was centered in the Fly River basin, including the Fly River itself, its upper tributaries the Palmer and Black rivers, and parts of the Strickland River as well as some of the lakes (notably Daviumbu Lake and Murray Lake). Access to the area had improved as a result of oil and mineral prospecting activities, with Oroville Camp (now Kiunga) a potentially useful staging point for the principal objective, exploration of the Hindenburg limestone range between the Fly River and the upper Sepik River. In addition to a ketch, the expedition also had a plane (used particularly out of Daru); but, on 9 July, the plane was—as already mentioned—wrecked at Port Moresby. The just-initiated mountain work therefore had to be abandoned. Instead the party, after much work in the middle Fly, spent its final three months in extensive exploration of the so-called TransFly region (between the lower Fly River and the Torres Straits and up to the international border). In addition to Archbold, Rand, and Brass, an American Museum mammologist, G. H. H. Tate (who would return on post–World War II Archbold Expeditions) was also in the party (animals, AMNH; plants, Harvard University Herbaria with duplicates elsewhere). Collecting stations include Rona (or Rouna) near Port Moresby, Daru, Mabaduan (notable for a granite outcrop geologically homologous with those in the Cape York Peninsula of Australia),

................. 16157$

$CH2

02-08-07 10:22:24

PS

PAGE 59

60 / david g. f r o d i n

Everill Junction, Oroville Camp, Palmer River (one month), Black River (two months), Lake Daviumbu (one month), Sturt Island (one month), Gaima, and (in the Trans-Fly) along the channels of the Wassi and Mai Kussa River, calling at Penzara, Tumbuke, and, in particular, Tarara; finally, Daru was once more visited. The Third Archbold Expedition (also known as the Netherlands IndiesAmerican Expedition), of thirteen months’ duration (April 1938–May 1939), was carried out jointly with Dutch interests. Working with Archbold, Rand, and Brass were entomologists L. J. Toxopeus and J. Olthof, another zoologist, W. B. Richardson, and forest botanists E. Meijer Drees and C. Versteegh. The expedition traversed in particular the Nassau Range from Mt Wilhelmina (now Mt Trikora) to the Lake Plain, partly by boat and land but mainly in a new amphibious plane, the Guba (one of two prototype PBYs, later the famous ‘‘Catalina’’ class of World War II and beyond). After fieldwork in the vicinity of Hollandia (now Jayapura) and the Cyclops Mts, a first mountain camp was established at Lake Habbema (3,225 m), with the Guba successfully landing and taking off from the water—then a new altitudinal record for a seaplane. From there camps were established at Letterbox (3,560 m) and Scree Valley (3,800 m), the latter near the summit of Mt Wilhelmina (now Mt Trikora). The expedition then worked its way northwards, with collecting stations at Moss Forest (upper Bele Valley, 2,800 m), Bele (2,200 m), and Baliem (1,600 m), the latter in the Grand Valley—a new ‘‘discovery,’’ of which parts were explored by the team. Then followed a detailed examination of the central-eastern part of the Nassau Range, with stations at Top (2,150 m), Mist (1,800 m), Sigi (1,500 m), Rattan (1,200 m), Araucaria (800 m), and, by the Idenburg (now Taritatu) River, Bernhard Camp (50 m), the last named in honor of Prince Bernhard of the Netherlands. Very extensive collections were amassed, including over 5,500 numbers of plants obtained by Brass, Meijer-Drees, and Versteegh (vertebrates at AMNH; insects at Leiden after processing at Bogor; plants in the Harvard University Herbaria with many duplicates in Bogor and elsewhere). Apart from its great geographical discovery, the expedition was the first to recognize the presence of Nothofagus in New Guinea—although specimens had been collected earlier, they initially were not correctly identified. The substantial Dutch representation in this expedition was a reflection not only of pride but also a return of better economic conditions in the Netherlands Indies. Already the Dutch had resumed exploration (and resource investigation) in their own right, a notable goal being to ‘‘fill in’’ the remaining ‘‘white spaces’’ on the map—clearly evident to the world when Klein’s Nieuw-Guine´e appeared. The use of radio and aircraft became standard, as elsewhere, and aerial photography was also strongly promoted. In 1936 A. H. Colijn, with J. J. Dozy and pilot F. J. Wissel, made a largely successful general survey of the Carstensz Mts complex (now Mt Jaya), reaching the top of Ngga Pulu (dense fog obscuring the Carstensz Pyramid), although much would remain for Harrer and Temple in 1961–1962. A valuable small plant collection was secured by Wissel (Leiden, Bogor). In 1938 Wissel made another discovery: the three Wissel (now Paniai) Lakes. The following year an aerially supplied

................. 16157$

$CH2

02-08-07 10:22:25

PS

PAGE 60

Biological Exploration of New Guinea / 61

government station, Enarotali, was opened; this inspired two more large expeditions even while the Third Archbold was in the field. The Royal Netherlands Geographical Society’s ‘‘Le Roux’’ Expedition of 1939, led by Stirling Expedition veteran C. C. F. M. le Roux with zoologist Prof. H. Boschma, collected insects and other animals in the Nassau Mts, Wissel (now Paniai) Lakes, and Etna Bay (Leiden). For at least part of the time they were accompanied by botanist P. J. Eyma and his assistant, E. Loupattij, who—over nearly a year (December 1938–November 1939, partly on their own account)— made extensive plant collections throughout the area (Bogor, Leiden; parts of the field data were, however, lost). Afterwards, explorer and controleur J. P. K. van Eechoud—one of two ‘‘Bapa Papua’’ (papa Papua)—collected birds, insects, and other animals during 1939–1940 in the Wissel (now Paniai) Lakes area as well as in the Mamberamo basin and the Van Rees Mts (Leiden).

zoology and entomology Reference has already been made to the major expeditions and cruises, from some of which there were copious zoological results. But, as I have indicated, it became more possible and efficient to sponsor individuals with specific aims, as Walter Rothschild had already done for some time before the war with Albert Meek in British New Guinea/Papua. From the United Kingdom, Rothschild continued—albeit on a smaller scale— his sponsorship, with collections going to his museum at Tring. Meek’s associates and successors, the Eichhorns, continued in Papua until 1923 and then worked in New Britain until 1925. Unfortunately, by 1932 personal circumstances forced Rothschild to sell most of his bird collection to the American Museum of Natural History in New York—five years before his death. Before then, however, he was able partly to support the expedition of Ernst Mayr (see below) and his subsequent studies, and in 1928 also to initiate sponsorship for Fred Shaw Mayer, so launching a career which would last nearly forty years (the present author met him in 1966) with stays in New Guinea of varying duration. The considerable collections of birds by Bu¨rgers on the German Sepik Expedition of 1912–1913, worked up, as already noted, by Stresemann in Berlin and published in 1923, filled in many gaps but a number of questions remained— notably concerning some ‘‘mystery’’ birds of paradise and bower birds collected from time to time by plume hunters before 1920 but not seen since. This led to joint American, British, and German sponsorship for a medical student turned zoologist, the future evolutionist, prolific writer, and ultimate centenarian Ernst Mayr (1904–2005). While still under supervision by Stresemann, Mayr traveled both to western and eastern New Guinea in 1928–1929, collecting many birds (Tring and AMNH) and mammals along with insects (Berlin) and some plants (partly lost, with duplicates in Bogor), ranging from the Vogelkop Peninsula (Siwi, Momu, and Anggi Lakes) to the Cyclops Mts (to the summit), the Huon Peninsula (including the Saruwaged Range, following in Keysser’s and Lane-Poole’s tracks to the summit), and the Herzog Mts south of Lae, crossing into the upper Snake

................. 16157$

$CH2

02-08-07 10:22:25

PS

PAGE 61

62 / david g. f r o d i n

Valley. Mayr’s results led in 1930 to a significant suggestion by Stresemann—that the ‘‘mystery’’ birds were of hybrid origin. Shaw Mayer’s early trips were for the Tring Museum, collecting birds and mammals. Contemporary localities included the Vogelkop Peninsula (1928), the Weyland Mts (1930), the Huon Peninsula (1931), and the Milne Bay Islands (1935). Later he amassed the first major collections from the Central Highlands of present-day Papua New Guinea as these became accessible (BMNH), an activity continuing after World War II. A number of other zoological collectors also were active in the 1920s and 1930s—again with American sponsorship now more in evidence. During the years to 1928, Goodfellow once more was in the field, in 1925 searching for vertebrates in southern Papua; T. Jackson was active around Merauke in 1920–1924 (birds, MCZ); while Wirz, in addition to his already-mentioned sojourn in the central mountains, visited some coastal areas in the early 1920s including swampy Frederik Hendrik (now Yos Sudarso) Island as well as Merauke in the south (animals, Leiden). Wirz, later established at Basel University, would undertake further, largely anthropological, trips over the next three decades, passing away in 1955 in the Maprik area. P. T. Putnam in 1927 collected amphibians and reptiles in the Merauke area (MCZ). L. S. Crandall and H. Hamlin in 1928 collected birds in the southeastern mountains (AMNH). From 1928 to 1933, in addition to Mayr (see above) zoological collectors included C. T. MacNamara in 1928–1930 on Mt Lamington (southeast of Popondetta; Mt Lamington later underwent a Pele´ean eruption which in 1951 killed 3,000 people and destroyed vegetation over a considerable area), focusing on beetles (Sydney, Adelaide); the Rev. L. Wagner in 1929, collecting beetles at Lutheran stations at Finschhafen, Wareo, and Komba, and in the Cromwell Range (Adelaide); W. G. N. van der Steen in the same year in the upper Digul River (insects, Amsterdam); J. T. Zimmer (see above) over 1929–1931, collecting birds on the Fly River (AMNH); W. J. C. Frost, obtaining in 1930 birds on the Vogelkop Peninsula and some of the western Papuan islands (Batanta, Waigeo, and Salawati); Dr and Mrs G. Stein in 1931 in the Vogelkop Peninsula, Weyland Mts, and Yapen (birds and some plants, Berlin and Bogor); S. L. Brug in 1932 on the southwest coast and in the Aru Islands (mosquitoes, Amsterdam, BMNH); and Herbert Stevens in 1932–1933, obtaining birds, insects, herpetofauna, and a few mammals (MCZ) in the upper Watut basin—the first to do so in this famous gold mining region (visiting among other places Wau, Mt Missim, Bulolo, and Bulowat). On the marine front, W. J. Eyerdam collected shells and corals (AMNH). In the Territory of Papua after 1935, Ivan Champion and C. T. J. Adamson, inspired by the concurrent Archbold Expeditions (by whom some aerial support was provided), collected animals on their 1936–1937 Bamu-Purari patrol through the Southern Highlands (including Lake Kutubu, first discovered in 1936). In 1938 there was a small Papuan-Australian Expedition (BMNH). Returning to Dutch New Guinea, E. Jacobson was active in 1936 on Waigeo (around the same time as was Cheeseman; see below), making collections of birds

................. 16157$

$CH2

02-08-07 10:22:26

PS

PAGE 62

Biological Exploration of New Guinea / 63

and insects (Bogor); and, while on the Denison-Crockett expedition in the schooner Chiva to the Vogelkop Peninsula and Raja Ampat Islands in 1937–1938, S. Dillon Ripley (a future Secretary of the Smithsonian Institution) collected birds in the Tamrau Mts and on Salawati, Batanta, and Misool (Philadelphia). Ripley later wrote a popular account of his trip (Trail of the Money Bird, 1942), including observations of Rabaul not long after its Pompeiianesque devastation. The 1930s also saw a renewal of independent entomological collecting in New Guinea, beginning in 1933 with the intrepid Englishwoman Evelyn Lucy Cheeseman, who over her several trips benefited from sponsorship by the British Museum (Natural History). She was the first extensively to sample for insects in many areas. Her adventures—including unwelcome obstructions on the part of Australian officialdom as well as long coastal journeys on foot—are related in a number of entertaining books: The Two Roads of Papua (on her 1933–1934 trip), Six-legged Snakes (on her 1936 tour, partly with W. Stu¨ber; see below), and Land of the Red Bird (on her 1938–1939 trip). In 1933–1934 Cheeseman collected insects at Kokoda, Orrori, and Oquali (on the north side of Owen Stanley Mts), Isurava (900 m), and at Mafulu (1,200 m), Mondo (1,500 m), Dieni (600 m), Mt Tafa (2,550 m), all on the southern fall of the Owen Stanley Mts. During part of that period she worked with the First Archbold Expedition (see above). In 1936 she collected briefly at Kavieng, New Ireland, but much more substantially around Hollandia (now Jayapura) with visits to the Bougainville Range (on the border) and particularly in the Cyclops Mts. In 1938– 1939 she worked on Waigeo Island and on Yapen, from there proceeding to Hollandia (and Humboldt Bay), then working in the Mandated Territory at Aitape (having largely walked from Hollandia!), the Torricelli Mts, the hills between Vanimo and Hollandia, and the Bewani Mts. Many species have been based on her collections (insects BMNH; 1939 collections from the Mandated Territory at Adelaide; plants BMNH, Kew, including cryptogams from 1936). Other entomological collectors worked on the mainland of western New Guinea, the Raja Ampat Islands, the Geelvink (now Cenderawasih) Bay islands, and elsewhere under Dutch control in this decade. These collectors included Jacobson (see above); Lt J. M. van Ravenswaay Claasen (Berau Peninsula, Vogelkop, in 1937, and Mappia, the Digul River, Merauke, and Ayamaru (insects; Leiden) in 1938); R. G. Wind (butterflies and other insects in 1939 along the south coast including Fakfak, Merauke, etc., sold to various museums); and, from 1930, the settler and professional collector W. Stu¨ber (in 1936 with Cheeseman) in the Hollandia (now Jayapura)-Sawia area and hills to the south and east, including Sentani, Krisa Road, Korime, Mamda, Ajiop, Tarafia (600 m), Kofio (Komfe) Hills, Vokwar, Njau, Bougainville Mts (400 m), Bewani Mts, Nonno (Japoe), Cyclops Mts, and the Pim River (insects, Bogor, Leiden, with particular attention to Odonata); he also collected orchids (see below). In the Mandated Territory, a new center for entomological activity was established at Kerevat outside Rabaul in 1928 when the Department of Agriculture set up an experiment station. There, J. L. Froggatt, B. A. O’Connor, and Gordon Dun

................. 16157$

$CH2

02-08-07 10:22:26

PS

PAGE 63

64 / david g. f r o d i n

were entomologists, but their collections are now mostly destroyed. Also in New Britain, G. F. Hill collected mosquitos and other insects (Macleay; CSIRO). But that big island would, beyond the Gazelle Peninsula, remain largely unknown until after World War II; nor was there much activity in the rest of the Bismarck Archipelago. In September 1939, World War II broke out and in May 1940, the Netherlands was overrun by Germany. In the two years before war arrived in the Pacific there was a final flurry of Dutch activities; these in particular improved knowledge of the poorly known Bomberai Peninsula. The Negumy Expedition in 1941 included insect collecting (Bogor, Leiden) by the forester E. Lundquist (see also below) on the Vogelkop, McCluer Gulf, Agonda, Bomberai (or Onin) Peninsula, Etna Bay, Oeta (or Uta), and Najeju (south coast). The Indonesian official (mantri) Anta from Bogor collected on the Digul in 1941 while with Wentholt (see below). J. J. van der Starre collected insects at Kaimana (southswest) in 1941 (Leiden).

plants In Dutch New Guinea, in 1920–1921 H. J. Lam—followed in 1926 by W. M. Docters van Leeuwen—made substantial plant collections on their respective expeditions (see above), Lam reaching the summit of Mt Doorman. Otherwise, new botanical activity in the Dutch districts was relatively limited, though the writing up of earlier collections continued. Among the few other contributors were some primarily engaged in zoological work (see above), notably Mayr in the southern Arfak Mts and Cyclops Mts in 1928–1929 (Berlin, Bogor, Harvard), and the Steins in 1931 (Berlin, Bogor). In the early 1930s both Cheeseman and Stu¨ber (see above) collected some plants, Cheeseman in Australian as well as Dutch territory (BMNH, Kew). Stu¨ber focused particularly on orchids for commerce, finding among many others the ‘‘Sepik Blue,’’ Dendrobium lasianthera J.J.Sm. (1932), while Cheeseman obtained, among others, mosses, ferns and grasses. In Australian Papua, collecting early resumed with a visit in 1918 by the newly appointed Queensland Government Botanist C. T. White (who was also consultant botanist to the territory, in succession to Bailey), on invitation by Lt Governor Murray (Smith, his semi-independent senior civil servant and nemesis (see above), being away in Europe) and taking advantage of vacation leave. Several hundred numbers were collected (Brisbane, BMNH), all from the then-Central Division, and a report and collection list published. White was relatively soon followed in both Papua and the Mandated Territory by the chief forestry officer for the Commonwealth of Australia, C. E. Lane-Poole (who would some forty years later open the main buildings of the Forestry School at Bulolo). In 1922–1924, as part of a forest assessment survey (see below), LanePoole collected extensively in areas with relatively easy access, yielding some hundreds of numbers (Brisbane, Kew). His results were the first of any significance in the former German territory since 1914, but would be the last under official auspices in both territories for most of the remaining years between World War I and World War II.

................. 16157$

$CH2

02-08-07 10:22:27

PS

PAGE 64

Biological Exploration of New Guinea / 65

In 1925–1926 Brass (see Archbold Expeditions, above), like White a Queenslander, also came to Papua, but under private sponsorship. His patron was C. S. Sargent—director since 1872 of the Arnold Arboretum of Harvard University. Thus began an association over some three decades of that institution with New Guinea. Brass collected extensively in the Gulf, Eastern, and Central Divisions, in the latter reaching the Owen Stanley Range watershed. 1,165 numbers were obtained (Harvard Herbaria). A second expedition, planned for 1929, unfortunately came to naught because of Sargent’s death; but in that year White reported in the Arnold Arboretum’s Journal on the majority of Brass’s collections. Later individual collectors included two later-famous British anthropologists. One, Gregory Bateson, a son of the geneticist William Bateson, author of Naven and other books, and once Margaret Mead’s husband, collected in 1931 in the Baining areas of New Britain and also in the Sepik basin (Kew). Another, Beatrice Blackwood, collected in 1936–1938 in the upper Watut Valley (Morobe) and, in 1937, when excluded from the Watut, along the south coast of New Britain including Kandrian and Arawe Islands (Kew). Also paying visits were a British gentleman-traveler, A. H. Batten Pooll (1940, Central Division; collections, Sydney); and the Japanese botanists R. Kanehira and S. Hatusima collecting in 1937 in Morobe and the Bismarck Archipelago and in 1940 in the Vogelkop Peninsula and from Nabire inland to Dalman in western New Guinea, the latter the site of a copal or dammar (Agathis labilliardieri) gum enterprise (collections, FU, BO and Harvard). The visit by the two Japanese in 1937 was hurried, limited to the schedule of a cruise ship which took in Kavieng, Rabaul, and Salamaua; a swift return flight from nearby Lae enabled a quick visit to Wau. Collections were accordingly relatively few but their results were soon published. Their 1940 collection was, with more than 2,800 numbers, rather more extensive, and gave rise to a considerable series of papers (unfortunately never completed), with the most substantial coverage for the Anggi Lakes after Gibbs. Appointments to the Botanical Gardens at Bogor under Takenoshin Nakai (director in 1943–1945, during World War II) certainly facilitated their research. But—save for Brass from 1933 with Archbold—all these efforts would be far outdone by three indefatigable plantsmen: the British/Malayan plantation manager and orchidophile C. E. Carr and the American missionary couple Joseph and Mary Clemens. Mrs Clemens in particular would over some six years be responsible for more than 10,000 numbers in the Mandated Territory (almost all in Morobe District). Carr’s 1935–1936 contribution, from a still poorly-known part of the Owen Stanley Range (in spite of earlier visits by Macgregor and others), was perhaps equally important, his over 5,000 numbers being better prepared and with more duplicates than Mrs Clemens’. Both undertakings resulted in very thorough sampling of their respective areas. Mrs Clemens first came to Finschhafen with her husband, a retired U.S. Army chaplain, in 1935, after work in the Philippines, Indochina, and Borneo. From then on they worked (after his death late in 1936 she continued on her own)

................. 16157$

$CH2

02-08-07 10:22:27

PS

PAGE 65

66 / david g. f r o d i n

extensively until being hurriedly evacuated in January 1942, collecting perhaps 14,000 numbers of plants. Stations included Malolo (near Salamaua), Wau, Lae, Kaiapit, Wantoat, Boana, Matap, Samanzing, Amieng, Mt Sarawaket, Sambanga (above present Kabwum), Ogeramnang (also visited by Mayr in 1929), Yunzaing, Sattelberg, Wareo, and Quembung, many of them Lutheran mission stations. Collections went until 1939 to Berlin (partly destroyed but duplicates elsewhere), and afterwards (until 1941) to University of Michigan Herbarium (MICH). With the Japanese invasion, all transport was cut off and some collections had to be abandoned at Boana or Finschhafen. The plants were said to have been destroyed but in the 1960s bundles of specimens were uncovered at the two major herbaria in Tokyo (Tokyo University (TI), National Science Museum (TNS)). Carr, sponsored in part by the British Museum (Natural History), collected plants extensively in the Port Moresby region and on the north and south flanks of the Owen Stanley Mts around Mt Victoria (though not reaching the summit). Localities included Kanosia, Koitaki, Boridi (all in present Central Province), ‘‘The Gap,’’ Alola, Lala River, Isurava Mts, Yodda Creek, and Kokoda (Oro Province). Over 5,500 non-orchid and some 1,000 orchid collections were obtained (BMNH, Singapore; many duplicates elsewhere, especially Canberra, Leiden). Both collections were reported on in similar fashion, mainly either by Diels and others in the later installments of the Beitra¨ge, or by Merrill, Perry, and their collaborators through the Archbold Reports (or elsewhere), but never as a whole; there simply were too many, particularly when all those obtained by Brass on the Archbold Expeditions were also flowing into botanical institutions. Moreover, some of Carr’s collections were not distributed until after World War II; his death in the field meant that this task fell to others. Even today, not all numbers have been fully documented. Some of the Clemens collections moved to Japan were published after 1950 in Japanese outlets. Among residents in both eastern territories, the most outstanding botanist was Fr Gerhard Peekel, who continued his collecting in New Ireland from stations at Lemakot and Ugana. From Ugana he partly ascended the Lelet Plateau (1,000 m), a highland pocket not, however, collected for plants until after World War II—and where, in contrast to highland New Britain, Nothofagus is absent. With advancing years, he also focused much of his attention on compiling his valuable Illustrierte Flora des Bismarck-Archipels fu¨r Naturfreunde, which he saved from destruction under his robes when fleeing combat in 1942, and completed in 1947. The manuscript was after his death deposited in his order’s mother house in Steyl (Germany) and later microfilmed. (A full translation was prepared by E. E. Henty at Lae in the 1970s and 1980s and published in 1985 as Flora of the Bismarck Archipelago for Naturalists; but it is by no means complete for the region.) Others also collected, including two ministers (R. Lister Turner and A. H. Lambton, both in Papua) and, later, a schoolmaster (J. H. L. Waterhouse, in Bougainville and northeast New Britain; K and MAD/WIS); however, Waterhouse’s New Britain collections are relatively few compared with those made in the Solomons. By contrast with all this non-official effort, government activity in the Mandated

................. 16157$

$CH2

02-08-07 10:22:28

PS

PAGE 66

Biological Exploration of New Guinea / 67

Territory after Lane-Poole’s visit remained, as already noted, negligible, rising only slightly from the mid-1930s. A herbarium was begun around 1934 at the Rabaul Botanic Garden (under the Department of Agriculture) but activity was limited and the small collection apparently was lost (a few non-forest tree duplicates survive at Kew and elsewhere). Plants (particularly forest trees) were also collected by J. B. McAdam (see also below) from his appointment in 1938 as a Forestry Officer through 1941, both in New Britain (there largely the work of J. L. d’Espeissis, another forestry officer) and around Wau (CANB, BRI), and by the entomologist J. L. Froggatt (BRI). But, as with insects (see above), outside the Gazelle Peninsula, New Britain would remain floristically almost unknown until the 1950s—and even now, large areas are still poorly explored including much of the south and the mountains.

‘‘economic’’ exploration A number of ‘‘economic’’ expeditions and surveys were also made between 1918 and 1942, not always with significant botanical or zoological collections. The majority were in the 1920s or onwards from 1935, run as a result of reconnaissance requirements or inspired by more favorable business conditions. After the League of Nations mandate was granted for the Territory of New Guinea in 1921, the Australian authorities organized some exploratory surveys. A geological-geographical reconnaissance was made on the Ramu River around 1922, while from 1922 to 1924 Lane-Poole undertook his already-mentioned extensive survey of more accessible forests (following relatively superficial pre–World War I surveys in German New Guinea and the Territory of Papua). Lane-Poole’s well-illustrated 1925 report, which includes a botanical section as well as observations on the forests and vegetation, remains a classic. Key areas visited included the Central, Northern, and Gulf provinces in southeastern New Guinea and in the Huon Peninsula, Ramu basin, and parts of the islands in the Mandated Territory. Of other ‘‘economic’’ undertakings, one of the more important—and also a pioneer with ‘‘flying machines’’—was a major new search for sugar cane (Saccharum) germplasm led by E. W. Brandes in 1928 for the United States Department of Agriculture. His associates included the Dutch botanist J. Jeswiet (Wageningen Agricultural College, the Netherlands), a sugar expert who also made general plant collections (WAG); C. E. Pemberton (of HSPA, Hawai’i), collecting insect pests of cane (Bishop Museum); and pilot R. K. Peck (a Stirling Expedition veteran). With their amphibious plane (furnished by a Chicago businessman), they ranged widely (including the Sepik basin) but did much of their work in the Fly River basin (Lake Daviumbu and the Fly, Strickland, and Oriomo Rivers), with Jeswiet also collecting in the Port Moresby region. The some 130 lots obtained were aimed particularly at improvement of the crop in the southern United States, but replicates also were deposited in Australia. In the 1930s, with new political developments as well as improved business and economic conditions, further ‘‘economic’’ biotic exploration took place. This

................. 16157$

$CH2

02-08-07 10:22:28

PS

PAGE 67

68 / david g. f r o d i n

included renewed attention to forest resources, but relatively little else; extensive land surveys were in the future. In western New Guinea, the first significant forest surveys were carried out at this time, particularly in areas fairly readily accessible by sea. First in the field was Z. Salverda, active over several months in 1936–1937 in the McCluer Gulf (now Bintuni Bay), Bomberai, and along the southwest coast. In 1939 Salverda was followed by L. J. van Dijk (with assistance from Bogor officials (mantris) Ae¨t and Idjan), based for five months at Manokwari. From there he made tours to Yapen, Biak, and (nearer Manokwari) Mios Num. Some 1,600 numbers, primarily of forest trees, eventuated from these two undertakings (Bogor). In 1939–1940 van Eechoud—in connection with the already-mentioned Dutch expeditions to Enarotali—collected some forest trees at the request of van Dijk, mainly near the Mamberamo (Bogor). In 1941 E. Lundquist (see also above) examined more closely some of the areas explored by Salverda; he was accompanied by Ae¨t (collections, Bogor). Land evaluation was also taken up. A pedologist, F. A. Wentholt, collected on three occasions in connection with agricultural surveys—part of proposed transmigration projects as well as other potential development. On part of his last tour of duty (1940–1941), Wentholt was accompanied in the Merauke region and on the Digul River by Anta (see above; collections, Bogor). In 1938 a forest service was established in the Mandated Territory, and, as already indicated, J. B. McAdam and J. L. d’Espeissis were engaged as its first officers; their collections are covered above under ‘‘Plants’’ (above). Both were involved in forest surveys and the establishment of plantations, including, for example, hoop pine (Araucaria cunninghamii) near Wau (pers. obs.). McAdam was to return after 1942 as officer-in-charge of the Australian forestry companies (World War II section, below) and, after World War II, as head of the Department of Forests in the Territory of Papua and New Guinea (TPNG; see Post–World War II Era section, below). As for other plants and trees, one of the few long-term legacies of the limited official effort was the collection (for the United States Department of Agriculture, as part of citrologist Walter Swingle’s comprehensive research) of germplasm and reference material of a close citrus relative, Clymenia polyandra (Tan.) Swingle (now Citrus polyandra Tan.). This tree was previously discovered by Peekel in New Ireland but not then definitely known elsewhere; it has since been found in New Britain and on the New Guinea mainland. The stock has been used towards improvement of citrus plantations in the United States and other countries. The outbreak of World War II in Asia and the Pacific in December 1941 ended all official efforts as well as activities in the private sector. By this time, however, in ‘‘metropolitan’’ countries enough collections had been accumulated from most of Papuasia to furnish sketchy but useful knowledge of the biota, particularly the higher plant flora and vegetation, and the mammal, bird, and butterfly fauna. But sensibility still was largely static and would largely remain so for some time before the biota could begin to yield more secrets in relation to dynamic concepts of

................. 16157$

$CH2

02-08-07 10:22:28

PS

PAGE 68

Biological Exploration of New Guinea / 69

biology, ecology, and biogeography. That would require much more field time and sampling—tasks which would be energetically pursued after 1945 (see the Post-World War II Era section below).

World War II (1941–1945) The onset of World War II in the Pacific soon brought much of New Guinea under Japanese control, Rabaul being surrendered in January 1942. Much information had already been gathered in advance, including from those Japanese naturalists who had visited in the preceding decade. All (or most) established local collections—particularly at Rabaul and Kerevat—were abandoned and, in time, destroyed, if not evacuated to Japan. Collecting, however, did not end with Japanese occupation or both sides’ military operations. With the duration and extent of both, not only was there a considerable demand for biological knowledge but, in addition, many individual servicemen would collect organisms on their own account. A wide range of areas as well as biota were thus sampled. Some lots did not survive the war or were otherwise lost, but several important collections have been preserved through deposit.

japanese contributors Some insect and plant collections made by Japanese naturalists just before the war, or transported to Japan following their occupation, have been noted in the preceding section. Most lots were obtained in western New Guinea where from early 1942 until 1944 there was comparatively little military action. Circumstances further resulted in a focus on the Vogelkop and neighboring areas (including Jazira Doberai, Bomberai, and the Wandammen Peninsula). The most considerable contributions were in entomology. M. Satake studied natural history in general in 1942 and 1943, particularly in the Wandammen Peninsula and the adjacent Bird’s Neck, later publishing a book in Japanese (1963, Tokyo). Dr S. Issiki collected insects in western New Guinea including the Vogelkop (Windesi, Majosi, and, in the Wandammen Peninsula, Wasior) and on Rumberpon Island (Taipei; partly reported upon by Gressitt). Professor Toyohi Okada was at Aitape as a soldier, presumably collecting insects. Yoko-oji collected birds at Manokwari, 1942–1944 (Tokyo; all but seven destroyed). The main botanical field contribution was by Takasi Tuyama (1943, Vogelkop and Yapen; bryophytes and other plants, Tokyo) but of lasting value has been the already-mentioned series of papers on the 1940 collections by R. Kanehira and S. Hatusima—many worked up and published during this period, with others perhaps elaborated. The two men’s retreat from Bogor, Indonesia, in 1945, where both had been active for some three years, as well as the surrender of Formosa (where Kanehira had been a professor in forest botany) and adverse conditions in Japan unfortunately brought a premature end to this project. Some further novel-

................. 16157$

$CH2

02-08-07 10:22:29

PS

PAGE 69

70 / david g. f r o d i n

ties based on these collections were, however, described by others after World War II, and a series index by P. van Royen appeared in 1983.

western allies contributors As with the Japanese, Allied servicemen made relatively few contributions in vertebrate zoology; for both, there were naturally considerable obstacles. The Harvard zoologist and biogeographer P. J. Darlington, Jr., collected many carabid beetles, ants, frogs, and other animals, primarily at Dobodura near Buna (Oro Province, PNG) but also—as military operations progressed—at Milne Bay, Aitape, and Sansapor (in the Vogelkop). He also collected on the slopes of Mt Wilhelm (Simbu Province, PNG) while on leave after being mauled by a crocodile in the lowlands (MCZ). J. Frank Cassel collected birds and herps (amphibians and reptiles) at Finschhafen in 1944 (Cornell). L. W. Jarcho, C. W. Moren, W. M. Beek, G. H. Penn, A. M. Keefe, and W. H. Stickel collected amphibians and reptiles along the north coast, mostly in 1944 (MCZ, USNM); and by Melvin Kurz (AMNH). The contributions of Harry Hoogstraal as well as W. V. King et al. are mentioned below in the context of entomological work, while those of D. F. Grether appear in connection with the activities of plant collectors. Entomological contributions—particularly from the United States and Australia—were extensive, often with results of lasting value through publication, either by themselves or by others. Early papers came particularly from J. N. Belkin, R. M. Bohart, Joanna Bonne-Wepster, Robert Domrow, D. S. Farner, J. L. Gressitt, D. J. Lee, Elizabeth Marks, C. B. Philip, Alan Stone, F. H. Taylor, and Herbert Womersley. Many others appeared in subsequent decades. Among servicemen from North America, K. V. Krombein collected Hymenoptera and other insects, primarily at Nadzab, Markham Valley, 1944 (USNM). E. S. Ross and S. G. Jewett collected all groups of insects at Finschhafen, Hollandia, Maffin Bay, or elsewhere, mostly in 1944 (CAS). Borys Malkin collected mostly beetles, from several areas (USNM). Harry Hoogstraal collected generally as well as mosquitoes and ectoparasites in the Cyclops Mts (right up to the summit) and elsewhere, 1944–1945 (CAS, FMNH, USNM); unfortunately, much of his collection was lost. Willard V. King, W. E. Brewer, H. W. Cook, J. Forbes, W. R. Fullen, D. P. Furman, Donald R. Johnson, W. T. Nailon, George H. Penn (see above), L. W. Saylor, C. J. Steinhauer, and J. P. Toffaleti collected many mosquitoes along north coast, 1944 (USNM and CSIRO). Carl Mohr and W. D. Fitzwater collected mites, ticks, and other medical arthropods in the Buna-Gona area, and at Owi Island and Sansapor in 1944–1945 (USNM). Glen Kohls and Cornelius B. Philip collected chiggers (ticks) and other arthropods, etc., at Dobodura, Purdy Island, or elsewhere, 1943–1944 (USNM). Kenneth L. Knight and Lloyd E. Rozeboom collected mosquitoes in several northern coastal areas and islands, 1944–1945 (USNM). Grether collected butterflies with W. H. Wagner, Jr. (USNM), but their main work was in the Bismarck Archipelago (see below). Many Australian and New Zealand servicemen also made insect and other invertebrate collections. Mosquitoes, chiggers (ticks), and other arthropods of medi-

................. 16157$

$CH2

02-08-07 10:22:29

PS

PAGE 70

Biological Exploration of New Guinea / 71

cal interest were collected in northeastern Papua (PNG) and elsewhere from 1943 through 1945 by Frank H. Taylor (who also was stationed at Wewak), Anthony R. Woodhill, Carl Gunther, Ian M. Mackerras (later an author in, and editor of, the definitive Insects of Australia), D. O. Atherton, D. A. C. Cameron, F. Chippendale, D. H. Colless, H. A. Grandall, R. N. McCullock, M. H. Wallace, and R. Harry Wharton (Queensland Museum, Brisbane; Macleay Museum (Sydney University); and the Division of Entomology, CSIRO, Canberra). Another contemporary collector was Carl Gunther, who collected mites, etc., at Bulolo both before and after World War II (Queensland Museum). In the major islands to the east and northeast of New Guinea—essentially a single theatre of war—L. J. Dumbleton collected mosquitoes, etc. on Nissan Island (CSIRO and Nelson), while in 1945 Marshall Laird studied mosquitoes and parasites at Jacquinot Bay, New Britain (Nelson). In the Solomon Islands George E. Bohart, J. M. Fritts, Ashley B. Gurney, Paul Hurd, L. A. Posekany, Barnard V. Travis, and George W. Wharton (USNM), E. Eldon Beck, E. Reimschu¨ssel, Harry P. Chandler, and Dorald Taylor (BYU), and others collected—with some of them (including Gurney) also active on Bougainville Island and on Reimschu¨ssel in the Admiralty Islands. The Admiralty Islands (especially Manus, with its great naval base) became notable particularly for the work of W. H. Wagner, Jr., and D. F. Grether in pteridophytes (UC), with Grether also collecting butterflies (USNM). Both of them (as well as A. H. Dark) subsequently reported on their collections. Botanical work by servicemen was understandably impeded by the bulkiness of vascular plant collections, and major contributions—apart from the ‘‘New Guinea Forces’’ series described below—were few. Perhaps the most considerable contribution among North Americans was, as already noted, that of Wagner (with Grether) in the Admiralty Islands. Other significant lots—largely comprising grasses—came from Lee Burcham (USNM) and John R. Reeder (A). In 1942–1943 Carl de Zeeuw (later at the College of Forestry at Syracuse University, New York) collected plants (particularly large forest trees) in various parts of Papua, with a particular interest in their wood (vouchers in MEL). The Australian botanist N. A. Wakefield (from Victoria) collected plants (principally pteridophytes) in various parts of eastern New Guinea and the Solomon Islands, 1943–1945 (MEL, BMNH). H. S. McKee (originally from Northern Ireland, but later at Sydney University and ultimately resident in New Caledonia) collected plants in 1944–1945 in various parts of northeastern New Guinea (Brisbane). These generally small individual contributions were, however, offset for forest trees by a fortunate combination of place and people. The effective re-occupation of Lae, Nadzab, and other airfields in present-day Morobe Province provided a mainland base for subsequent military and other operations (including relief of the isolated Highlands, which had remained under Australian administration). Early on C. T. White and J. B. McAdam (both with considerable pre-war experience—see under the section covering the years between World War I and World War II, above) convinced the authorities of the need for a better understanding of the tree flora and the uses and properties of individual species, which was at that

................. 16157$

$CH2

02-08-07 10:22:29

PS

PAGE 71

72 / david g. f r o d i n

time still quite patchy. In 1944 the New Guinea Forces (NGF) series of collections was begun at Butibum near Lae under the Australian Forces forestry unit directed by McAdam. Over 2,000 numbers of collections were made before cessation of operations in the latter part of 1945 (BRI, with replicates in LAE and elsewhere). In 1946 the series was resumed by the Department of Forests, Territory of Papua New Guinea (TPNG; see section on the Flora of Eastern New Guinea, below). A number of novelties would in the ensuing years be described from these 1944– 1945 collections.

The Post–World War II Era (since 1945)

integrated expeditions and surveys (since 1945) The disruptions to most developed countries as a result of World War II were such that it would be several years before substantial expeditions again entered the field. Conditions on the ground in most of New Guinea were also difficult—the war had destroyed or severely damaged infrastructure in most coastal settlements. Papua was also affected by political changes in the Dutch Indies, including the emergence of Indonesia as a state. In the settlement of 1949 the Dutch contrived to retain control of Papua. In the Territory of Papua and New Guinea (TPNG)— created in 1945 with a unified administration at Port Moresby—Australia retained control, but reconstruction was at first slow. Developments in the sciences also tended towards greater specialization; at the same time—particularly in the first quarter-century after the war—the sciences enjoyed considerable political favor, with state funds relatively forthcoming. But such expansion could not continue indefinitely; in more recent decades financing has been much harder to obtain, and then only for more targeted, short-term work. Political, economic, and social developments have also been major factors. All this has also had an effect on recruitment into the sciences, including the maintenance of taxonomic expertise, as several recent reports have indicated.

Western New Guinea (West Irian, Irian Jaya, Papua) In what is now Papua, the Dutch also had to be seen to be doing something—even though costs outweighed the (not particularly high) level of return. Economic development in the Territory of Papua and New Guinea (TPNG) was also at first relatively slow but, in the 1960s and with greater external pressure, a firmer political commitment by Australia on the polity’s future had to be made. Before the establishment of Netherlands New Guinea as a separate territory there was one expedition that took in the Raja Ampat Islands as well as the Vogelkop Peninsula. In 1948–1949 an extended ‘‘Swedish-Netherlands Expedition’’ led by Sten Bergman, an ornithologist and natural historian, and accompanied by M. A. Lieftinck, D. R. Pleyte, Sjo¨qvist, and E. Lundquist (for more on Lundquist, see also the section on the years between the World Wars, above) along with Indonesian officials (mantris) Main and Djamhari visited the Raja Ampat Islands (including Misool, Salawati, Batanta, and Waigeo), the Sorong region, and parts

................. 16157$

$CH2

02-08-07 10:22:30

PS

PAGE 72

Biological Exploration of New Guinea / 73

of the southwestern coast. From mid-1949 Bergman spent some months on the eastern side of the Vogelkop Peninsula, including a visit to the Anggi Lakes via Ransiki as well as a visit to the Wandammen Peninsula. Over the course of a year and four months following their arrival, visits at different times were made in the Sorong region (their logistical base) as well as in Raja Ampat. Pleyte and two officials (mantris) focused on Misool as well as around Sorong, but left the field before the end of 1948—possibly in advance of imminent political changes. Apart from Pleyte’s botanical collections, most of the harvest was zoological (birds, Stockholm; insects, Bogor, Leiden) although Bergman did collect some plants around the Anggi Lakes. After 1949 and the advent of a separate Dutch administration, most efforts in biotic exploration were until 1963—with one exception below—individual (or in small specialist teams) or through state bodies such as the Forestry Service (Boswezen). Being largely disciplinary, they are taken up under the sections on Flora and Fauna of Western New Guinea, below. There was only one multidisciplinary ‘‘large’’ expedition in the old style, traversing an extensive area and with geographical as well as natural science objectives— that of April–August 1959 to the Star Mountains. In addition to air support (as in 1938–1939), there now was helicopter transport (though one of the two helicopters was destroyed during operations). Leading the expedition were the zoologist L. D. Brongersma (then also director of the zoological museum at Leiden, now the Naturalis Museum) and G. F. Venema. Participating botanists included C. Kalkman, B. O. van Zanten, and J. J. F. E. de Wilde, while W. Vervoort (as well as Brongersma) collected animals (and, with Kalkman, plants). A naval surgeon, M. O. Tissing, took part in the ascent of the principal objective, the 4,640-meter ice-capped Juliana Top (now Mt Mandala)—there collecting a few plants. Kalkman and van Zanten reached Mt Antares (3,380 m, in the western Star Mountains). From the expedition base in the Sibil Valley (east of the Baliem), where there was already an airstrip, two members (Ba¨r and Danselaar) afterwards pushed north, completing a land crossing—the first on the western side of the border, at the island’s widest point (the east had been crossed by land in 1927). The collections (Leiden) included, among the plants, a good representation of bryophytes (a speciality of van Zanten); these accordingly went first to Groningen. Results appeared in a relaunched Nova Guinea and elsewhere. A popular account by the two leaders is Het Witte Hart van Nieuw-Guinea (undated; in English as To the Mountains of the Stars, 1963). The expedition was a kind of ‘‘finale,’’ and the Dutch knew it. They did, however, thus fill perhaps the last significant ‘‘white spot’’ on the world map (apart from much of Antarctica), the discoveries of Cerro Neblina (Venezuela/Brazil) and in the Vilcambamba (Peru) being just prior. This drama was enhanced by the nearly-simultaneous cross-island trek (also from south to north, but slightly west of the Dutch route) of a French film crew under J.-Y. Gaisseau; this resulted in the much-appreciated The Sky Above, The Mud Below—an experience of most of us who have done any real traveling in Papuasia. International politics and sensibilities—and American pressure—now brought

................. 16157$

$CH2

02-08-07 10:22:30

PS

PAGE 73

74 / david g. f r o d i n

about an end to three-and-a-half centuries of the Dutch in the East Indies. In late 1962 control of Papua passed to a United Nations Temporary Executive Administration (UNTEA); on 1 May 1963 Indonesia took control. The territory was given provincial status and initially named Irian Barat; not long afterwards, this changed to Irian Jaya and remained so until the end of the 1990s. The Dutch administrative seat, Hollandia, was renamed Sukarnopura and, later, Jayapura. Although Indonesian sovereignty awaited final determination by plebiscite, the Act of Free Choice (Pepera: Penentuan Pendapat Rakyat) in 1969, higher education was given priority, and soon Cenderawasih University was built, its main campus at Abepura (outside Jayapura) with agriculture and forestry in Manokwari. For some years afterwards there were no effective outside contacts. Only after 1969 and the Act of Free Choice plebiscite (Pepera: Penentuan Pendapat Rakyat) was there a renewal of visits by scientific teams, mainly from Europe, North America, and Australasia. During 1971–1973 the Australian Universities’ Expeditions—organized at Melbourne University—undertook in two operations glaciological and biological investigations on and around the Mt Jaya (formerly Mt Carstensz) glaciers. Geoffrey S. Hope and Judy A. Peterson (Canberra) were the team biologists. The expeditions’ work was summarized in The Equatorial Glaciers of New Guinea (1976) edited by Hope, Peterson, L. Allison, and U. Radok. In 1972 the former King Leopold III made his second expedition to Papua (the first had been, as already indicated, in 1929 with van Straelen). With J. Raynal (Paris), collecting continued into 1973 near the Mt Jaya area, Baliem Valley, and other places (Brussels, Bogor, Paris, Leiden). In 1974–1976 a multidisciplinary expedition from Germany worked in the vicinity of the upper Eipomek River in the eastern Nassau Range—home to the Eipo, an isolated outlier group of the Mountain Ok of central New Guinea—under the title ‘‘interdisziplina¨re Erforschung von Mensch, Kultur und Umwelt im zentral Hochland von West-Irian (Neuguinea).’’ With support from the Deutschen Forschungsgemeinschaft and organized through the Museum fu¨r Volkerkunde in Berlin (already enriched by Moszkowski’s, the Behrmann expedition’s, and other ethnographic collections), a considerable team of specialists was active over some two years in various parts of the Eipomek River, including its eastern and western tributaries (and surviving two serious earthquakes, locally very destructive); their botanical collections are in Berlin. Their reports have since 1979 appeared in a special series ‘‘Mensch, Kultur und Umwelt in zentralen Bergland von WestNeuguinea’’ (Berlin). The 1980s represented another quiet period, with the next interdisciplinary contributions being collections of papers—only a minority biological—rather than expedition reports, although some reflected recent fieldwork. Both involved the Irian Jaya Studies Programme in the Netherlands, and had as a primary focus the Vogelkop Peninsula. They were Perspectives on the Bird’s Head of Irian Jaya, Indonesia (1997, Rodopi), ed. J. Miedema, C. Ode´, and R. A. C. Dam; and Bird’s Head Approaches (1998, Balkema, as number 15 in their series Modern Quaternary Research in Southeast Asia), ed. G. J. Bartstra. The biological papers are all general

................. 16157$

$CH2

02-08-07 10:22:31

PS

PAGE 74

Biological Exploration of New Guinea / 75

in nature; however, of interest for exploration history is a chapter in Perspectives on the 1907–1915 Dutch Military Expeditions (Militaire Exploratie). By the 1990s and the advent of the Convention on Biological Diversity, however, there was renewed interest in how much—or how little—was known of the biota of New Guinea, both in east and west. For Papua this led notably to a 1997 workshop in Biak, sponsored by Conservation International in Indonesia and the United States; a Laporan Akhir/Final Report appeared in 1999 with a number of maps (and a CD-ROM with several database files) depicting perceived priority areas, for different biotic groups as well as in general. Following the workshop, Conservation International’s Rapid Assessment Program (RAP) became active on the ground. In 1998 fieldwork was carried out on the Wapoga River area of Yapen-Waropen regency (southeast of Waren), an area—without much prior attention—inland from Olifant-berg, one of A.B. Meyer’s 1873 localities. All sites were at 1,100 m or less save one (1,890 m, reached by helicopter), and partly accessible because of previous prospecting in the area by the U.S. mining firm Freeport McMoRan (see also below). The results appeared in RAP’s Report 14. A later Marine RAP was active in the Raja Ampat Islands (Report 22). The Raja Ampat island group now attracted the attention of another U.S. organization, The Nature Conservancy (TNC). In 2002 a team from TNC carried out a several weeks’ survey with the aid of a 75-foot twin outboard-motored speedboat (the Pindito); of plants, 550 numbers were collected (Bogor in first instance). The 15-strong expedition (a third from media) was led by R. Salm (TNC); its zoologists were G. Allen (fish), D. Ivereigh (birds), A. Sumule and E. Turak, and botanists included J. P. Mogea and W. Takeuchi with support also from F. Liuw and D. Neville. Islands visited included Misool, Kofiau, Batanta, and Salawati along with the ultramafic Kawe´—and, near to it, the partly ultramafic Waigeo. A first botanical report appeared in 2003 (Takeuchi in Sida 20: 1093–1116). As a result of these surveys, the very high marine diversity of the Raja Ampat Islands—already partially known due to the Siboga and other earlier oceanographic work—now became effectively recognized, together with their distinctive geological history. At the same time, a much better idea was obtained of the plant and forest cover and its potential survival—more problematic in the drier islands. Yet the effects of long biological neglect were reflected in the number of new records (and even novelties)—some of relatively common taxa. As Takeuchi wrote, such were ‘‘indications of the undercollected status of the limestone, and show how poorly documented this flora still remains even after more than a century of . . . exploration’’ (and over two centuries if the partly unpublished collections of Labilliardie`re and perhaps other French explorers ever are fully accounted for). Parts of the rest of Papua are considered better known botanically (if a 1950 average for the Raja Ampat Islands of 25 collections/100 km2 —with not much more at least until recent years—is deemed satisfactory); but some island groups, let alone individual islands, may still be undercollected. These and other multidisciplinary undertakings—if now more focused (schwer-

................. 16157$

$CH2

02-08-07 10:22:31

PS

PAGE 75

76 / david g. f r o d i n

punktlich) than in the past, being shorter in duration and covering a smaller area—have once more become a main vehicle for natural history work in Papua. There have been relatively few long visits by single workers or small specialist teams, as was the case for van Royen and Sleumer in the years before 1962 or, more recently, for Widjaja, Mangen, and Milliken. Logistics, national sensibilities, and particularly security remain major current concerns. For decades all, or nearly all, of Papua has been a military zone, the armed forces active in parallel with civil administration. Moreover, those researchers in employment are under rather tighter constraints than in the past.

Eastern New Guinea In the Territory of Papua and New Guinea (TPNG)—now under one administration at Port Moresby—a number of multidisciplinary efforts together amassed large amounts of material and covered a number of areas previously poorly known. These include the four Archbold Expeditions (nos. 4–7) of 1953–1964 and the CSIRO Land Research regional surveys of 1953–1970—the latter an extension of similar activities in northern Australia.

Archbold Expeditions The Fourth Archbold New Guinea Expedition worked in eastern Papua between Collingwood Bay and Mt Dayman as well as on Goodenough Island—the last hardly before visited. Brass was leader and collected insects as well as plants. H. M. van Deusen (AMNH) was mammalogist and G. M. M. Tate (AMNH) was general collector; however, after six months Tate was evacuated, fatally ill (animals AMNH; plants Harvard). The Fifth Archbold New Guinea Expedition was a continuation of the Fourth, but focused on an effective survey of many of the remaining Milne Bay islands. They continued in the D’Entrecasteaux group (Fergusson and Normanby islands) and worked also in the Trobriand Islands, Woodlark (where Montrouzier had preceded them over a century before), and the Louisiades (Misima, Tagula (or Sudest), and Rossel) and also visited Milne Bay on the mainland. Brass again was leader and collected plants and some insects, while R. F. Peterson was mammalogist (animals, AMNH; plants, Leiden). The Sixth Archbold New Guinea Expedition focused on the northeastern mainland, taking advantage of the developing road network and other infrastructure to cover a fairly wide area. They worked in the present Morobe, Madang, Eastern Highlands, and Simbu provinces; localities visited included Mt Wilhelm, Mt Otto (Mt Sagueti), Mt Michael, Mt Elandora, around Okaba, Kassam Pass, and the Markham Valley as well as points from Lae to Mt Kaindi and Edie Creek. Again Brass was leader, with van Deusen as mammalogist. Other participants included J. Womersley, J. D. Collins, and T. C. Maa (the latter from the Bishop Museum), Maa remaining for a month during the Kassam Pass and Okaba sojourns making a collection of vertebrate ectoparasites (Bishop Museum; other vertebrates, AMNH; plants, USNM and Lae).

................. 16157$

$CH2

02-08-07 10:22:32

PS

PAGE 76

Biological Exploration of New Guinea / 77

The Seventh Archbold New Guinea Expedition was led by van Deusen and focused on the Huon Peninsula. Participants included S. A. Grierson (as general zoologist and photographer), R. G. Zweifel (herpetology) and R. D. Hoogland (plants). Work was done in the Rawlinson Range, the Cromwell Mountains, and near Finschhafen, with Hoogland also reaching the Saruwaged Mts (animals, AMNH; plants, CANB). Logistical and other support was generally in the hands of local residents under contract (e.g., Collins for the 1959 Archbold Expedition), and, as noted, local professional scientists and others sometimes accompanied these parties. Substantial general reports were published for the expeditions of 1953, 1956, and 1959, but preparation of that from 1964 lagged and was eventually abandoned. Zoological results continued to be published for the most part in the American Museum of Natural History Bulletin, and an ‘‘Archbold Office’’ remained there at least until the late twentieth century (under H. van Deusen and then K. Koopman). But botanical results (J. Arnold Arboretum) persisted only until 1953, by which time there had been a general turn by U.S. tropical botanists to the Americas. Overall, though, the Archbold reports have been, and are, the largest American contribution in the tradition set by other major undertakings, individually or collectively, including the reports of the Challenger, Siboga, and the first Leopold expedition as well as Nova Guinea and Beitra¨ge zur Flora von Papuasien, all with treatments by specialists.

CSIRO Expeditions While the Archbold expeditions remained renowned, other sponsors were not to be outdone. The markedly improving infrastructure of the Territory of Papua and New Guinea (TPNG), including its expanding road and air network, was an attraction. Particularly worthy of note in the post–World War II decades was the work of the Commonwealth Scientific and Industrial Research Organisation of Australia (CSIRO)—hitherto without any presence in New Guinea—and in particular its Land Research Division. Following a tradition first espoused by Linnaeus with his royally sponsored expeditions in Sweden, followed by others of the kind in the nineteenth century, the CSIRO Regional Surveys of 1953–1970—like those in still poorly-known northern Australia—were interdisciplinary. Participants included R. D. Hoogland (through 1966; later with the Taxonomy Unit, Research School of Biological Sciences, Australian National University, Canberra, before returning to Europe), P. Darbyshire, L. Craven, R. Pullen, R. Schodde, P. C. Heyligers, K. Paijmans, A. Kanis, and J. C. Saunders. The teams were active in most of the modern provinces (save in the Bismarck Archipelago), and on all surveys collections of biota were made (CSIRO, Canberra). Fourteen survey reports appeared between 1964 and 1976, with three further syntheses on particular aspects including vegetation (1975) and some books (1976–1983). All the reports and synthesis featured illustrations, diagrams of land systems, and maps. A key aim was assessment for potential agricultural develop-

................. 16157$

$CH2

02-08-07 10:22:32

PS

PAGE 77

78 / david g. f r o d i n

ment; there was no similar mandate for conservation. But because there were no related botanical and zoological series, biotic results are by now widely scattered (though partially synthesized for some groups).

Noona Dan and Alpha Helix Expeditions Two multidisciplinary ocean expeditions visited in the Territory of Papua and New Guinea (TPNG) in the 1960s, the first being the Danish Noona Dan (1962). This had as its general focus the western Pacific. After calls in Palawan and the Sula Archipelago they headed for the Bismarck Archipelago and were there for just over six months (January–June 1962). Stops were made in New Ireland (Kavieng and vicinity, also elsewhere including the already-mentioned Lelet Plateau), Dyaul Island (off the south of New Ireland), Mussau Island (one month), Lavongai (also called New Hanover), Manus Island, the western Admiralties including the Hermit Islands, the outer northeastern atolls, parts of New Britain including Hoskins, the Baining Islands, Blanche Bay (Rabaul and vicinity), and Credner Island, and the Duke of York Islands. They thence proceeded to the Solomon Islands (with a particular interest in Rennell) before returning to Denmark. Botanical collectors included S.-E. Sandermann Olsen, M. E. Køie, H. Dissing, S. F. Christiansen, and T. L. Wolff (Wolff was scientific leader). Several publications resulted. Botanically the Mussau Island call was the most useful—there had been no previous collecting there, the Emirau (also known as Squally) group (in the Bismarck Archipelago), or Tench Island. Perhaps the Mencke incident (under the section on Northeastern New Guinea (1875–1914), above) was a factor in making local relations difficult, but by 1962 Christianity was well established. In all, 19 scientists participated (at different times); substantial collections were made (Copenhagen). Also of comparatively long duration was the R/V Alpha Helix New Guinea Expedition of May–November 1969. Sponsored by the U.S. National Science Foundation and active mostly on or by the northeastern mainland and in the Bismarck Archipelago, it involved scientists from universities and museums in several countries, especially the United States and Australia. Research included studies on physiology and ecology, notably in vertebrates (C. Sibley, R. Zweifel), on bioluminescence in fireflies, etc. (J. P. Buck, J. E. Lloyd), as well as on marine life and fungi. Australian collaborators included J. Calaby, H. Cogger, and R. Schodde (USNM, AMNH, Yale, CSIRO, AM, etc.). (Sibley would later contribute significantly to a recasting of bird phylogeny, partly utilizing evidence from genomic sequences.)

Other Multidisciplinary Undertakings, Including Biological Stations Multidisciplinary efforts over the last four decades of the twentieth century in eastern New Guinea and the Bismarcks have usually had quite limited, more indepth geographical objectives, or have been ethnobiological. Almost all are nonofficial, some longer-term (e.g. the Wildlife Conservation Society (New York) at Crater Mountain and elsewhere, or the PABITRA initiative of ICSU’s ‘‘Diversitas’’ program) and some ad-hoc (including at least one Conservation International

................. 16157$

$CH2

02-08-07 10:22:33

PS

PAGE 78

Biological Exploration of New Guinea / 79

RAP survey as well as work in the Hunstein Range partly sponsored by the National Geographic Society in the United States). Also, a number of local stations have been established. Apart from the stations of the government at Lae, Bulolo, Kanudi, and elsewhere, independently-sponsored stations (including those related to educational institutions) have been established at Wau (from 1961 as the Bishop Museum Field Station, becoming in 1971 the Wau Ecology Institute, or WEI); in Madang Province including the Christensen Research Institute (1980s–1990s), the Leopold III Research Station at Laing Island (1970s onwards, with numerous contributions to its credit and collections in Belgium, PNG, and elsewhere), and the Parataxonomy Center (1990s onwards); Motupore Station in Central Province (for UPNG; 1971 onwards); Ivimka Station by the Lakekamu River in Gulf Province; and Crater Mountain in the Eastern Highlands (1990s onwards). More ‘‘informal’’ sites also exist, usually in association with local communities. Space, however, forbids a detailed account of their activities. (For people and activities at Wau in the 1970s, see Frodin and Gressitt (1982), in the References section, below).

flora of western new guinea and associated islands (since 1945) Nieuw Guinea, Residency of Netherlands East Indies (1945–1949); Netherlands New Guinea (1949–1962); UNTEA (1962–1963) Botanical activity in western New Guinea after 1945 became largely a state undertaking, not unnaturally focusing on the woody flora and other plants of economic interest. Nevertheless, some expeditions, primarily those from the Rijksherbarium (now Leiden branch, National Herbarium of the Netherlands), had a more general remit. In this they were well supported by two successive professors of systematic botany, H. J. Lam (who himself had been in New Guinea with the Kremer expedition in 1920) and, after him, C. G. G. J. van Steenis (with a strong interest in the whole flora of Malesia, but notably that of the mountains). The years prior to establishment of Netherlands New Guinea as a separate polity saw only relatively limited activity. In 1948 A. J. G. H. Kostermans with Weygers continued the forest reconnaissance work in Bomberai and the Vogelkop peninsulas begun before World War II, visiting areas bordering the east coast of the Vogelkop but also collecting in the Namtui Mts (particularly for Cryptocarya massoy, a spice tree then still poorly understood but now known to be in patchy stands around mainland New Guinea) and for 12 days around Anggi Lakes. With the separation of Papua from Indonesia, autochthonous internal services came into being, including the Forestry Service (Boswezen) and Agriculture Service (Landbouw). An agricultural station was initially developed at Kota Nica near Lake Sentani, nearby areas having been the site of a transmigration scheme. The research sections (including forestry) were, however, in a few years moved from there (and Hollandia) to Amban near Manokwari. There, over the period 1953–1962, a research station was developed. Resident botanists in the Forestry Service (Boswezen) included Ch. Versteegh (an associate of Brass during the Third Archbold Expedi-

................. 16157$

$CH2

02-08-07 10:22:34

PS

PAGE 79

80 / david g. f r o d i n

tion in 1938–1939), C. Kalkman, and later W. Vink; they were assisted by Peter and Gerrit Iwanggin, Cris Koster, and F. A. W. Schram. In charge of surveys was Forester J. F. U. Zieck. From a start in 1953, a relatively good representation of the lowland tree flora was collected in the BW-series (Boswezen Nieuw Guinea) during forest assessment surveys; but for reasons of economic accessibility not that much from above 1,000 m was obtained. Associated material including wood samples was also gathered. In the last two years of Dutch rule, however, the forest botanists broadened their collecting, a wide variety of plants being obtained before cessation of activities (Manokwari, Leiden, Bogor, Kew, and elsewhere) with numbers reaching just short of 16,000. Only some have so far found their way into contributions and revisions; however, the replicates at MAN have since been at least partially included in a database. In the Vogelkop Peninsula, the Forestry Service (Boswezen) surveys paid particular attention to the Warsamson Valley (east of Sorong), Sausapor (north coast), the Kebar Valley (Araucaria cunninghamii and Agathis labilliardieri being present), the Arfak Plain and its deltas (west of Manokwari), Oransbari, Momu, and Ransiki (all on the east coast, and earlier visited by Kostermans and Weygers), Tisi and Muturi near Bintuni (at the head of the eponymous gulf), the Ayamaru Lakes (in the center towards Ayawasi), and Beriat (near Teminabuan)—this last with sandstone outcrops and white-sand lands (the latter with a higher-than-usual percentage of Myrtaceae and Dipterocarpaceae). In 1954 Zieck and Versteegh reached the Anggi Lakes, partly to examine stands of Agathis from which (as at Dalman; see above) copal was being extracted and traded. In the Raja Ampat Islands, Kaloal (on Salawati Island) was also surveyed, but otherwise those islands remained botanically neglected. Coverage elsewhere was relatively limited. The first outside botanist was P. van Royen from Leiden with the first Rijksherbarium expedition (1954–1955). Partly with Versteegh (and, for a short time in October 1954, Lam) he explored many areas of the Vogelkop Peninsula as well as Batanta Island in the Raja Ampats; in the south he collected around Merauke and from there to the Fly River; he then visited the Cyclops Mts in the north. In 1955 he worked in Waigeo, obtaining materials for a valuable baseline report (1960). He also visited eastern New Guinea, partly to make formal contacts with the Division of Botany (see below). In 1955 and later Gressitt (see next section) collected a few plants (Bishop). In 1957 C. O. Grassl, on a sugar cane germplasm expedition, collected grasses in lowlands and at Anggi and Wissel (now Paniai) Lakes (Leiden). In 1959 Kalkman participated in the Star Mountains expedition (see section on Integrated Expeditions, above), but made somewhat fewer collections than might have been expected. In 1961 P. van Royen with H. O. Sleumer, comprising the second Rijksherbarium expedition, did valuable work in the Vogelkop Peninsula, visiting the Kebar Valley, Tamrau Mts, the Nettoti Range, and the northern Arfak Mts, and also climbed into the Cyclops Range (Leiden). Some results from the two Rijksherbarium expeditions appeared in Nova Guinea, but no full account of the plants resulted.

................. 16157$

$CH2

02-08-07 10:22:34

PS

PAGE 80

Biological Exploration of New Guinea / 81

Not much collecting was done in the main ranges during the remaining years of Dutch rule. In 1958 Bergman collected plants in the Swart Valley, including the curious Papilionopsis stylidioides Steenis—revealed in 1977 as an artifact: an inflorescence of a legume, Desmodium (now Hylodesmum) repandum inserted into a sterile tuft of a monocot, Burmannia disticha. A few plants were obtained on the 1961 New Zealand expedition to the Carstensz (now Jaya) Mountains by D. E. Cooper and Philip Temple, and then in 1962 by Temple and Heinrich Harrer (Auckland). This latter party was the first to reach the summit of the Pyramid, now known to be the highest point in New Guinea and thus the goal of expeditions for more than fifty years. Major collections from this area were, however, not made until after 1970, and then facilitated by improved access consequent to reconnaissance and establishment of the copper mines at Ertsberg and Grasberg by Freeport Sulfur (now Freeport McMoRan).

Irian Jaya, Later Papua (since 1963) Following the advent of Indonesian administration and in connection with the founding of Cenderawasih University, an agriculture and forestry college was set up in 1964 at Amban near Manokwari, close to the already-mentioned experiment station. This opened up more opportunities for local education in applied biology and related fields. In due course the forest herbarium was transferred to the Manokwari campus of Cenderawasih University, with which it remains associated. However, the forest herbarium suffered from relative neglect until the last decade or so. From the 1990s, however, there have been substantial additions as well as rehabilitation, and MAN is now functioning as the leading botanical collection in Papua. In 1966, plants (Bogor) were collected by W. Soegeng Reksodihardjo (then at Bogor) with Kostermans around Sukarnopura (formerly Hollandia, now Jayapura) and its environs (including Abepura), Lake Sentani, the Cyclops foothills (including Deplanchea glabra) and in the Baliem Valley (including Wamena and Wellesey, ascending to 2,500 m); a visit was also made to Biak Island before return to Java. In 1967 Soegeng, together with the Bogor official (mantri) Nedi, returned to New Guinea as his country’s botanical representative to the IndonesianAustralian border survey party. They collected (Bogor) from April to June at different localities from the foothills of the Star Mountains south to the coast: Ok Walimkan River (Papua), Ingembit (on border), Yat, Angarmaruk, and Weam (PNG), and Bensbach (Papua). In 1968–1970 a Japanese expedition under Y. Kobayashi investigated lower plants in the Wamena area of the Baliem Valley and elsewhere (Tokyo). Vink (see above) made a short return visit in 1968, adding a few more BW-numbers from the Warsamson Valley and elsewhere. Following the plebiscite, the Act of Free Choice (Pepera: Penentuan Pendapat Rakyat) of 1969 with its ostensible confirmation of Indonesian sovereignty, came a renewal of visits from European, American, and Australasian scientists; but at least through the 1970s their botanical work was generally undertaken in association with interdisciplinary undertakings (see section on Integrated Expeditions,

................. 16157$

$CH2

02-08-07 10:22:35

PS

PAGE 81

82 / david g. f r o d i n

above). The Australian Universities’ Expeditions of 1971–1973 obtained a fair number of plants, mostly from G. S. Hope and J. A. Patterson (CANB). These teams were soon followed by J. Raynal, a member of the Leopold III expedition from Belgium (see also the Integrated Expeditions section, above). Raynal collected during 1972–1973 in the Baliem Valley and near Mt Jaya (Bogor, Brussels, Paris, Leiden). (This undertaking later inspired the creation of the Belgian research station on Laing Island; see the Integrated Expeditions section, above.) In 1976 P. Hiepko, W. Schultze-Motel, and W. Schiefenhovel (the latter concerned with medicinal plants) were in the Eipomek Valley (see the Integrated Expeditions section, above). In the 1980s E. Widjaja visited Tembagapura and vicinity, below Mt Jaya. Later she collected in the Vogelkop Peninsula. Also in that decade came the flora and vegetation studies undertaken by J.-M. Mangen in the Jayawijaya Mountains and in particular near Mt Trikora (formerly Mt Wilhelmina). Over three sorties (via the Baliem Valley—three days’ walk to the northeast of his study areas) in 1982– 1984 Mangen carried out extensive topographical, floristic, and vegetation studies (reported upon in French in 1986 and in English in 1993; collections in Luxembourg). It was a continuation, in more detail, of Brass’s 1938–1939 work. Later, Mangen made an expedition to the Valentijn Mountains—a range not otherwise botanically examined, but partly continuous with high mountains near the Yali lands (see below). A different kind of expedition was that made by William Milliken (with Sertu Very Bakaru) in September–October 1992 to the Yali area (northeast of the Baliem Valley and west of the Eipomek Valley). The object was ethnobotanical documentation, and a considerable number of vouchers were obtained (Kew). The work formed part of the International Scientific Support Trust’s ‘‘Expedition to West Papua 1992,’’ and a report was circulated. Soon after in the 1990s came two considerable undertakings, the first at various points in the Vogelkop Peninsula (1994–1995) with support from the John D. and Catherine T. MacArthur Foundation in the United States, the second (1998–2000, locally supported by Freeport McMoRan) in the main ranges along the access road to Tembagapura and at points beyond, including the alpine areas. For both expeditions the major metropolitan participant was the Royal Botanic Gardens, Kew; scientists from Bogor, Manokwari, and Tembagapura also took part. From the work of 1994–1995, some 2,000 collections were made. These collections have not been worked up into a definitive synthetic work (and no plans for one exist) although some background works of limited circulation and interim lists have appeared, including Checklist of the Flowering Plants of N.E. Kepala Burung (Vogelkop), Irian Jaya (1997)—the first synthesis, albeit avowedly provisional, for any part of the Vogelkop Peninsula. The 1998–2000 field campaigns yielded the largest new collections from the Mt Jaya region, exceeding all before. A considerable portion of these new collections (along with, where relevant, earlier records— particularly those of Boden Kloss in 1912–1913) have been a primary basis for a florula, The Alpine and Subalpine Flora of Mount Jaya—covering areas over 3,000

................. 16157$

$CH2

02-08-07 10:22:35

PS

PAGE 82

Biological Exploration of New Guinea / 83

m with a small, somewhat lower area near Tembagapura featuring a subalpine facies. More recently, Kew botanists have collaborated with those at MAN, Lae, and elsewhere in work towards an account of New Guinea palms; in connection with this W. R. Baker visited at points in the Vogelkop and Wandammen Peninsulas. D. Hicks was for periods of time based at MAN as part of a U.K. Darwin Initiative assistance program. Other work has, for logistical, security, and other reasons, been to a considerable extent associated with integrated area surveys (see Integrated Expeditions section, above).

fauna of western new guinea and associated islands (since 1945) Nieuw Guinea, Residency of Netherlands East Indies (1945–1949); Netherlands New Guinea (1949–1962); UNTEA (1962–1963) As in botany, there were a few zoologists who returned to the field in western New Guinea before its separation from the new state of Indonesia. Most notable were S. Bergman and M. A. Lieftinck (see above) in 1948–1949, active in Raja Ampat, Sorong, and the eastern side of the Vogelkop Peninsula. But in the 1950s as civil developments proceeded and western New Guinea entered its last period of relative stability, more sustained field activity got under way. In particular, with economic and human development, entomology came more to the fore than in the past, moving beyond a concern primarily with Coleoptera and the showier Lepidoptera. In 1952, L. D. Brongersma (with W. J. Roosdorp) collected animals in the north, in the Vogelkop Peninsula and at Wissel (now Paniai) Lakes and Merauke (Leiden). In that same year Bergman returned (remaining until 1953) with a further visit in 1958, this last inclusive of the Swart Valley (collections, Stockholm); he also published popular works covering these travels as well as on that of 1948– 1949. In 1954 L. van der Hammen collected Acarina (ticks and mites) etc. in the Vogelkop Peninsula, on Geelvink Bay, the Wissel Lakes, and in the vicinity of Hollandia (now Jayapura; including Lake Sentani). In 1954–1955 Brongersma (with M. Boeseman and L. B. Holthuis) again collected animals (Leiden), taking in many localities in the Vogelkop Peninsula, on the islands in Geelvink (now Cenderawasih) Bay (Yapen, Biak), and along the Digul River. Brongersma also collected animals, with M. Boeseman and L. B. Holthuis, at many localities in the Vogelkop Peninsula, along the Digul River, and on Biak, Yapen, and elsewhere. In 1959 Brongersma returned to lead the scientific team under the Star Mountains expedition (see Integrated Expeditions section, above). In 1960–1961 S. Dillon Ripley (with his wife) came once more to western New Guinea, collecting birds on the northern slopes of the Snow Mountains (Yale). The 1950s also saw the beginnings of several years of entomological surveys by the Bishop Museum (Honolulu, Hawai’i, U.S.) under the direction of the present writer’s previous co-author, the late J. L. Gressitt (collections therein). In 1955

................. 16157$

$CH2

02-08-07 10:22:36

PS

PAGE 83

84 / david g. f r o d i n

Gressitt (largely in company with R. T. Simon Thomas, then recently arrived as a government agricultural entomologist, working out of Manokwari) collected insects for a month—mainly in the Wissel Lakes area (including Kamo Valley) but also at Hollandia (now Jayapura), Sentani, the Cyclops Mountains, and Biak Island. In 1957 Gressitt revisited the Cyclops and Biak; in 1958 he spent three weeks in the Swart Valley (Nassau Range)—visited by Lam in 1920—and areas to the west. In 1959 he collected with T. C. Maa in the Cyclops, on Biak Island, and at Fakfak (including caves) and inland on the Bomberai Peninsula. In 1962 he collected in the Cyclops Mts, Biak, Nabire, and Wissel Lakes (partly in company with J. Sedlacek, N. Wilson, and H. C. Clissold). Afterwards Gressitt would become more involved with Papua New Guinea (where from 1961 he had established for the Museum a field station at Wau, which is now the Wau Ecology Institute). Gressitt did, however, make two final trips (1977 and 1979) to Papua, mainly to establish contacts at the Abepura (outside Jayapura) and Manokwari campuses of Cenderawasih University, but also visiting Biak, Sorong, and the Arfak Mountains. As an associate in Gressitt’s program—but separately—D. E. Hardy in 1957 collected at Manokwari, Anggi Lakes, and elsewhere on the Vogelkop, and Sentani and Hollandia (now Jayapura). In 1959 Maa collected at Waris, Sarmi, Holmafin, the Baliem Valley (then recently opened to outsiders), and Merauke, as well as on the Vogelkop Peninsula, besides the above-mentioned. In 1961 L. W. and Stella Quate collected at Bokondini, Lake Archbold, and Ok Sibil (in the Star Mountains). In 1962 and early 1963, in addition to Sedlacek, Wilson, and Clissold, collecting was done by L. P. Richards, Max C. Thompson, Philip Temple (a participant in expeditions to Mt Jaya, including that of Harrer), Heinrich Holtmann, and Ray Straatman (all at Bishop Museum). (Straatman later moved to Papua New Guinea.) On the applied front, during the final years of Dutch rule, Simon Thomas continued as agricultural entomologist (collections, Manokwari). Active as medical entomologists were Rudolph Sloof, Dirk Metzelaar, Johannes van den Assem, and Willem J. O. M. van Dijk; their main concerns were with malaria and mosquitoes (collections, Amsterdam).

Irian Jaya; Papua, Province of Indonesia (since 1963) After effective (and later de jure) power passed to Indonesia, little animal collecting was done for some two decades. Major activities during this time mostly centered around three surveys in the 1960s. In 1963–1964 the ‘‘Cenderawasih Expedition’’ comprising S. Soemadikarta (from Bogor) and Boeadi collected some animals east of Wissel Lakes (Bogor). Around the same time a Kyoto University science expedition worked from Wissel Lakes to Ngga Pulu; its chief biologist was Y. Yasue. In 1964 came a joint Cenderawasih University/Explorers’ Club of Nanzan University (Japan) climbing trip to Ngga Pulu (on the Mt Jaya massif); one of its participants, Yosii (Nanzan), collected Collembola. In 1977 and 1979 Gressitt made, as already noted, relatively brief visits; while in 1979 Prof. Jared Diamond (see also Fauna of

................. 16157$

$CH2

02-08-07 10:22:36

PS

PAGE 84

Biological Exploration of New Guinea / 85

Eastern New Guinea section, below) studied birds along the Mamberamo River and in the Foja (also known as Gauttier) Mts. In spite of an apparent lull in surveys and collecting, however, official recognition of the need for continuing zoological studies in Province of Irian Jaya came in 1971 with the establishment of a museum on the Abepura campus of Cenderawasih University. This now houses representative collections from the several integrated surveys of more recent decades (see Integrated Expeditions section, above).

flora of eastern new guinea: territory of papua and new guinea (tpng) (1946 –1971); papua new guinea, independent from 1975 (since 1971) Government: Division of Botany and National Herbarium, Lae The leading contributions to our stock of collections from eastern New Guinea have since 1945 been made through the Division of Botany, Department of Forests (now PNG Forest Research Institute, or FRI). Its herbarium was in the 1970s designated as the National Herbarium, and its upkeep and enhancement (along with that of the Botanic Garden), as well as research, remain the Division’s (now Section’s) core activities. The loss due to World War II of all earlier (though not numerous) local collections, the more than 2,000 made in 1944–1945 by forestry services companies (initially taught by C. T. White from the Queensland Herbarium), and the appointment of McAdam (these forces’ commander) as Director for Forests for the postwar Territory of Papua and New Guinea (TPNG) all led to a favorable climate for establishment of a botanical division within the new Department. Although having in its early years an emphasis on forest trees, relatively soon the Department of Agriculture transferred to it all its botanical interests (except for pathogenic fungi); it thus received collections from its staff as well. Established in 1946 on the 1944–1945 site and in 1949 moved to the new Botanic Garden near the War Cemetery (both sited on a former plantation), the Division was until 1975 directed by J. S. Womersley. From the second set of the Forces’ collections, left behind at war’s end, its herbarium (LAE) and associated collections were strongly developed, along with a program of botanical illustration. A purpose-designed building was opened in 1965. Over time acquisitions included many duplicates from pre-war decades. LAE now is home to a collection of some 300,000 or more higher plants and bryophytes from Papua New Guinea and neighboring areas (particularly Papua and Solomon Islands), and has hosted (and also jointly sponsored) many botanical expeditions from abroad. The staff was at its largest in the 1960s and 1970s. In the latter part of the 1980s the Division became a section within the Forest Research Institute on its incorporation, which also consolidated various Departmental research units within an adjacent new building, constructed (with more herbarium space) with Japanese assistance. During the years to 1980 (and sometimes beyond), the staff included M. Galore (chief from 1975 to 1983), A. G. Floyd, E. E. Henty (finally senior botanist and, as

................. 16157$

$CH2

02-08-07 10:22:37

PS

PAGE 85

86 / david g. f r o d i n

we have seen, translator and editor of Peekel’s florula of the Bismarck Archipelago), A. Millar (active in particular with the Garden, but from 1971 in Port Moresby), P. van Royen (later at Bishop), D. Sayers (in the Garden, later in Britain), T. G. Hartley (as an associate; later Harvard and then CSIRO, Canberra), A. Gillison (later Bulolo and then CSIRO, Canberra), J. Buderus, D. G. Frodin (the present writer, later at UPNG and Kew, and currently Chelsea Physic Garden), K. Woolliams (in the Garden, later at Waimea Arboretum, Hawai’i), C. E. Ridsdale (later Leiden), A. W. Dockrill (later CSIRO, Atherton, Queensland), M. J. E. Coode (later Kew), J. Vandenberg, D. B. Foreman (later Melbourne), R. J. Johns (later Bulolo, then PNG University of Technology and Kew), P. F. Stevens (later Harvard, then University of Missouri–St. Louis/Missouri Botanical Garden), G. Leach (later UPNG, then Darwin), W. R. Barker (later Adelaide), N. Clunie, J. Croft (later Canberra), B. Conn (later Melbourne and Sydney), P. Katik, Yakas Lelean, Kipira Damas, and J. Wiakabu. In more recent years K. Kerenga, R. Kiapranis, O. Gideon (now UPNG), and R. Banka, and, as an associate, W. Takeuchi, have been active. In excess of 60,000 numbers in the institutional New Guinea Forests (NGF, later LAE) series—continuing on from numbers of collections started in 1944—were collected over some forty years from all parts of the country, at localities too numerous to mention but with many ‘‘firsts.’’ Some of these, including a number of mountain summits, were in association with outside expeditions (see below, as well as the section on Collections, below). The Division has attracted many visitors over the five decades of its existence, some remaining for considerable periods of time and undertaking their own fieldwork or taxonomic or other studies. Some are mentioned below under ‘‘Other Collectors.’’ A significant recent project has been a revision of New Guinean palms (together with botanists in Manokwari as well as Kew botanists J. Dransfield and W. Baker).

Government: Bulolo, Konedobu Other herbaria also came into being in PNG to meet particular needs, both within (see below) and outside of the government. In 1957 the forerunner of the Forestry College at Bulolo was established within the Department of Forests and, with teaching an imperative, an herbarium was developed (with contributions by J. J. Havel, H. Streimann (later in Canberra), A. Kairo, A. Gillison, R. J. Johns, B. Conn, A. Hay (both presently in Sydney), Lawong Balun, and others). In recent years the College has moved into the higher education sector (under PNGUT at Lae). Active collecting has been carried out in the Wau-Bulolo region (for which in 1983 Streimann published a checklist of the lichenized fungi, bryophytes, and higher plants) and elsewhere. At the adjacent Forest Research Station work in forest pathology commenced about 1969; the fungus herbarium started there currently has the best local collection of macrofungi. Contributors included J. Simpson and F. Arentz. In the 1980s this unit moved to Lae when the units of the Research Station were absorbed into FRI. Another of the Station’s staff, the silviculturalist N. Howcroft, likewise moved

................. 16157$

$CH2

02-08-07 10:22:37

PS

PAGE 86

Biological Exploration of New Guinea / 87

but continued his considerable side interest in Orchidaceae (notably the grounddwelling Spathoglottis). At Konedobu, the headquarters of the Department of Agriculture, Stock, and Fisheries (DASF; later the Department of Primary Industry (DPI) and, from the mid-1980s, Department of Agriculture and Stock (DAS)), a Plant Pathology herbarium was begun about 1955. It was built up largely by Dorothy E. Shaw and more recently has been under G. R. Kula. Its contents consist largely of pathogenic microfungi, but small collections of other thallomorphic plants are also held.

Nongovernmental Institutions At the University of Papua New Guinea (UPNG), the herbarium was started in 1968 by R. Robbins and M. Pulsford but from 1971 was greatly enlarged by Frodin (until 1985) with contributions from others (including A. Millar, A. Gebo, J. Powell, C. Huxley, J. Dodd, I. Johnstone, G. Leach, and P. L. Osborne). In 1978 the herbarium was badly damaged by an accidental fire, but most collections were saved. New facilities were constructed during 1983–1984 as part of a complex known as the Natural Science Resource Centre. A principal aim was (and is) the study and documentation of the distinctive seasonal flora of the Port Moresby region (including the already-mentioned offshore island of Motupore), but research into other areas was also carried out including mangroves (including a manual), seagrasses, ant-plants (Rubiaceae), and freshwater aquatic macrophytes (again with publication, in 1985, of a manual by Leach and Osborne). After Frodin’s departure, H. Fortune Hopkins was (through 1990) lecturer-in-charge, continuing among other activities work on the Motupore florula, published jointly with Menzies (see below) in the mid-1990s. In the latter part of that decade (following completion of his PhD), O. Gideon took up this post, transferring from Lae. Technical staff—most of whom also collected—have included A. Gebo, K. Naoni, A. Vinas, M. Kuduk, and P. Piskaut, and many students similarly contributed. Undergraduate courses in plant diversity continue to be taught. A herbarium was also set up at the University of Technology (PNGUT) by R. Johns after his accession to the professorship of forestry, the full degree course of which was taught there from 1980. This served as a depository for vouchers from forestry plots and vegetation studies as well as a teaching resource. Subsequently it came under P. Siaguru.

Outside Expeditions In the three decades after World War II, numerous externally-funded botanical expeditions were conducted. With most was associated, as already indicated, the Division of Botany, where original or duplicate sets of their collections have been deposited (though in a few cases these are also, or only, at UPNG). The largest undertakings incorporating botanical work were the four later Archbold Expeditions (1953–1964), on which Brass was botanist for the first three and Hoogland for the last, and the several CSIRO land surveys (mostly 1953–1969; for both, see section on Integrated Expeditions, above) but there were many others, principally

................. 16157$

$CH2

02-08-07 10:22:37

PS

PAGE 87

88 / david g. f r o d i n

from Europe. Some comprised a single individual who might make Lae a base and run field trips from there over several months to a few years. Of particular interest for some of the external expeditions were, not unnaturally, the high mountains; and in general the montane flora with its distinct Gondwanan affinities was attractive (and now more accessible). In 1956 Hoogland and Pullen worked on Mt Wilhelm (first named, as already indicated, by Zo¨ller in 1888, but only later ascertained as the highest in eastern New Guinea), where they were joined by Womersley and Galore (LAE). Their collections were the first of any real extent to be made on this peak (in 1970 the subject of a florula, and where for a time ANU maintained a field station). In 1957 R. C. Robbins worked on vegetation on Mts Wilhelm, Hagen, and Giluwe, and, also in 1957, J. C. Saunders was in the Kubor Mts. Several mountains in the Eastern Highlands and Simbu were visited by Brass in 1959 with the Sixth Archbold Expedition, while in 1964 Hoogland climbed high into the Saruwageds with the Seventh Archbold Expedition. In 1960 Hoogland, Schodde, and Robbins worked on Mt Sugarloaf (west of Mt Hagen); in 1961 Hoogland and Darbyshire worked in the Torricelli Mts, and in 1966 Hoogland and Craven spent some time in the Hunstein (also known as Sumset) Range—making the first plant collections there since Ledermann. By the 1980s such expeditions, along with those of locally-employed scientists, had covered most parts of the country; but such was the terrain as well as issues of access that many gaps still remained; also, the unusual flora present over some substrates was only gradually recognized for what it was. Moreover, a number of areas worked by German collectors before 1914 had never been revisited. In the last two decades, there has been some gap-filling—particularly under outside sponsorship and often as part of more general bioinventories. A byproduct of the CSIRO botanical work was an extensive card file (maintained through 1968) of taxa in the New Guinea flora with literature references compiled by Hoogland (CSIRO, Canberra; two copies exist in book form—one in FRI, Lae). It has, however, yet to be included in a database or even digitally imaged in the way that the large Hu card index for China at Harvard has been. But for the most part the very substantial botanical collections were not worked up into separate reports but simply have joined the general stock drawn upon for Flora Malesiana, its precursors, and individual revisions. Very many remain to be documented, with some of them as-yet-undescribed novelties. A large proportion of the distributed collections has been entered into databases, but as part of other projects.

Other Collectors C. R. Stonor, in the course of his ornithological work, in 1948–1949 collected plants on Mt Hagen and Mt Wilhelm (Edinburgh). In 1954 H. S. McKee collected in the Wahgi Valley (Brisbane, Lae). In 1955, in continuation of his long expedition in western New Guinea, P. van Royen collected at Bulolo, the Bulldog Track south of Wau, Lae, and near Port Moresby (Leiden). From 1958 (continuing until 1979) Dr D. Carleton Gajdusek (National Institutes of Health, United States) col-

................. 16157$

$CH2

02-08-07 10:22:38

PS

PAGE 88

Biological Exploration of New Guinea / 89

lected plant and animal material as vouchers for food and drugs in the Eastern Highlands and many other areas while studying the spongiform encephalopathic disease, Kuru (related to ‘‘mad cow,’’ or BSE, and nvCJD), and related topics, for which he became a Nobel Laureate. In 1960 J. A. R. Anderson collected 22 numbers on Mt Wilhelm (Edinburgh) and in the same year F. R. Fosberg collected in the vicinity of Goroka, Lae, and Rabaul (USNM). The relative trickle of the 1950s now turned into a flood, and substantial collections in all groups were made. In the first half of the 1960s P. van Royen (while on the LAE staff) collected with H. O. Sleumer on Mt Wilhelm and in New Britain (Sleumer particularly interested in Rhododendron), on Mt Wilhelm with the plant physiologist F. W. Went (then director of the Missouri Botanical Garden), and on the Huon Peninsula with S. Carlquist (RSA). The 1962 Noona Dan expedition (see section on Integrated Expeditions, above) made a considerable plant collection in the Bismarcks. In 1963, W. Vink (Leiden) and Pullen (CSIRO) collected on the Kubor Range, while Carlquist collected with Henty (LAE) at Mt Piora. F. Kleckham of DASF collected plants on Mt Strong; G. Rosenberg collected plants and insects on Mt Amungwiwa; and R. Heim collected fungi in the Central Highlands (Paris). In 1964–1965 A. Clive Jermy (BMNH) led a BMNH/University of Newcastle-upon-Tyne expedition, initially working in the Finisterres—not visited since before 1914—and then in parts of the Highlands including Okapa, Mt Elandora (Kra¨tke Range), and Mt Wilhelm as well as in the Herzog Ranges south of Lae. He was accompanied by T. G. Walker (who later collected in New Britain), A. Eddy, P. W. James, and M. E. Bacchus; for the Finisterres they were joined by Sayers (LAE) and Pullen (CSIRO). In the first serious undertaking of its kind, pteridophytes and non-vascular cryptogams were emphasized; but some 1,000 numbers of phanerogams were also obtained. During 1962–1965 T. G. Hartley—connected to CSIRO but not part of the land survey program—was engaged in systematic sampling for alkaloids and other medicinal properties (an extension of similar work on the Australian flora); several localities, mainly in Morobe and Eastern Highlands, were visited (plants, CSIRO, Canberra). His final report, A Survey of New Guinea Plants for Alkaloids, appeared in 1973 (Lloydia 38: 217–319), covering, in taxonomic sequence, all his collections. (Later he joined the organization as a taxonomist in the present Australian National Herbarium and remains active in Malesian and Australasian taxonomic research.) The opening of the new herbarium for LAE in early 1965 facilitated the work of outside visitors as well as the staff. In that year Gillison, van Royen, and Buderus (LAE) worked on Mt Biota, west of Mt Albert Edward, and B. Craig (later with the National Museum in Port Moresby) in the Star Mts. Later M. J. van Balgooy visited the Territory of Papua and New Guinea (TPNG), notably working on Mt Wilhelm (Leiden, his collections particularly fine). His trip was the first of a number organized from Leiden in pursuit of the notable high-mountain and alpine flora (eventually a major basis for van Royen’s Alpine Flora of New Guinea). Balgooy was followed in 1966 by C. Kalkman and W. Vink. With Gillison and Frodin

................. 16157$

$CH2

02-08-07 10:22:38

PS

PAGE 89

90 / david g. f r o d i n

(LAE) as local counterparts, they spent some three months in the Doma Peaks, followed by a short visit to the Telefomin area including part of the Hindenburg Range (Leiden, Lae). In August 1968, on a privately financed expedition, P. Woods (Edinburgh) and M. Black with Ridsdale (LAE) collected towards Murray Pass and on Mt Albert Edward, with Woods and Black specializing in orchids (Leiden; Lae). In 1969–1970 M. J. S. Sands (Kew) with Coode (LAE) worked on New Ireland, obtaining the first significant collections from its southern mountains (Kew, Lae). The strong pace carried on well into the 1970s, the last decade of relatively liberal public funding for fieldwork. In 1970–1971 Y. Kobayashi collected lower plants on Mt Wilhelm, Mt Hagen, and at Oksapmin. In 1971 F. R. Mitchell (from the United States) came from Christchurch and collected plants in several areas (DSIR, Lae). In 1972 there was another substantial joint Leiden-Lae mountain massif expedition, this time to Mt Suckling; members included J. F. Veldkamp (Leiden), Stevens, Frodin (UPNG), N. Cruttwell, R. Pullen, F. Essig, and entomologists T. L. Fenner (DASF, Konedobu) and Gressitt (plants, Lae, Leiden; insects, Konedobu, Bishop, etc.). In 1973 M. Jacobs (Leiden) and K. Paijmans (CSIRO) collected on Mt Bosavi. In 1974 Craven and Croft worked on Mt Victoria. In 1975 the last of the major Leiden-Lae expeditions worked in the eastern Star Mts with Veldkamp and A. Touw (Leiden) and Barker, Conn and Croft from LAE (Leiden, Lae). In the same year Sands was again in New Ireland (with G. A. Pattison and J. J. Wood) as well as on the mainland (Kew, Lae), while a group of Japanese botanists led by S. Kurokawa and M. Inoue collected mainly cryptogams on Mt Albert Edward, the Wau area, and the Saruwaged Mts. The Bishop Museum in 1976 collaborated with WEI in a primarily botanical expedition to several high mountain areas, with van Royen as leader (collections at Bishop and WEI). Visits were made to Mt Victoria (Royen, P. Kores, Frodin, R. T. Corlett); Mt Amungwiwa (Royen, B. Gagne´); the Finisterres (Royen, Kores, B. Gagne´, and Gressitt), and the Victor Emanuel Mts (Royen, Kores, B. and W. Gagne´, Gressitt). The previous year, Corlett (ANU) also collected on Mt Giluwe, Mt Amungwiwa, and elsewhere, while the Gagne´s continued to collect insects as well as plants in various places, 1976–1980 (with a tour to the Louisiades in 1979). In 1977 Veldkamp worked in the Western Highlands (Leiden). Churchmen were also active, as in the past. Canon Norman E. G. Cruttwell (1916–1995) was an Anglican missionary from 1946 until the late 1980s. He was long stationed at Dagwa (eastern Papua) and from 1976 at Goroka, and developed a particular interest in rhododendrons and orchids. He visited, among other high spots, Mt Simpson in 1947 and 1968, Mt Dayman in 1951, 1974, etc., Mt Amata in 1959, and Mt Suckling (1972, see above). Outside Goroka he set up a high altitude botanic garden (Lipizauga Sanctuary) in the Gahavisuka Park—one of only two such parks in the country relatively near urban centers. Among other missionaries doing botanical work were the Rev. H. A. (Bert) Brown, based at Elema but also traveling to the Kunimaipa (near Mt Strong); and, until the end of the 1970s, Br. William Borrell at Kairiru off Wewak. In 1960 Fr. E. Borgmann

................. 16157$

$CH2

02-08-07 10:22:39

PS

PAGE 90

Biological Exploration of New Guinea / 91

(Mingende, Simbu Province) studied chromosome numbers of Mt Wilhelm plants, shortly thereafter publishing a landmark paper. Anthropologists also made substantial plant collections. R. Bulmer (and Ian Saem Majnep, active in the Simbai District of Madang Province) were particularly notable for thoroughness of collecting and documentation; so, too, was N. Bowers (Western Highlands). This work has been further pursued by R. Gardner (Auckland) in more recent years. The geographers H. E. Street and H. Manner in 1967 collected in the Koinambe area of the western Bismarcks (Lae, Leiden). Medicinal plants were studied over a number of years by D. K. Holdsworth (UPNG), with some collections deposited in that herbarium (in 1991 he contributed to the first volume of a general work on PNG medicinal plants). In the 1960s, P. Eddowes made collections in connection with wood-anatomical work (Lae, UPNG). With respect to particular plant groups, the collecting of orchids has been accomplished by—among others—A. Millar, P. Woods (see above), N. Howcroft (see above), T. Reeve (Laiagam), J. Dodd (see above), and, for a few years from 1988, P. O’Byrne (Port Moresby). O’Byrne’s field and related work soon led to his Lowland Orchids of Papua New Guinea (1994)—the first major work on the New Guinea orchid flora since those of Schlechter (an English edition of which was published in 1982), although Orchids of Papua New Guinea: An Introduction by Millar had first appeared in 1978. Over the last decade a full revision of New Guinea orchids has been under way under E. de Vogel (Leiden), resulting in a series of CD-ROMs; some groups were also studied at Kew. A major survey of Dendrobium by H. P. Wood appeared in 2006. Henty focused in grasses, his work leading to A Manual of the Grasses of New Guinea (1969). R. E. Holttum also studied bamboos in addition to ferns (see below). B. Verdcourt carried out fieldwork in PNG in 1976 and 1978 under a U.K. government grant, resulting in Legumes of New Guinea (1979). In the early 1970s G. Argent (Edinburgh) worked on Musa; in addition to the assembly of a living collection of bananas in Lae his work led to a still-standard taxonomic revision. Sleumer specialized in rhododendrons, and before long rising interest in the Vireya group (to which all native species belong) and other mountain Ericaceae brought others: Woods and Black in the late 1960s (see above), H. F. Winters and J. J. Higgins (USDA) in several areas in 1970 (U.S. National Arboretum), and, in 1978–1979, K. Arisumi and associates (Kagoshima). Croft and Johns have specialized in ferns, a group in connection with which (in addition to Jermy and Walker; see above) Holttum and B. Parris(-Croxall) also paid visits, the latter—now in New Zealand—collecting in several areas, 1971– 1972 and 1977 (Cambridge, Lae). There were also other expeditions specializing in bryophytes and other plant and fungal groups (including lichenized fungi) as well as some resident activity in these groups (e.g. by Dorothy Shaw, Heiner Streimann, and Peter Lambley). A considerable proportion of their studies has been published, with all those named here producing annotated checklists in their groups. In the 1960s and 1970s a number of botanists were aided in the field by Aubita

................. 16157$

$CH2

02-08-07 10:22:39

PS

PAGE 91

92 / david g. f r o d i n

Kairo, an assistant at Bulolo Forestry College, who could from sterile material recognize most genera of trees and who identified many in the WEI arboretum. Paul Katik, Artis Vinas (until 1979, then UPNG and Bulolo), Yakas Lelean, and (in earlier years) Michael Galore and John Koibua also assisted many (including the writer). Katik also acquired an ability to recognize almost all genera and many species of seed plants at sight, so ranking with Indonesian official (mantri) Nedi at Bogor. But, due to retrenchment, Katik had to leave the FRI in 1992 and since has worked privately. Other local assistants were associated with UPNG.

Ecological and Paleoecological Investigations The vegetation history and pollen analysis studies carried out from 1960 until the late 1970s under the direction of Professor Donald Walker (ANU) involved substantial fieldwork. Early studies were largely carried out in Western Highlands and Enga provinces. In 1965 Walker established the Mt Wilhelm research laboratory at Pindaunde (3,300 m) with help from Bishop Museum and the Territory of Papua and New Guinea (TPNG) administration. Much research was done here on vegetation, palynology, and climate. In 1974, the station was ceded to the National Parks Board. Some specific studies include: 1965, Walker and J. R. Flenley on pollen deposits in Enga Province (Gressitt 1982: chapter III, 1); 1966, Jocelyn Wheeler Powell on Eastern Highlands deposits (Gressitt 1982: chapter II, 4); 1966–1967, intensive studies on Mt Wilhelm by D. McVean and L. K. Wade; 1968, Walker, B. O. van Zanten, W. A. Weber (Colorado), McVean, and Johns (prior to his Government and PNGUT service) on vegetation studies including the cryptogamic flora at Mt Wilhelm; 1968–1969, G. S. and Jeannette Hope on vegetation history and animal ecology at Mt Wilhelm, G. S. also working at Mt Giluwe and later in the Owen Stanley Mts (as well as at Mt Jaya; see sections above on Integrated Expeditions and Flora of Western New Guinea); and 1972, J. A. Peterson (see also the justmentioned sections) on glacial history, and J. M. B. Smith on alpine plant ecology at Mt Wilhelm and on the Kubor Range (Gressitt 1982: chapter III, 2). In 1976 S. Garrett-Jones worked at Lake Wanum (near Lae; ‘‘inland’’ mangal remnants are known in the vicinity). In 1977 planned fieldwork at Lake Trist (east of Wau) was aborted, but some studies were carried out at Lake Kutubu and a small lake near Mt Ialibu. Other vegetation studies include those by R. Hynes, J. Ash (Gressitt 1982: chapter III, 5) and N. Clunie on Nothofagus (Clunie also on New Britain); N. Enright on Araucaria (Gressitt 1982: chapter III, 6), P. C. Heyligers in the Port Moresby region, 1963–1967 (Gressitt 1982: chapter III, 8); K. Paijmans on Mt Albert Edward in 1970 and in Galley Reach in 1975; and P. Edwards and P. J. Grubb near Mt Kerigomna above Marafunga in 1970–1971. For some twenty years from 1970, Johns, while at Bulolo and Lae, sampled numerous vegetation plots (references in P.N.G. Conservation Needs Assessment, 2: 55–60. 1993).

................. 16157$

$CH2

02-08-07 10:22:40

PS

PAGE 92

Biological Exploration of New Guinea / 93

The 1980s and Beyond After the mid-1980s, with changing interests, increasing financial stringency (and irregularities), ‘‘privatizations,’’ a decrease in the quality of life (including security) in many more closely settled areas (including the cities and towns), more regulations, and—notably—an insecure central government, collecting declined notably. Only a few people have done anything substantial, notably W. Takeuchi in many areas under varying sponsorship since the late 1980s, and the present writer (with G. Morren, Rutgers University) in 1992 and 1993 in the Telefomin District, this latter undertaking with particular reference to uses and local ecology among the Miyanmin—an area not otherwise sampled (Kew, Lae). In 1992, with E. Gabir, Frodin and Morren reached some Miyanmin lowland areas along the Iwa (also known as May) River (Fiak Airstrip and Hotmin—the latter ca 10 km or so south of the furthest point reached by the Behrmann expedition). There have also been a number of more targeted field undertakings, including (as already mentioned) those for palms and, as of late 2005, for orchids (E. de Vogel, pers. comm.). Government activity has now dropped to relatively low levels. Takeuchi has indicated that since 1989 additions in the NGF/LAE institutional numbering sequence have averaged only some 400 numbers/year—well below earlier rates, particularly in the 1960s and early 1970s. This reflects significant staff cuts since the 1970s; the focus is presently on maintenance of existing resources. The alpine flora was gathered together in an impressive series of volumes (1980–1983) by P. van Royen, following upon his 1976 expedition. This must, however, be but a starting point for further research on this fragile ecosystem; probably only at Mt Wilhelm (and Mt Jaya) has collecting reached a relatively advanced level—almost everywhere else visits have been at scattered times. The Mt Victoria complex, for example—although the first to be visited (in 1889)—has only rarely been studied since (e.g., in 1976; see above). At lesser elevations, knowledge remains patchy. Effective inventory of any given area will come about only with repeated, relatively sustained visits and where conditions (including local relations) are favorable, and most likely with outside or nongovernmental organization (NGO) support. A fair idea of our present knowledge may be had from the various recent conservation needs assessments (see References section, below). Formal documentation is, however, likely to remain a slow process (and to many recondite in language and style) without radically different approaches.

fauna of eastern new guinea: territory of papua and new guinea (tpng) (1946 –1971); papua new guinea, independent from 1975 (since 1971) In contrast to the plant world over much of this period, there was no single government body—nor perhaps could there have been—responsible for fauna, there being many different stakeholders. As a result, a number of separate official collections came into being, some of them before re-establishment (in 1954) of the old Papuan territorial museum as the National Museum. Even afterwards the Muse-

................. 16157$

$CH2

02-08-07 10:22:40

PS

PAGE 93

94 / david g. f r o d i n

um’s primary focus in research was on ethnography and archeology, although gradually a zoological collection, predominately of higher vertebrates, was built up. Of agency collections, DASF (later DPI) accumulated important holdings of fish (and other aquatic organisms) as well as insects, while the Department of Forests developed a collection of forest insects at Bulolo (later moved to Lae). Collections were also built up at Wau Ecology Institute (see below) and the universities (particularly the University of Papua New Guinea). There is also a collection at the Parataxonomy Center in Madang.

Vertebrates N. B. Blood collected birds in the highlands 1945–1947 and later (AM). Fred Shaw Mayer (see above) collected birds in the Mt Hagen area in 1946–1947 and on Mt Wilhelm in 1949. In 1948 Blood set up at Nondugl in the Wahgi Valley the bird of paradise (plus other wildlife) sanctuary partly funded by Sir Edward Hallstrom. Its management was passed in 1954 to Shaw Meyer and in 1960 to M. J. Tyler (see below), In the early 1960s it was turned over to the government and moved to suitable forested land by the Baiyer River, also in the Western Highlands. It is now a National Park, with as well some botanical significance. The first two managers at the new location were Graeme George and Roy Mackay. Another government manager of this period, Angus Hutton (of the tea plantation at Garaina and previously resident in India), collected animals at Garaina and elsewhere in southeastern Morobe. A contemporary of Blood and Shaw Mayer was E. Thomas Gilliard of AMNH, there an associate of A. L. Rand. Gilliard made several trips (1948–1959) to collect birds (AMNH): Astrolabe Range (1948), Wilhelm, Giluwe (1950), the Kubor Range (1952), the Victor Emanuel and Hindenburg ranges (1954), Mt Hagen and the Finisterres (1956), New Britain (1958–1959) including a climb into the remote Whiteman Range, and the Adelbert Range (1959). He also collected butterflies (AMNH), obtaining new species of Delias, and a few plants (Harvard). Gilliard was partly sponsored by the National Geographic Society of Washington DC, and wrote and photographed for National Geographic. Apart from formal papers, much of his work is summarized in his Birds of Paradise and Bower Birds (1965) which also contains useful historical material. He was in addition a co-author (with Rand) of Handbook of New Guinea Birds (1967). The Gyldenstolpe expedition of 1951 in the Hagen/Wahgi area concentrated on birds but also collected other animals and plants. Its principals were the Swedes Nils Gyldenstolpe and his wife (Stockholm; results in Ark. Zool. 8(1), 1955). In 1954 Ellis Troughton, assisted by Norman Camps as well as by Blood, collected mammals in the Western Highlands (AM; Australian Mus. Mag. 11: 246). Blood and Camps also assisted Gilliard and Gressitt. In 1955 Rev. O. Shelly collected some frogs in the Wahgi Valley (AMNH). In 1959 Reimer collected some animals in Kikori area, Papua Gulf (Mu¨nchen, Frankfurt). Turning to the 1960s, the establishment of the Bishop Museum Field Station (later the Wau Ecology Institute) at Wau (1961) provided a new focus (see also

................. 16157$

$CH2

02-08-07 10:22:41

PS

PAGE 94

Biological Exploration of New Guinea / 95

Integrated Expeditions and Surveys section, above). Vertebrate research also figured in its activities, with mammals and their ectoparasites of particular interest. Collections were made by M. C. Thompson and P. Temple; R. Traub (USNM) with Abid Beg Mirza, M. Nadchatram, E. Mann, Wilson, Ziegler, R. Greene (Bishop; WEI), D. Schlitter, and S. Williams (also Carnegie). Early outside visitors included, in 1962, Alden H. Miller (birds) and W. Z. Lidicker, Jr. (mammals), focusing on the Wau-Bulolo area (MVZ). Separately, Professor T. C. Schultze-Westrum (a nephew of L. Schultze-Jena; see above) in 1964 and 1970 studied mammals and conservation at Mt Bosavi and elsewhere for the IUCN and the Territory of Papua and New Guinea (TPNG) authorities. Also in 1964 and continuing over several visits, Jared Diamond (most recently author of Guns, Germs and Steel (1997) and Collapse (2004)) undertook his detailed surveys of birds of certain mountains, including Mt Karimui and the Torricellis, pioneering certain aspects of ecology and biogeography and at the same time making recommendations for conservation (collections mainly AMNH). In 1972 Diamond published his critical Avifauna of the Eastern Highlands of New Guinea, a notable contribution to the literature. During the CSIRO surveys (see above, Integrated Expeditions section) R. Schodde collected many birds (and some insects) as well as plants, for example, with S. Schodde at Lake Kutubu and Mt Giluwe (1961) and in Central Province (1962). W. B. Hitchcock collected birds in 1963 with botanists W. Vink and R. Pullen (CSIRO), Pullen also collecting some insects (CSIRO). The Noona Dan expedition of 1962 (see Integrated Expeditions section, above) in its survey of the Bismarck Archipelago also collected animals, with Torben Wolff, Leif Linneborg, Finn Salomonsen, and Wm. Buch particularly interested in birds, insects, and fresh water and marine organisms (Copenhagen). More detailed herpetological work had its advent in the 1960s. Fred Parker, a government officer, from 1960–1978 made large herpetological collections in many parts of the Territory of Papua and New Guinea (MCZ, AM, etc.). Harold Cogger collected reptiles and amphibians in several areas, especially in the former Territory of Papua. Michael J. Tyler, Shaw Mayer’s successor at Nondugl, from 1960 collected many frogs and reptiles in Wahgi Valley and on the Wahgi-Sepik divide; and in 1967 he made a frog survey of New Britain (Adelaide, AM, BMNH, AMNH). Richard Zweifel in 1964, 1968, and 1969 collected many frogs in several areas in the Highlands, the Wau area, and along the north coast (AMNH). In 1969 Harold Heatwole, while on the Fairbridge New Guinea coral reef expedition, studied reptiles on some smaller islands in eastern Papua. George Zug collected reptiles and amphibians in 1971 at Mt Kaindi and elsewhere (USNM). In 1968 James Menzies joined the staff of UPNG, beginning his long association with New Guinea herpetology (as well as mammalogy and, with collaborators, in botany); after a period in Africa in the 1980s he returned, first to the National Museum, then back to UPNG (retiring in 2001). In more recent years Tim Flannery (now in Adelaide, and author of Mammals of New Guinea (1990; 2nd ed., 1995) and Mammals of the South-West Pacific and

................. 16157$

$CH2

02-08-07 10:22:41

PS

PAGE 95

96 / david g. f r o d i n

Moluccan Islands (1995) as well as The Future Eaters (1994) and Throwim Way Leg (1998), the last an account of his fieldwork from 1981 onwards) has been the most prominent in mammology; but Menzies also made significant contributions including A Handbook of New Guinea Marsupials and Monotremes (1991). In ornithology B. Beehler (now Washington DC), B.J. Coates (now Queensland), T. K. Pratt (now Hawai’i), and M. LeCroy (New York) have been very active since the 1970s; and resident ‘‘twitchers’’ (especially B. W. Finch, P. Gregory, K. D. Bishop) have continued their observations. Most notable, perhaps, were the doctoral and postdoctoral ornithological field studies conducted on Crater Mountain and Mt Missim in the 1970s and 1980s by A. Mack, D. Wright, B. Beehler, T. Pratt, and S. Pruett-Jones. These encompass the most significant fieldwork completed to date on New Guinea’s birdlife, and at Crater Mountain led the way towards long-term involvement (see section on Integrated Expeditions, above). Beehler and Pratt (with D. A. Zimmerman) have published what is now a standard handbook (Birds of New Guinea, 1986; see also Burung-burung di Kawasan Papua, 2001). The Bismarcks (with the Solomons) were quite recently also the subject of a monograph, The Birds of Northern Melanesia (2002) by Mayr and Diamond. In herpetology, besides Cogger, Tyler, and Menzies (particularly in amphibians), M. O’Shea has studied reptiles (notably snakes). Gerald Allen (Perth) has been active in ichthyology, publishing moreover two field guides, respectively in 1991 (freshwater fish) and 1993 (reef fish)—both through the Christensen Research Institute. At the present time, the vertebrates of Papua New Guinea (or New Guinea as a whole) are relatively well covered in field and technical guides, all dating from the 1980s or later and to some of which specific reference has been made. Nevertheless, as the various conservation assessment reports (see References section, below) indicate, many gaps remain.

Arthropoda (Insects and Spiders) As with birds, an early sponsor in entomology after World War II was Hallstrom—in effect carrying on the Rothschild tradition. He engaged Wm. W. Brandt, earlier from Europe, to make a large collection of Lepidoptera—primarily butterflies—in the Territory of Papua and New Guinea (TPNG) from 1949 to 1955, covering many localities. Hallstrom then terminated the operation and gave the collection for the future national collections (while depositing it for safe-keeping with CSIRO at Canberra, in the manner of Macgregor long before with his official collections, sent to the Queensland Museum). Brandt then collected half-time for the Bishop Museum (miscellaneous insects) from 1956 to 1960 (Bishop), the other half of his time being spent on Lepidoptera for the Territory of Papua and New Guinea (and adding to the collections deposited by Hallstrom; CSIRO). Later on he was employed by CSIRO to curate the collection and to do some further collecting in the Territory of Papua and New Guinea (CSIRO). In the 1950s Stan Christian, Harry A. Standfast, Wallace Peters, and others worked on mosquitoes and built up a reference collection (now at UPNG) as well as making studies of malaria transmission and teaching mosquito control. Gordon

................. 16157$

$CH2

02-08-07 10:22:41

PS

PAGE 96

Biological Exploration of New Guinea / 97

Dun of the Mandated Territory service (see section Between World War I and World War II, above) carried on with DASF to about 1960. In 1954 he was joined by Dr Joseph J. H. Szent-Ivany, a refugee from Hungary (where he had been a curator of Lepidoptera at Budapest and university lecturer in zoogeography) who had migrated to Australia. On field trips and vacations until in 1966 retiring from government service (and later at Wau Ecology Institute; see below) Szent-Ivany collected extensively in various areas, greatly adding to the DASF collection (Konedobu). Other government entomologists who helped add to the DASF collections included J. H. Barrett, T. L. Fenner (see also Flora of Eastern New Guinea section, above), J. Healy, L. Smee, A. Catley, R. M. Stevens, T. V. Bourke, G. Baker, E. Hassan, Stuart Smith, J. N. L. Stibick, and Jan Greve. Further additions were made in 1962–1963 by J. Allen, J. M. Carlisle, D. Hutton, I. Johnson, D. Price, M. Stevens, M. Erben, and G. Rosenberg who collected insects on various mountaintops in those years. In the 1970s Donald Sands (later chief entomologist) collected butterflies as well as other insects (DASF; CSIRO). In 1955 Edward O. Wilson, then a graduate fellow at Harvard University, collected and observed ants in various areas, mainly in the northern and southern lowlands but also ascending to the top of the Saruwaged Range (MCZ). From this work he developed the concept of ‘‘the taxon cycle’’ as an evolutionary geographical process. Elements of E. O. Wilson’s work in New Guinea and in the Solomon Islands also feature in his many popular books (e.g., The Diversity of Life, 1992; Naturalist, 1994). In that same year J. L. Gressitt (see also above) commenced his 26 years of annual visits or extended stays, focusing on insects: 1955 in the Eastern and Western Highlands, Mt Otto, Mt Wilhelm, the Jimi Valley, the north coast, Manus, New Ireland, and New Britain; 1956 in Kokoda, New Britain (including a coconut beetle study for DASF), and the Solomons; 1957 in the Solomons, the north coast, and Biak; 1958 in Southern Highlands and Baiyer River; 1959 on the north coast, in the Cyclops Mts, and at Fakfak; 1960 in the Oriomo region and Cape Rodney; 1961 (with wife Margaret) to many highland areas, Mt Karimui, and Mt Giluwe, as well as Maprik, Dreikikir, Lae, and Popondetta. With longer-term surveys and studies in mind, during his 1961 visit Gressitt established the Bishop Museum Field Station at Wau (see also section on Integrated Expeditions, as well as under Vertebrates above). There, he settled in residence Joseph Sedlacek, his wife Marie, and their son J. H. Sedlacek. There then ensued continuous insect collecting for the Bishop Museum, along with the already-mentioned collection of parasites with their vertebrate hosts as well as plants for identification of insect hosts. Besides the Sedlaceks’ decade of work in succession to Brandt, Ray Straatman (after migrating from Netherlands New Guinea) collected 1961–1966, and Abid Beg Mirza 1967–1974 (and in 1975–1980 part-time while manager of WEI). For shorter periods during the 1960s many others (not all entomologists) participated, including T. C. Maa, Wallace A. Steffan, G. A. Samuelson, L. W. and S. Quate,

................. 16157$

$CH2

02-08-07 10:22:42

PS

PAGE 97

98 / david g. f r o d i n

Y. M. Huang, S. Sirivanakarn, Nixon Wilson, F. J. Radovsky, and A. C. Ziegler (all associated with the Bishop Museum research staff); Dr and Mrs Szent-Ivany as associates for varying periods of time; and E. J. Ford, Jr., Peter Shanahan, Geoff Monteith, P. H. Colman, N. L. H. Krauss, and H. Clissold as well as local assistants Tawi Bukam, Rennie, Spanis, Wita, Gewise, G. Nalu, and others. Cooperating researchers from other institutions largely funded by or through Bishop Museum included Drs C. D. Michener, E. N. Marks (see section on the World War II era and also below), John Smart, Peter Mattingly, D. Eimo Hardy, Robert Traub, M. Nadchatram, J. Balogh (1969 trip only), and Y. Hirashima. Field areas examined by Gressitt included, in 1963 and 1966–1967, Enga, Tari, the Kubor Mts, and Mt Michael; in 1965 (with T. C. Maa) to Mt Ialibu; in 1966 (with E. C. and R. Gressitt) to Mt Wilhelm and the Kubor and Schrader ranges; in 1969 to the Bulldog Road, Garaina (with M. and E. Gressitt), Mt Wilhelm, and Angoram (with Balogh and Hirashima); and in other years many other areas (all Bishop Museum, with some reference material retained in Wau). From 1970 work at Wau gradually shifted to ecological and other research (covering a wider field) as well as, in time, education, conservation, and handbook preparation. In 1971 the Field Station became separately incorporated as Wau Ecology Institute; but many of those mentioned in this section continued to work or be based there, and others would join them in various capacities for targeted research (among them Hampton Carson on Drosophila diversification). In the later 1970s Gressitt was succeeded as Director by H. Sakulas, but until Gressitt’s tragic death in April 1982 in China he remained closely associated. From 1962 another important local collection began its development, this time at the Forest Research Station in Bulolo. Barry Gray and Ross Wylie, and later Peter Shanahan, John Dobunaba, and Dr H. Roberts, built up a large collection and much data for forest insects. In the latter 1980s the collection was moved to Lae as part of the Forest Research Institute (see section on Flora of Eastern New Guinea, above), where it remains. Many scolytid beetles were described from this collection by Schedl. From the 1970s a reference collection was also assembled at the Insect Farming and Trading Agency in Bulolo; for some years it was headed by the lepidopterist Michael Parsons who while there also developed extensive data on distribution—later part of a substantial book (The Butterflies of Papua New Guinea, 1998). Parsons also studied some of the host plants. A laboratory for study of the screw-worm fly was set up around 1964 by CSIRO—run in connection with related activities in Australia, and their first longer-term presence in the Territory of Papua and New Guinea (TPNG). P. Spradbery, D. Sands, and R. Tozer were entomologists there, the last-named active into the 1980s. A wide range of insects was collected by others, both within and outside of government and by amateurs as well as professionals. From the latter part of the 1950s onwards beetles in particular continued to attract amateurs, so continuing the tradition of von Benningsen (see section on New Guinea and the Bismarck Archipelago, 1875–1914, above). Sir Alan Mann, in the 1950s Chief Justice of the

................. 16157$

$CH2

02-08-07 10:22:42

PS

PAGE 98

Biological Exploration of New Guinea / 99

Territory of Papua and New Guinea, collected many beetles as well as other insects. A somewhat later (1962–1974) beetle collector was Dr R. W. Hornabrook, a government physician initially at Okapa studying Kuru (see above) and later first director of the Institute of Human Biology at Goroka (now the Institute of Medical Research). Most of his collections were from the Highlands and in the Huon Peninsula (Wellington; some types in BMNH and Bishop). Another amateur, Henry Ohlmus, from 1961–1980 assembled much material from the highlands and elsewhere (mostly Canberra and Brisbane; some types in Bishop). A Canadian amateur, R. Parrott, in 1970 made a large collection in the northeast. Janos Balogh (partly with I. Loksa) on six visits (1956–1980) made many collections, mostly of soil arthropods (Budapest). In 1957 Drs Eugene Munroe and George Holland from Ottawa made extensive collections of moths, mostly at highland localities (CNC). In 1964 M. E. Bacchus, as a member of the BMNHUniversity of Newcastle-upon-Tyne expedition (see section on Flora of Eastern New Guinea, above), collected beetles (especially water-beetles) near Wau, Huon Peninsula, etc. In 1966 Prof. J. Illies (general editor of the Junk series Monographiae Biologicae) searched for Trichoptera in the Eastern Highlands (Schlitz). In 1968–1969 Pierre Jolivet studied ecological aspects of several arthropod groups, including parasites, in the Eastern Highlands and elsewhere; later he published an introduction to New Guinea entomology (see section on References, below). In 1978 P. Deharveng collected Collembola, both at Wau and elsewhere (Toulouse). From Australia D. K. McAlpine in 1963–1964 collected Diptera (AM). Elizabeth Marks made several visits collecting mosquitos, mostly in Papua between 1957 and 1979 (QIMR). G. Monteith in the 1960s collected insects, especially Hemiptera on several visits (Bishop, Brisbane). I. W. B. Thornton (Latrobe University) and C. N. Smithers (AM) in 1970 collected Psocoptera in several areas of the mainland and in 1974 in the Bismarcks. In recent years active entomologists have included Scott Miller (Bishop, then Smithsonian) and D. Polhemus (Smithsonian), the latter interested in particular in aquatic insects.

Other Invertebrates F. R. Allison collected Protozoa (NZ). D. F. McMichael collected land and freshwater mollusks at many areas, both east and west on two trips (1955, 1956) as a Bishop Museum fellow (AM). Y. Kondo collected some land mollusks, 1965 (Bishop). But much activity here was associated with marine expeditions (see section on Integrated Expeditions, above).

paleohistory: brief remarks (since 1945) Animal fossils were studied in the Bulolo-Watut area by Michael Plane (CSIRO). Ethnological and archeological animal material was studied in the highlands (Jimi Valley, 1955, etc.) by Ralph and Susan Bulmer (AM, Auckland; see also Gressitt 1982: chapter II, 3). Archeological animals were also studied by J. Hope in 1969 (ANU; see also section on Flora of Eastern New Guinea, above) and James Men-

................. 16157$

$CH2

02-08-07 10:22:42

PS

PAGE 99

100 / david g. f r o d i n

zies in the 1970s (UPNG). Vertebrate fossils were also obtained and studied by T. Flannery (working with Plane), with a contribution on the Pureni area (Southern Highlands). Plant fossils (notably pollen) were the subject of several years of work by D. Walker and his associates on the Quaternary era (see Flora of Eastern New Guinea, above). D. Haig (UPNG) examined Foraminifera in the 1970s and 1980s. All these (with other contributions too numerous to mention here) have now provided a clearer, though still sketchy, view of the pre-Tertiary, Tertiary, and Quaternary paleobiotic history of New Guinea in train with contemporary changes in geological and biological thought.

Collections

major collections of new guinea biota Collections within Papua or Papua New Guinea are underlined. For plants abbreviations in parentheses follow usage in Index Herbariorum I: Herbaria of the World (8th ed. 1990, New York). An asterisk (*) indicates institutions with more or less significant holdings of pre-1942 plant material. (The majority of the Berlin herbarium was lost or destroyed during World War II, but pteridophytes, a few flowering plant families, and many types are intact, as well as replicates, e.g., of Clemens, not yet distributed before the war. Plants were also lost from the British Museum (Natural History) in 1940 likewise due to bombing.) All groups (mainly historical collections): MNHN, Paris. Vascular plants: Arnold (A),* BMNH (BM),* Berlin (B), Bishop (BISH), Bogor (BO),* Queensland Herbarium, Brisbane (BRI),* CSIRO, Canberra (CANB), Edinburgh (E), Firenze (FI),* Fort Worth, Texas (BRIT), Gray Herbarium (GH),* Kew (K), Lae, PNG (LAE),* Leiden (L),* Manokwari, Papua (MAN), Melbourne (MEL),* New York Botanical Garden (NY),* Sydney (NSW),* USNM (US),* Waigani (Port Moresby), PNG (UPNG), Utrecht (U),* Wroclaw, Poland (WRSL).* Non-vascular plants and fungi (in addition to institutions indicated above): Farlow (FH),* Geneva (G),* Tokyo (TNS), and Nichinan, Japan (NICH), general; Bulolo (now Lae) and Konebobu (Port Moresby), PNG, fungi; Budapest (BP) and Uppsala, lichenized fungi (lichens); Helsinki (H) and Jena (JE), bryophytes; Berkeley, California (UC), algae. Vertebrates: Abepura (Papua), Berlin, Bogor, Genoa, Waigani (general); AMNH, BMNH, Bishop (mammals); AMNH, BMNH, Bishop, CSIRO, Genoa (birds); AM, AMNH, Bishop, Leiden, MCZ (reptiles); AM, AMNH, Adelaide, Bishop, Leiden (amphibians); AM, Leiden, BMNH, Copenhagen, Leiden, USNM (freshwater fish); Kanudi (Port Moresby), PNG (marine fish and other vertebrates). Insects and other arthropods: AM, AMNH, BMNH, Berlin, Bishop, Bogor, Bulolo (partly now Lae), CAS, CSIRO, Genoa, Copenhagen, Kanudi, Konedobu, Leiden, Lyon, Manokwari, UPNG, WEI (general); BMNH, Bishop, Leiden (Odonata); BMNH, Bishop, CNC, CSIRO, Leiden (Lepidoptera); AM, BMNH,

................. 16157$

$CH2

02-08-07 10:22:43

PS

PAGE 100

Biological Exploration of New Guinea / 101

Bishop, Dresden, Leiden (Coleoptera, Diptera, etc.); TMDU (muscoid Diptera); Adelaide, Bishop, BMNH, Leiden, USNM, Budapest, Amsterdam (arachnids and parasites); Budapest, Bishop (soil arthropods); Budapest (freshwater Crustacea). Mollusks (terrestrial): AM, BMNH, Leiden. Marine invertebrates: BMNH, Leiden, Brussels.

principal repositories AM AMNH ANU Abepura Adelaide Amsterdam Arnold Auckland BMNH

BYU Berlin

Bishop Bogor Brisbane

Brussels Budapest Bulolo

Australian Museum (incl. parts of the colls. of Macleay Museum, Sydney University) (Sydney, N.S.W., Australia) American Museum of Natural History (New York, N.Y., U.S.) Australian National University (Canberra, A.C.T., Australia) Cenderawasih University (Abepura, Jayapura, Papua, Indonesia) South Australian Museum (Adelaide, South Australia) Natura Artis Magistra (Zoological Museum) (Amsterdam, the Netherlands) Arnold Arboretum of Harvard University (Cambridge, Mass., U.S.) Auckland Institute and Museum (Auckland, New Zealand) Natural History Museum (London (South Kensington), England, U.K.) [Formerly British Museum (Natural History); name changed in 1989.] Brigham Young University (Provo, Utah, U.S.) Botanisches Museum (Free Univ.); Naturhistorisches Museum (Humboldt-Univ.) (Berlin, Germany, the Botanisches Museum in Dahlem) Bernice P. Bishop Museum (Honolulu, Hawai’i, U.S.) Herbarium Bogoriense/Museum Zoologicum Bogoriense, PPPB, LIPI (near Bogor, West Java, Indonesia) Queensland Herbarium (Mt Coo-tha); Queensland Institute of Medical Research (Herston); Queensland Museum; University of Queensland (St. Lucia) (all Brisbane, Queensland, Australia) Institut Royale d’Histoire Naturelle de Belgique (Bruxelles/ Brussel, Belgium) Termeszettudomanyi Museum (Hungarian National Museum) (Budapest, Hungary) Forestry College Herbarium (Bulolo, P.N.G.). [For Department of Forests collections of fungi and insects, see Lae. Collections of the Insect Farming and Trading Agency remain at Bulolo.]

................. 16157$

$CH2

02-08-07 10:22:44

PS

PAGE 101

102 / david g. f r o d i n

CAS CNC CSIRO

Canberra Copenhagen DAS DASF

DFMR DPI

DSIR

Dresden Edinburgh Farlow Field Firenze Fort Worth Frankfurt Geneva Gray Harvard

California Academy of Sciences (San Francisco, Calif., U.S.) Canadian National Collection (Ottawa, Canada). [Insects at Agriculture; others at National Museums of Canada.] Commonwealth Scientific and Industrial Research Organization (Canberra, A.C.T., Australia). [Includes Australian National Herbarium (Div. Plant Industry) and Div. Entomology.] See ANU, CSIRO Zoological Museum, Univ. Copenhagen (København (Copenhagen), Denmark) Department of Agriculture and Stock, P.N.G. (formerly part of DASF, then DPI). See Konedobu. former Department of Agriculture, Stock and Fisheries, P.N.G. (later DPI, now Depts. of Agriculture and Stock (DAS) and Fisheries and Marine Resources (DFMR)). See Kanudi, Konebobu. Department of Fisheries and Marine Resources (formerly part of DASF, then DPI), PNG. See Kanudi. former Department of Primary Industries, PNG (DPI, formerly DASF, now Depts. of Agriculture and Stock (DAS) and Fisheries and Marine Resources (DFMR)). See Kanudi, Konedobu. former Department of Scientific and Industrial Research (Lincoln, Canterbury, New Zealand). [Biological collections now under Landcare Research, same address.] Staatliches Museum fur Tierkunde (Dresden, Sa¨chsen, Germany) Royal Botanic Garden, Edinburgh, Scotland Farlow Herbarium of Harvard University (Cambridge, Mass., U.S.) Field Museum of Natural History (Chicago, Ill., U.S.) Museo Botanico, Univ. Firenze (Florence, Italy). [Colls. Beccari, particularly Palmae, Pandanaceae] Botanical Research Institute of Texas (Fort Worth, Texas, U.S.) Senckenbergisches Institut und Museum (Frankfurt-amMain, Germany) Conservatoire et Jardin Botaniques; Museum d’Histoire Naturelle (both Ville de Gene`ve, Switzerland) Gray Herbarium of Harvard University (Cambridge, Mass., U.S.) Harvard University Museums and Herbaria. See Arnold, Farlow, Gray, MCZ

................. 16157$

$CH2

02-08-07 10:22:44

PS

PAGE 102

Biological Exploration of New Guinea / 103

Helsinki Honolulu Kanudi

Kew Konedobu Lae

Leiden

MCZ MVZ Macleay Manokwari

Melbourne Mu¨nchen Nelson New York Nichinan Paris Port Moresby QIMR RSA Sydney

Botanical Museum, Univ. Helsinki (Helsinki, Finland) Hawaiian Sugar Planters Association Experiment Station, Honolulu, Hawai’i, U.S. (insects, now Bishop) collections (fish and other marine organisms) of DFMR (earlier DASF, DPI) (Kanudi, Port Moresby, PNG) [Catalogue, 1974: A catalogue of the fish reference collection at the Kanudi Fisheries Research Laboratory, Port Moresby by P.J. Kailola. DASF Res. Bull. 16.] Royal Botanic Gardens, Kew (Richmond, Surrey, England, U.K.) collections (insects, fungi) of DAS (earlier DASF, DPI) (Konedobu, Port Moresby, PNG) Herbarium, Div. Botany, Forestry Research Institute (FRI); also collections (formerly at Bulolo) of forest fungi and insects, FRI (Lae, Morobe Province, PNG) National Herbarium of the Netherlands (Leiden branch); Naturalis (formerly Rijksmuseum van Natuurlijke Historie) (Leiden, the Netherlands) Museum of Comparative Zoology, Harvard University (Cambridge, Mass., U.S.) Museum of Vertebrate Zoology, University of California, Berkeley (Calif., U.S.) See AM. [Some material remains in Macleay.] Agriculture College of Cenderawasih University; Forestry and Agriculture Experiment Station (Amban, Manokwari, Papua, Indonesia) National Herbarium of Victoria (South Yarra); Museum of Victoria (Melbourne, Vic., Australia) Zoologische Staatsammlung der Bayerische Staat (Munich, Bavaria, Germany) Cawthron Institute (Nelson, New Zealand; now part of Landcare NZ (formerly DSIR). [Insects to Auckland.] New York Botanical Garden (Bronx, New York, N.Y., U.S.) Hattori Botanical Laboratory (Obi, Nichinan, Miyazaki Pref., Japan) Muse´um National d’Histoire Naturelle (Paris, France) See Kanudi, Konedobu, Waigani Queensland Institute of Medical Research. See Brisbane Rancho Santa Ana Botanic Garden Herbarium (Claremont, Calif., U.S.) Australian Museum (see also AM); National Herbarium of New South Wales; University of Sydney, Newtown (all Sydney, N.S.W., Australia)

................. 16157$

$CH2

02-08-07 10:22:45

PS

PAGE 103

104 / david g. f r o d i n

TMDU Tokyo Toulouse Tring

UC UniTech UPNG

USNM

Uppsala Utrecht WEI Waigani

Wellington

Tokyo Medical and Dental University (Bunkyo-ku, Tokyo) (Muscoid Diptera) National Science Museum (Shinjuku-ku, Tokyo, Japan) Universite´ Paul Sabatier (Toulouse, France) (Collembola) Rothschild Museum (Tring, Herts., England, U.K.), now a BMNH branch. [Most bird collections sold in 1932 to AMNH. Remainder of museum merged with BMNH in 1937 under Rothschild’s will. Insects now in London; BMNH birds at Tring.] University of California Herbarium (Berkeley, Calif., U.S.) P.N.G. University of Technology (Lae, Morobe Prov., PNG) Natural Sciences Resource Centre, University of Papua New Guinea (Waigani, Port Moresby, PNG) [Includes herbarium along with entomological and zoological collections.] National Museum of Natural History, Smithsonian Institution (Washington, D.C., U.S.). [Includes U.S. National Herbarium as well as entomological and zoological collections.] Evolutionary Biology Centre, Univ. Uppsala (Uppsala, Sweden) National Herbarium of the Netherlands (Utrecht branch) (Utrecht, the Netherlands) Wau Ecology Institute, Wau, Morobe Prov., PNG. [Formerly Bishop Museum Field Station.] Papua New Guinea National Museum and Art Gallery (Waigani, Port Moresby, P.N.G) (Vertebrates); University of Papua New Guinea (see UPNG) National (formerly Dominion) Museum (Wellington, New Zealand)

Selected References Because a bibliography of all historical sources of information as well as the results of collecting mentioned in this review would fill a large volume, most works mentioned by author or title in the text are not repeated here. Only some more general sources are accounted for below.

bibliographies The most substantial older twentieth-century bibliographies are Toa kyo-ei-ken sigenkagaku bunken mokuroku 1: [New Guinea] (1942, Dept. Education, Japan) and An annotated bibliography of the southwest Pacific and adjacent areas 1 and 2 (1944, Allied Geographical Section) along with a general bibliography in five volumes issued in the 1980s by the University of Papua New Guinea Library. For Papua there is also West Irian: a bibliography (1984, Dordrecht: Foris, as KITLV

................. 16157$

$CH2

02-08-07 10:22:45

PS

PAGE 104

Biological Exploration of New Guinea / 105

Bibl. Ser. 15) by J. van Baal, K. W. Galis, and R. M. Koentjaranigrat (a revision of Bibliographie van Nederlands-Nieuw-Guinea (1962) by Galis). For specific biota there have been published Bibliography of New Guinea entomology (1968, as Pacific Insects Monogr. 18) by J. L. Gressitt and J. J. H. Szent-Ivany; Bibliography of invertebrate animals from New Guinea (1973, in Science in New Guinea 1: 41–46) by W. H. Ewers; Papua New Guinea fisheries bibliography (1985, Port Moresby, as Dept. Primary Industry Tech. Rept. 85/3) by J. M. Lock and D. C. Waites; Bibliography of freshwater ecology in Papua New Guinea (1988, Waigani (Port Moresby), as Dept. Biology (UPNG) Occ. Pap. 9) by P. L. Osborne; and A bibliography of the flora and vegetation of Papua New Guinea (1996, in Papua New Guinea Journal of Agriculture, Forestry and Fisheries 39(2): 20–168) by S. Saulei. The entomological bibliography contains taxonomic and subject indices; those of Lock/Waites and Saulei have broadly circumscribed subdivisions. More or less extensive lists of references furthermore appear in several of the other works listed here.

encyclopedic works, general monographs, and other reviews General encyclopedic works on, or inclusive of, New Guinea are Neu-Guinea ([1899], Berlin: Schall, in their series Bibliothek der La¨nderkunde), ed. M. Krieger; Deutsches Kolonialreich (1910, Leipzig), ed. H. J. Meyer; Deutsch-Neu-Guinea (1911, Berlin: Reimer) by R. Neuhauss; Deutsches Kolonial-Lexikon (1920, Berlin) [now available on the Web]; Nieuw-Guine´e (1935–38, the Hague) and its second edition, Nieuw-Guinea (1953–1954, the Hague), both ed. W. C. Klein and islandwide in coverage; and Encyclopaedia of Papua New Guinea (1972, Melbourne). Very important for biota in general is Biogeography and Ecology of New Guinea (1982, Junk, as Monographiae biologicae 42), ed. J. L. Gressitt; a wide range of groups (and some ecosystems) is encompassed. Earlier coverage for flora and vegetation may be found in an essay by H. J. Lam in Nieuw-Guine´e (1: 187–210) and one (with references) by C. G. G. J. van Steenis in its successor (2: 218–275); others have appeared since, including Documentation of the flora of New Guinea (pp. 123–156) by B. J. Conn in Biodiversity and terrestrial ecosystems (1994, Taipei, as Mon. Inst. Bot. Acad. Sin. 14), ed. C.-I. Peng and C.-H. Chou. For entomology reference may be made to La Nouvelle-Guine´e Australienne: introduction ´ecologique et entomologique by P. Jolivet (1971; in Cahiers du Pacifique (Paris) 15: 41–70). More recent reviews, with particular reference to conservation, include Papua New Guinea Conservation Needs Assessment (1993, [Port Moresby]), ed. B. M. Beehler; Papua New Guinea Country Study on Biological Diversity (1994, [Port Moresby]), ed. N. Sekhran and S. Miller; and Lokakarya penentuan prioritas konservasi keanekaragaman hayati Irian Jaya [Biak, 7–12 January 1997]: laporan akhir/ The Irian Jaya Biodiversity Conservation Priority-Setting Workshop [Biak, 7–12 January 1997]: Final Report (1999, Washington, D.C.), ed. J. Supriatna. All are generously furnished with tables and many maps from which some idea of the present state of knowledge can be gained; they form parts of chapters (or sections) on biodiversity (the 1993 report features treatments of individual biotic groups or

................. 16157$

$CH2

02-08-07 10:22:45

PS

PAGE 105

106 / david g. f r o d i n

environments). For plants may be added the chapter on New Guinea by P. F. Stevens (pp. 120–132) in Floristic inventory of tropical countries (1989, New York), ed. D. G. Campbell and H. D. Hammond.

serial publications and periodicals Important general series are Nova Guinea (1909–1966, Leiden); Results of the Archbold Expeditions (American Museum of Natural History, and (for plants) the New York Botanical Garden and the Arnold Arboretum of Harvard University)—the papers in the AMNH Bulletin being numbered; the several numbers of the CSIRO Land Research Series pertaining to present-day Papua New Guinea; the Christensen Research Institute Publications; and the Wau Ecology Institute Handbooks. Useful for exploration and topography (and rich in maps) are Mitteilungen aus den deutschen Schutzgebieten (1888–1929, Berlin) and its Erga¨nzungshefte (1908–1930, Berlin) and, for German New Guinea, Nachrichten u¨ber Kaiser-Wilhelms-Land und den Bismarck-Archipel (1885–1998). For British New Guinea (and Papua) the Annual Reports for 1884–1885 to 1913–1914 similarly contain much that is useful, with some specifically biotic appendices. Current and past outlets in Australia include the publications of Queensland Museum, Queensland Herbarium (Austrobaileya), University of Queensland, Australian Museum, South Australian Museum, National Herbarium of New South Wales (Telopea), Australian Journal of Systematic Botany, and the Linnean Society of New South Wales. Elsewhere, for Asia Gardens Bulletin, Singapore, and Reinwardtia and its predecessors (Bogor) may also be mentioned, along with the Bulletins of the National Science Museum (Tokyo) and Journal of the Hattori Botanical Laboratory. In Britain Kew Bulletin as well as the Bulletins of the British Museum (Natural History)/ Natural History Museum have run results based in whole or in part on collections from New Guinea. On the European continent Flora Malesiana (1948–, now based at Leiden) is a key reference but still far from complete; for the nearer term, much research has appeared in the serial Blumea, another Leiden-based publication. Beitra¨ge zur Flora von Papuasien (1912–1942, with 150 contributions in Botanische Jahrbu¨cher fu¨r Systematik, Pflanzengeschichte und Pflanzengeographie) remains historically important, along with some other German publications including Pflanzenreich as well as those from the Museum fu¨r Naturkunde in Berlin. For bryophytes, Acta Botanica Fennica (Helsinki) embodies a substantial amount of recent research. In the United States of America, for entomology mention should be made of several Bishop Museum series (Pacific Insects; Pacific Insects Monographs; Journal of Medical Entomology; and Insects of Micronesia). For zoology in general there have been many outlets, but because of the Archbold Expeditions the Bulletin of the American Museum of Natural History has been very significant. Local journals include those of the (now defunct) Papua New Guinea Scientific Society; the PNG Agricultural Journal (now PNG Journal of Agriculture, Forestry and Fisheries); the PNG Medical Journal; and Science in New Guinea. The PNG

................. 16157$

$CH2

02-08-07 10:22:46

PS

PAGE 106

Biological Exploration of New Guinea / 107

Wildlife Division reports as well as the Botanical Bulletins of the PNG Division of Botany may also be mentioned.

exploration For general exploration through the early 1960s, the best single reference is New Guinea: the last unknown by G. Souter (1963, Sydney). A useful earlier review (to 1934) by C. C. F. M. le Roux appears in Klein’s Nieuw-Guine´e (1, 1935; see above). Valuable for expeditions, individuals and localities up to 1902 is the very detailed Entdeckungsgeschichte von Neu-Guinea by A. Wichmann (1909–1912, Leiden; in Nova Guinea, 1–2). All three works are well documented. Botanical exploration (for Malesia in general) is covered by M. J. van SteenisKruseman in series I of Flora Malesiana (1, 1950; 5, 1958; 7, 1974) with later coverage in Flora Malesiana Bulletin. For zoology, entomology, and marine biology, there is no single source, although monographs and reviews of major groups sometimes include historical sections. ‘‘Bioinventory’’—a current successor to ‘‘primary’’ and most ‘‘secondary’’ biotic exploration—is usefully aired in W. Takeuchi and M. Golman, Floristic documentation imperatives: some conclusions from contemporary surveys in Papua New Guinea (in Sida 19: 445–468. 2001) and A. Allison, Biological surveys—new perspectives in the Pacific (in Organisms, Diversity and Evolution 3: 103–110. 2003). Finally, reference may be made to The natural world of New Guinea: hopes, realities and legacies by the present writer (pp. 89–138 in Nature in its greatest extent: Western science in the Pacific (1988, Honolulu), ed. R. MacLeod and P. F. Rehbock).

Acknowledgments For their 1982 chapter, Frodin and Gressitt were indebted for assistance to Drs. F. R. Fosberg, L. B. Holthuis, P. Raven, P. van Royen, J. J. H. Szent-Ivany, and to curators and librarians of the Bishop Museum, California Academy of Sciences, American Museum of Natural History, Museum of Comparative Zoology, the University of Papua New Guinea, the Rijksherbarium (Leiden), and others. The present revision has been entirely the work of the writer, who here acknowledges the facilities of the Herbarium and Library of the Royal Botanic Gardens, Kew, as well as some assistance through personal communications.

................. 16157$

$CH2

02-08-07 10:22:46

PS

PAGE 107

1.3. The Socio-cultural Plurality of Papuan Society j. r. mansoben The Socio-cultural Diversity of Papuan People h e pro v i nc e o f pa p u a (as construed to include all of western New Guinea) is the largest in Indonesia (416,000 km2, or three times the size of Java). Within this expansive area, two million people live at the lowest population density in Indonesia (approximately 4 persons per km2; 1999 census data). Despite this relatively small population size, Papua exhibits a much greater diversity of ethnicities and cultures than any other Indonesian province. This chapter is an overview of the variation in language, social structure, leadership systems, religion, livelihood systems, land tenure system, orientation of cultural values, and work ethic in this highly diverse province. Although a complete review of these diverse elements would fill many volumes, the information given here may help provide some insight into Papua’s rich cultural and social heritage, and may assist in the design of suitable and sustainable programs for the development of Papua and its peoples.

T

Languages According to language experts at the Summer Institute for Linguistics, approximately 269 living local languages are spoken in Papua (Ethnologue Website). Language provides a means of communication as well as a symbol of group identity, suggesting that Papua contains a minimum of 269 distinct ethnic groups. Papua’s languages are typically classified into two large groups, or mother languages: Austronesian and Non-Austronesian (often called Papuan). The Austronesian mother language group is comprised of languages spoken by coastal communities (e.g., Biak, Wandammen, Waropen, Maya). The Non-Austronesian (or Papuan) language group contains languages spoken by people that live in remote areas in the center of the island, from the western Vogelkop to the eastern tip of New Guinea (e.g., Meybrat, Dani, Ekari, Asmat, Muyu, and Sentani). The Papuan language groups are divided into ten phyla: The Trans New Guinea Phylum, West Papuan Phylum, Sepik-Ramu Phylum, Torricelli Phylum, Sko Phylum, Kwomtari Phylum, Arai (Left May) Phylum, Amto-Musian Phylum, Geelvink Bay Phylum, and East Bird’s Head (Vogelkop) Phylum. These phyla are further split into language families. Hence, one phylum consists of several language families, each containing several local languages or dialects. This language classification was initiated by Voorhoeve and McElhanon. The high linguistic diversity on Papua provides a unique opportunity for the study of languages and language evolution. Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

108

................. 16157$

$CH3

02-08-07 10:22:06

PS

PAGE 108

Socio-cultural Plurality of Papuan Society / 109

The diversity of languages in Papua poses a challenge to development efforts in this area, since mutual understanding and communication is difficult when so many different languages are spoken in one province. However, the diversity of languages in Papua has led in recent decades to the widespread adoption of Bahasa Indonesia (and Melayu languages in the past), which serves as an intermediary language that is used among different ethnic groups within Papua and with people from other parts of Indonesia. Therefore, despite the high linguistic diversity in Papua, most Papuans speak and understand Bahasa Indonesia. Indeed a higher proportion of Papuans are fluent in this language than people in most other Indonesian provinces. This common language helps to offset some of the challenges inherent to the high linguistic diversity in Papua, and should aid development efforts in Papua in the future.

Social Structure In this context, social structure refers to the patterns of social relationships that maintain group cohesion and social unity. Typically, these patterns of social relationships are organized around kinship, and can be characterized by the terminology used to refer to family members and their inheritance systems. Kinship terminology is extremely useful in understanding social structure because kinship terms frequently convey details about the roles of family members and the interactions among them, and about social rights and responsibilities, all of which may differ greatly among groups. Pouwer (1966) suggests that the people of Papua can be divided into at least four groups based on their systems of kinship terminology. The first group uses a kinship terminology system similar to the Iroquois, a Native American nation (Iroquois type). The Iroquois system classifies cousins in parallel with siblings, and uses a different expression for cross-cousins. Another characteristic is the use of the same expression for father and all male brothers from both maternal and paternal sides (uncles). The languages of Biak, Iha, Waropen, Senggi, and MarindAnim, Humboldt (Yos Sudarso) Bay, and Me are included within this group. The second group uses kinship terminology similar to those used by native peoples in Hawai’i (Hawai’ian type). In this system, the same expression is used for siblings and all parallel and cross-cousins. Ethnic groups using this kinship terminology include Mairasi, Mimika, Hattam-Manikion, Asmat, Kimam, and Pantai Timur Sarmi people. The third group uses the Omaha type kinship system. Omaha type is a system that uses different terms for matrilineal and patrilineal cross-cousins and incorporates information about generations into their kinship terminology in an asymmetric way. On the maternal side, cross-cousins are raised a generation while those on the paternal side are lowered a generation. Hence, the expression for mother’s brother’s son is the same as mother’s brother and the expression for father’s sister’s son is the same as sister’s son. Included in this group are the people of Auwyu, Dani, Meybrat, Mek in the Star Mountains, and Muyu.

................. 16157$

$CH3

02-08-07 10:22:06

PS

PAGE 109

110 / j. r. mansoben

The fourth group contains people that use the Iroquois-Hawai’ian type of kinship terminology. Included in this system are people of Bintuni, Tor, and West Coast of Sarmi (Pouwer 1966). Papuan peoples can also be classified according to which of two major inheritance systems they recognize. The first inheritance system used in Papua is patrilineal, where inheritance is from father to son or among other male kin. This system is used by the people of Meybrat, Me, Dani, Biak, Waropen, Wandammen, Sentani, Marind-Anim, and Nimboran. The second major inheritance system used in Papua is the matrilineal system, in which inheritance is passed through the female kin. Some Papuan people use a system that is intermediate between these two major types. In such bilateral systems, inheritance is either through the father’s or the mother’s kin. This type is used by remote communities in Sarmi. Similarly, some communities practice ambilateral or ambilineal structure, for example in the communities of Mimika, Mappi, and Manikion, inheritance is sometimes through the maternal line (Mappi and Mimika) and sometimes through the paternal line (Manikion) according to individual choice (Bruijn 1959; Pouwer 1966). One additional notable characteristic differs among social structures found in Papua. Some peoples group the community into a phratry (a group of clans tracing descent to a common ancestor) while others practice moiety (dividing the group into two halves for ritual purposes). Among Papuan people who use moiety groups are Asmat (aipmu and aipem), Dani (waita and waya), and Waropen (buriworai and buriferai). However, there are also ethnic groups that do not recognize this principle, for example the people Muyu and Biak (Heider 1979; Held 1947; Kamma 1972).

Land Tenure Systems Two major types of property rights and land use practices are common among Papuan peoples: communal ownership systems and individual ownership systems. In communal ownership systems the land that provides the main resources necessary for livelihood are owned communally. Two types of communal ownership systems are found in Papua, those based on small clans or lineages and those based on large clans or villages (kampung). In the communal ownership system based on clans, all members of the clan (marga, keret), including unmarried women, have equal rights to use the clan lands for their livelihood. Although all members have the same rights in principle, individuals do not have the freedom to decide where they want to conduct their economic activities (e.g., to open new agricultural land or collect certain forest products). The clan head (kepala marga) regulates and monitors the clan’s land use, but often makes decisions about land usage in conjunction with other clan members. Importantly, no member of the clan (including the head) has the authority to cede ownership of the land to an outside party (e.g., the government or a private company). Such decisions must be made communally, and any proceeds

................. 16157$

$CH3

02-08-07 10:22:07

PS

PAGE 110

Socio-cultural Plurality of Papuan Society / 111

from the sale or lease of such land are shared equally among all clan members. Clan-based communal ownership system can be found in the following ethnic groups: Dani, Meybrat (Ayamaru), Muyu, Marind-Anim, Auwyu, Wandammen, Simuri, Irarutu, Biak, and Waropen. In large clan (kampung) based communal ownership systems, land ownership rights are held by the community head, who has the authority to make land-use decisions in conjunction with clan leaders (e.g., in Sentani the authority to manage the land is jointly held by community heads (yo-ondoafi) and clan heads (khoselo). Neither clan nor community leaders can make decisions alone, and plans for community development, land use, and ownership must be made jointly. Any proceeds from land sale are distributed within the village according to the internally recognized differences in land rights and authority of different members. Frequently, individuals deliberately ‘‘forget’’ the relative authority or rights of a particular individual, which can lead to conflict within the community.

Political Systems Political systems are also highly variable in Papua. To understand the traditional political systems used by the Papua people, Mansoben (1985) applied the continuum model suggested by Sahlins (1963) to available ethnographic data and identified four political systems in Papua. These four systems are big man (or powerful man) systems, kingdom systems, ondoafi systems, and mixed systems. Sahlins (1963) suggested that political systems could be analyzed along a continuum. On one end of the continuum the political system is characterized by ascription or inheritance; while on the other end the political system is characterized by achievement. On the ascription end of the continuum are chief (head of ethnic group) systems, while on the achievement end are the big man systems. Applying this continuum to the political systems in Papua, Mansoben (1985) found that in addition to the two systems at the extremes of the continuum, some Papuan political systems contain elements of both systems and belong somewhere in between. These systems are called mixed systems. Furthermore, two distinct types can be identified in Sahlin’s ascription leadership system: kingdom and ondoafi. The major differences between Papuan political systems are the geographical extent of their power and their political orientations. Below I briefly discuss the principle characteristics and major differences among the four most common political systems in Papua. In big man political systems the leadership position is based on individual achievement. The source of power in this political system is derived from the big man’s personal abilities or achievements, such as success in allocating and distributing wealth, diplomatic or oratory skills, courage on the battlefield, physical strength, or generosity (Sahlins 1963; Koentjaraningrat 1970). Big men characteristically hold a substantial amount of personal power, and have autonomy to make important decisions single-handedly. The Dani, Asmat, Me, Meybrat, and Muyu peoples have big man political systems.

................. 16157$

$CH3

02-08-07 10:22:07

PS

PAGE 111

112 / j. r. mansoben

The kingdom system is primarily characterized by ascribed political status or inherited leadership position. Political power is conferred on individuals due to their family membership and birth order. In kingdom systems power is passed patrilineally, and patrilineal lines form traditional bureaucracies in which individuals have clearly-defined roles, responsibilities, and authority. All positions of authority are passed down through the male line, and if a man’s first son is inappropriate for the job, it is passed to another clan member. The kingdom system is common among communities in southwest Papua, including the Raja Ampat Islands, Onin Peninsula (on Bomberai peninsula), McCluer Gulf (Berau Gulf), and Kaimana. The ondoafi system is similar to the kingdom system in that leadership positions are inherited and traditional bureaucracies are utilized. However, the ondoafi system differs from the kingdom system in its geographic range of power and political orientation. The power of an ondoafi leader is limited to a single village (yo), and the social unit consists of one ethnic group or subgroup. In contrast, the authority of leaders in kingdom systems is not limited to one village, but covers a wider area. Also, in kingdom systems the social units consist of several ethnic groups. Other characteristic of ondoafi is its alliance system, in which several villages act as a unit, trace their ancestry to a single individual, and acknowledge a single leader for the larger community. This ‘‘great leader’’ typically comes from a village in the center of the geographic range encompassed by the larger community. Finally, while the principle focus of the kingdom political system is on trade, in the ondoafi system the center of political orientation is religion. Ondoafi systems are practiced in northeast of Papua, by the people of Sentani, Genyem (Nimboran), Humboldt (Yos Sudarso) Bay, Tabla, Yaona, Yakari-Skou, and Arso-Waris. The final political system found in Papua is the leadership mixed system, in which leadership is obtained through either inheritance or achievement. In other words, an individual can be a leader based on his personal ability, achievement, or birthright. Leaders that gain their authority based on achievement usually appear during times of stress, such as war, famine, epidemic disease, or cultural decadence. Such leaders are known as situational leaders, since the leader is chosen based on his ability to overcome the particular challenge facing the community. In mixed systems power is usually inherited during ‘‘safe’’ times, when external and internal threats are low. During such stable times, the leaders are chosen from the traditionally powerful family. In contrast to the kingdom and ondoafi systems, bureaucracy is not found in mixed systems. Mixed systems are common in the people who live around Cenderawasih Bay, such as the people of Biak, Wandammen, Waropen, Yawa, and Maya.

Religion and Belief Systems Before Islam and Christianity were introduced to Papua, each ethnic group had its own traditional belief system. Although traditional belief systems varied among groups, most groups believed in a single Goddess or God that held supreme power

................. 16157$

$CH3

02-08-07 10:22:07

PS

PAGE 112

Socio-cultural Plurality of Papuan Society / 113

over other deities. This God or Goddess had different names in different groups. For example in Biak-Numfoor culture this highest Goddess is called Manseren Nanggi or the Sky God, Moi people recognize Fun Nah, Seget people use the name Fun Naha, Waropen people refer to their supreme god as Naninggi, Wandammen recognize Syen Allah, Marind-Anim people Dema, Asmat people Mbiwiripitsy, and the Me people Ugatame. Ethnographic accounts of traditional belief systems in Papua indicate that the principal Goddess or the Highest God is considered to be the creator and to have absolute authority over human destiny. In addition, most followers of these traditional religious systems believe that the power of this God is manifest in natural forces, such as wind, rain, and thunder; or believe that the power resides in natural objects near human settlement, such as large trees, streams, river currents or eddies, the bottom of the sea, or certain bays. Since it is believed that these spirits have the power to control people’s lives, they are feared and respected. Various activities express this fear and respect, in the form of offerings and rituals. These behaviors demonstrate an acknowledgement of the existence and power of the spirits and are thought to foster good relationships between humans and the spirits. The Papuan people also believe that the spirits of dead ancestors are given power by the Creator God to control people that are still alive. Hence, the living must maintain positive relationships with their ancestors to protect themselves from disasters that may occur if deceased relatives are angered. This is the basis for ancestor worship, which is expressed in various forms, for example, the praise of korwar statues and the mon ceremony in Biak-Numfoor culture, the skeleton payment ritual of Meybrat people, the mbis ceremony of the Asmat. Since the arrival of Islam and Christianity traditional practices, including ancestor worship, are becoming less common. However, when traumatic events occur, such as accidents, illness, and death, many Papua people still seek solace and inspiration from traditional practices. Major religions, such as Islam and Christianity, arrived in Papua at different times. Islam entered Papua first, brought to the Raja Ampat Islands and Fakfak by traders from the Maluku Islands in the 13th century (Leeden 1980). Christianity was introduced to Papua in Mansinam Island on February 5, 1855, by two missionaries, Ottow and Geissler who were sent by the Dutch Bible institution Utrechtsche Zendingsvereniging (Kamma 1953). Catholicism came to Papua in 1892. Other Christian denominations, such as Pentecostal, Adventist, and Christian and Missionary Alliance (CMA), came to Papua after World War II. Hinduism entered Papua in the 1960s. According to 1980 census data, 12% of Papuans were Islamic (132,930 people), 23% were Catholic (256,209 people), and 65% were from other Christian denominations (768,279 people).

Ecology and Subsistence Systems In Papua four broad categories of ecological environments can be recognized: swampy areas, coastal lowlands, foothills and small valleys, and highlands (Walker

................. 16157$

$CH3

02-08-07 10:22:08

PS

PAGE 113

114 / j. r. mansoben

and Mansoben 1990). Each of these zones supports different subsistence systems. In swampy areas people primarily depend on sago as a staple food, supplemented by fish (e.g., Asmat, Mimika, and Waropen). For people living in coastal and riverine zones (e.g., Biak, Wandammen, Moi, Simuri, Maya, the Raja Ampat Islands), fishing, sago cultivation, and agriculture are the primary economic activities, while hunting serves as an alternative strategy. People living in the foothills and small valleys depend primarily on agriculture and sago, supplemented by hunting and animal farming (e.g., Muyu, Genyem, and Arso). Finally, in the highlands, farming and raising pigs is the primary subsistence strategy (e.g., Dani and Me). The natural environment influences the technological adaptations and culture (e.g., social organization, belief systems) of the people living there. For example, in the highlands zone, communities live in large houses and have relationships with their extended families, forming clan networks and more complex federations (e.g., Dani). In coastal areas, islands, and riverine zones, people tend to live in small, independent, nuclear family groups of four to five individuals (e.g., North Coast; Koentjaraningrat 1970), although some groups (e.g., the people of Kimaam in Kolepom [Yos Sudarso] Island, and Meren-Vlakte) the people live in more extended families (about 10–15 individuals; Koentjaraningrat 1970). For several decades researchers have examined the relationship between ecological environments and the diversity of Papuan people. J. van Baal observed that Papuan people who subsist on sago in swampy and riverine zones tend to have larger and more frequent celebratory religious ceremonies compared to the Papuan people who eat root plants or live in the highland zone, and implied that the complexity of rituals and belief systems of the Papuan people is influenced by environmental conditions. Ecological factors also influence the degree of mobility of groups. V. de Bruijn suggested that the inability of the Biak-Numfoor Islands to support people led the communities that lived there to sail, trade, and headhunt along the coasts of north coastal Papua and eastern Indonesia (Maluku and Sulawesi Islands), following which they resettled in various places in north coastal Papua (e.g., Vogelkop, Raja Ampat Islands, Halmahera; de Bruijn 1959). Similarly, W. H. K. Feuilletau de Bruijn argued that the ecology of the Biak-Numfoor Islands prohibited productive farming, and led the peoples living there to develop a more advanced knowledge of astronomy and boat building than is found anywhere else in Papua.

Population Distribution Census data from 1995 showed that Papua was inhabited by 2,031,620 people, 27% of whom lived in cities and 73% of whom lived in villages, indicating that the majority of Papuans live in rural areas. A slight change in population distribution is indicated by census data from 2000, when 39% lived in towns and 61% lived in rural areas (BPS 2002). The population distribution is changing rapidly,

................. 16157$

$CH3

02-08-07 10:22:09

PS

PAGE 114

Socio-cultural Plurality of Papuan Society / 115

as government has developed new regencies (kabupaten) from formerly 13 regencies to 26 regencies and municipalities now.

Philosophy of Life Cultural values, manifest in social norms, ethics, regulations, and laws differ substantially among communities. A characteristic that may be highly valued by ethnic group A may not be considered good by ethnic group B; important obligations recognized by ethnic group C may not considered important for the ethnic group D, and so on. Nevertheless, it has been suggested that different cultures can be characterized by their approach to five basic concepts. Concept of the Meaning of Life All cultures in the world have their own concepts about the meaning of life, the ultimate purpose of existence, and how the journey of life should be traveled. Religions usually provide guidance that shape beliefs about the meaning of life. Concepts of the meaning of life vary tremendously; for example, some people view life as a misery that cannot be avoided, others see life is a way to redeem past sins, still other as an opportunity to accept and cherish oneself and others regardless of faults. Perception of Work Cultures differ widely in their views on the meaning of work. In some cultures work is centrally important, and is one of the ways in which one finds meaning in life. Other cultures view work as a way to gain respect from fellow citizens, still others view it as a way to serve others. Concept of Human’s Relationship with Nature Some cultures view nature as a resource solely for human use, others believe that there must be a balance between humanity and the natural world, and that natural laws need to be obeyed to maintain harmony. Still others see nature as an allpowerful force to which humans must submit. Perceptions of Time Various cultures have various perceptions of time, particularly in regards planning for the future. Some peoples focus on the present, and have a relatively narrow conception of time. Others are more oriented to the future, and place value on planning for events to come. Perceptions of Fellow Humans There are cultures that highly value the vertical relationships in society. Such cultures respect leaders and senior people, and look to them as guides for the community. Other cultures have a more horizontal view, and focus on interpersonal

................. 16157$

$CH3

02-08-07 10:22:10

PS

PAGE 115

116 / j. r. mansoben

relationships. Some cultures emphasize self-reliance, while others stress the interdependence of people and the need to cooperate. These five principles can be used to assess the attitudes of people or communities, and are especially important in determining how a culture or community feels about the future, and how they view interactions with the outside world. These are important considerations for those interested in community development. Koentjaraningrat noted that some cultural values are particular assets that facilitate future community development: (1) orientation towards the future; (2) intention to explore the natural environment; (3) placing a highly value on human work; and (4) consideration of fellow humans (Koentjaraningrat 1974).

cultural values of papuan people Various analyses have suggested that Papuan culture is not particularly innovative when it comes to nature exploration. This may be due to the traditional (and still active) belief that nature has spiritual powers over human life and destiny that are to be feared and respected. This cultural value may contribute to the passivity towards the natural environment. In one way, this passivity is beneficial for nature preservation, as it suggests a harmony with nature that is largely environmentallyfriendly. In other ways, this belief may hinder creativity and innovation. For example, although knowledge of the natural world (e.g., traditional medicine) is substantial in many Papuan cultures, this knowledge is typically limited to a few individuals or ethnic groups and is not developed or refined using modern scientific techniques. This limits the scientific, economic, and health benefits that can be derived from utilization of the natural world. However, this generalization is by no means universally true, as some Papuan cultures (e.g., those in Cenderawasih Bay) do have a history of exploration of the environment and expeditions within and outside of Papua. The tendency towards exploration led to expertise in the technology of shipbuilding and navigation in these cultures, as noted above. Although many Papuan cultures appear to be relatively passive towards largescale exploration of nature, they do highly value individual efforts and respect individuals who are industrious and innovative. The cultural value placed on individual effort encourages many Papuans to work harder, which in turn benefits the group as a whole. It also builds a sense of independence and self-confidence in some individuals, and in others builds a sense of responsibility. As mentioned above, this cultural value can be a major asset for community development and improvement. If we use the traditional leadership systems and cultural values as a ‘‘window’’ into Papua’s culture, we can see that the premium placed on hard work and industriousness is beneficial in many Papuan cultures. For example, as discussed above, in many big man systems (e.g., Meybrat, Me, Muyu, Dani, and Asmat) the leader gains power through personal abilities and achievements (achieved status). In the Meybrat culture, a person is highly respected if he is successful in devising and implementing systems for the exchange of ceremonial cloth. A successful Meybrat

................. 16157$

$CH3

02-08-07 10:22:10

PS

PAGE 116

Socio-cultural Plurality of Papuan Society / 117

man would be called bobot to acknowledge this achievement. Similarly, in the Me people a person who has good diplomatic skills, is kind and honest, and economically successful (i.e., acquires much agricultural land, many pigs and wives, and cowrie shell ‘‘money’’) is highly respected and recognized as the leader of the community. Such a person is known as leader (tonowi) and rich man (sonowi). The Muyu people respect and confer power upon men with talents in organizing large pig feasts and ceremonies. In cultures where warfare is of central economic and ritual importance (e.g., Dani, Asmat), men with courage and strength during battle are respected and become leaders. Anthropological analysis of Papuan cultures shows that two very different attitudes towards interpersonal relationships are found in Papua. First, some cultures are strongly vertically-oriented. Cultures with the kingdom leadership system (e.g., those on Onin (part of the Bomberai) Peninsula, Kowiai area, the Raja Ampat Islands) strongly exhibit this orientation, as do cultures in northeast Papua that practice the ondoafi leadership system (e.g., Tabla, Skow, Nimboran, Sentani, and the people of Yos Sudarso Bay). In these cultures, the leader is viewed as a descendant of a mythical ancestor who plays a special role as mediator between the real and the supernatural worlds. As these leaders are believed to have magical powers, they are widely respected and consulted by all community members. Koentjaraningrat (1974) believed that vertically-oriented systems tend to reduce an individual’s sense of independence and self-confidence. He also argued that these systems reduce self-discipline, since individuals can come to feel that they must only obey cultural rules and laws only if they are being watched by more powerful individuals. Finally, leaders having considerable power to dictate the lives of other community members can reduce the sense of responsibility felt by those with little power (Koentjaraningrat 1974). Although some of the byproducts of such systems (e.g., lack of discipline and responsibility, lack of innovation) are detrimental to community development, there are some positive outcomes as well. For example, a strong leader can efficiently organize and motivate community members to participate in development projects. A second view of interpersonal relationships has a more horizontal orientation, as noted above. In cultures that hold such beliefs (e.g., Biak), the relationships among the community members in a clan are very strong, and group needs are prioritized above individual needs. Among the clan members, solidarity is high, based on the view that ‘‘a part is the whole’’ (pars-prototo). This view creates a sense of security for clan members because they know that they will always be helped in difficult times. However such a system means that community members have a strong obligation to continually maintain good relationships and share resources with neighbors. This obligation to share means that individuals or communities cannot accumulate or save capital for future investment. Thus, a strong horizontal orientation does not facilitate community development.

work ethic A community’s work ethic encapsulates the social norms and general attitudes towards work. In common parlance, work ethic can be defined as the level of

................. 16157$

$CH3

02-08-07 10:22:10

PS

PAGE 117

118 / j. r. mansoben

enthusiasm and industriousness typical of a person or group. As noted above, Papuan cultures tend to value work highly, and work is thought of as something that produces a product that can be enjoyed by oneself and others. In this view, unproductive people have low social status. However, the definition of productivity varies substantially among groups, based on methods required to make a living in different environments. The work ethic of communities that live collectively in swampy areas and depend on sago for subsistence (e.g., Asmat, Kamoro, Waropen, Bauzi, and Inawatan) is different from the work ethic of other Papuan people who depend on agriculture. In collective societies the work ethic focuses on efforts to fulfill immediate needs (e.g., collecting enough food for a single day) and does not stress work as an investment in the future. People in these groups do not value hard work beyond what is necessary to enjoy life in the present. This view is quite compatible with their social organization and lifestyle, because each family is an element in a production group in which each family does the same work. Collective societies gather products that are immediately available in nature, and do not need to focus on producing or maintaining the resources they utilize. This work ethic is successful and well-adapted to the ecological conditions under which these societies live, but makes collectively-living communities poorly suited to entering the market economy. Such people require support and training in modern technology to allow them to enter the market economy, and empowerment to manage and harvest their resources in a way that will allow the products to be sold at central markets. In contrast to collective societies, the people who practice agriculture have a work ethic that is more geared towards investment for anticipated future returns. Opening land for agriculture is a long process that includes tree-cutting, planting, tending gardens, weeding, and harvesting. This process can take from six to ten months, sometimes more, depending on the types of the plants being grown. This protracted process requires perseverance and diligence, and indicates that Papuan agricultural societies have a particularly strong work ethic. It should be noted that this strong work ethic not only results in sustenance for the household, it also provides products for the market economy. The emphasis on investment for the future is an asset to agricultural societies as they enter the market economy. Many Papuan cultures also are highly competitive. Most Papuan ethnic groups (e.g., Meybrat (Ayamaru), Me, Muyu, Biak, Dani, and Waropen and Serui in Cenderawasih Bay) are inherently competitive. Individuals compete to become powerful and successful members of their groups by accumulating wealth or demonstrating diplomatic expertise, proficiency in warfare, organization abilities, or magical abilities. Therefore Papuans are well-endowed with a competitive spirit, which is valued in the modern world and should prove to be an asset as they enter the global economy. The challenge now is to determine how to help Papuans enhance these cultural assets and apply them for the development of Papua and the rest of Indonesia.

................. 16157$

$CH3

02-08-07 10:22:11

PS

PAGE 118

Socio-cultural Plurality of Papuan Society / 119

Culturally- and Ecologically-sensitive Development One important issue is ascertaining how to help develop and empower Papuan people in a climate that is so ethnically and culturally diverse. Clearly, the correct approaches will depend heavily on the particular situation and society, on the ethnic, religious, and economic status of communities, and the type of development program to be conducted. It should be stressed that any development program must be implemented in a way that is sensitive to Papua’s cultural diversity. For instance, in order to increase the income of people who live in areas with rich ecological potential for sago (e.g., Asmat, Waropen, Inanwatan, and Babo), it would be useful to provide these communities with the skills and technology necessary to increase the quality and quantity of product that they are able to extract. In addition, the government should provide the infrastructure necessary to support the marketing of these products at local or regional markets. Similarly, people who live in the coastal zones with potential for fishing (e.g., Raja Ampat Islands, coastal areas, and islands in the Cenderawasih Bay), need to be given practical knowledge about fishing and preservation techniques that will allow them to transport fish to market. Furthermore, people living in the highland/foothills ecological zone need to be provided with technology to enhance the efficiency of their agricultural practices. It is also important to provide information about modern techniques of cattle farming in order to improve and increase production, especially for people who live in the Central Mountains (e.g., Dani and Me) who have typically been pig farmers. For Dani and Me people, pig farms are usually maintained by the women, so women would need to be centrally involved in these activities.

Literature Cited BPS (Badan Pusat Statistik; Central Bureau of Statistics). 2002. Papua Dalam Angka 2002. BPS Papua, Jayapura. de Bruijn, J.V. 1959. Anthropological research in Netherlands New Guinea since 1950 by Bureau for Native Affair, Hollandia, Netherlands New Guinea. University of Sydney, Sydney. de Bruijn, J.V. 1970. Ekagi land tenure. In Ploeg, A. (ed.) Land Tenure in West Irian. New Guinea Research Bulletin 38. Australian National University, Canberra. Ethnologue (website). 2005. Papuan language diversity. www.ethnologue.com/show_ country.asp?nameIDP. Galis, K.W. 1970. Land tenure in the Biak-Numfor area. In Ploeg, A. (ed.) Land Tenure in West Irian. New Guinea Research Bulletin 38. Australian National University, Canberra. Godschalk, J.A. 1993. Sela Valley: ethnography of Mek society in the Eastern Highlands, Irian Jaya, Indonesia. Ph.D. diss., Amsterdam. Heider, K. 1979. Grand Valley Dani, Peaceful Warriors. Holt, Rinehart and Winston, New York. Held, G.J. 1947. Papoea’s van Waropen. Brill, Leiden

................. 16157$

$CH3

02-08-07 10:22:11

PS

PAGE 119

120 / j. r. mansoben Kamma, F.C. 1953. Zending. In Klein, W.C. (ed.) Koreri: Messianic Movements in the Biak-Numfor Culture Area. Nijhoff, The Hague. Klein, W.C. (ed.). 1953–1954. Nieuw Guinea: De Ontwikkeling op Economisch, Sociaal en Cultuur Gebied, in Nederlands en Australisch Nieuw Guinea. [3 Vol.]. Staatsdrukkerijen Uitgeverijbedrijf, ’s Gravenhage. Koentjaraningrat, R.M. 1970. Keseragaman dan Anekawarna Masyarakat Irian Barat. Lembaga Pengatahuan Indonesia, Jakarta. Koentjaraningrat, R.M. 1974. Kebudayaan, Mentalitas dan Pembangunan. PT Gramedia Pustaka Utama, Jakarta. Koentjaraningrat, R.M. 1984. Kepemimpinan dan kekuasaan: tradisional, masa kini, resmi dan tak resmi. In Budiarjo, M. (ed.) Aneka pikiran tentang kuasa dan wibawa. Sinar Harapan, Jakarta. Leeden, A.C. van der. 1955. Verwantschapstermen en-verhoudingen in het Sarmische. Adatrechtbundel 45: 436–454. Leeden, A.C. van der. 1980.Report on anthropological field research. In Masinambow, E.K.M. (ed.) Halmahera dan Raja Ampat: konsep dan strategi penelitian. LEKNAS-LIPI, Jakarta. Mansoben, J.R. 1957. Kultuur en Kultuurveranderingen in het Moejoegebied. Voorhoeve, The Hague. Mansoben, J.R. 1970. Muyu land tenure. In Ploeg, A. (ed.) Land Tenure in West Irian. New Guinea Research Bulletin 38. Australian National University, Canberra. Mansoben, J.R. 1985. Sistem politik pria berwibawa di Irian Jaya: suatu studi komparatif terhadap lima suku-bangsa. Universitas Indonesia (Thesis S-2), Jakarta. Mansoben, J.R. 1994. Sistem politik tradisional di Irian Jaya. Ph.D. diss., Leiden. Mansoben, J.R., and M.T. Walker. 1990. Irian Jaya cultures: an overview. IBJD 18: 1–16. Masinambow, E.K.M. 1980. Halmahera dan Raja Ampat: konsep dan strategi penelitian. LEKNAS-LIPI, Jakarta. Ploeg, A.1970. Land Tenure in West Irian. New Guinea Research Bulletin 38. Australian National University, Canberra. Pouwer, J. 1966. Toward a configurational approach to society and culture in New Guinea. Journal of Polynesian Society 75: 267–286. Sahlins, M.D. 1963. Poor man, rich man, big-man, chief: political types in Melanesia and Polynesia. Comparative Studies in Society and History 5: 285–303. Sensus Penduduk. 1995. Sensus penduduk Irian Jaya 1995. Seri I, BPS, Jakarta. Silzer, P.J., H. Heikkinen, and D. Clouse. 1986. Peta lokasi bahasa-bahasa daerah di Propinsi Irian Jaya [Publikasi Khusus Bahasa-Bahasa Daerah, Serie D. No.1]. SILUNCEN, Jayapura. Studies in Cultural Anthropology. 1980. Kinship based social categories of Grand Valley Dani. In Cook, E.A., and D. O’Brien (eds.) Blood Semen. University of Michigan Press, Ann Arbor. Verschueren, J. 1970. Marind-Anim land tenure. In Ploeg, A. (ed.) Land Tenure in West Irian. New Guinea Research Bulletin 38. Australian National University, Canberra. Weber, M. 1970/1924. The Theory of Social and Economic Organization. Free Press, Glencoe, Illinois.

................. 16157$

$CH3

02-08-07 10:22:12

PS

PAGE 120

1.4. Prehistoric Human Presence in Papua and Adjacent Areas juliette pasveer Prehistoric Human Occupation in Areas Surrounding Papua h r ou g h m uc h of the Late Pleistocene, because of lowered sea levels, New Guinea was united with Australia, comprising the large landmass known as Sahul (White and O’Connell 1982; Figure 1.4.1). Based on radiocarbon dating, many Australian archeological sites have produced ages in the range 35–40,000 bp (years before present). Re-dating of some sites using a variety of alternative methods suggests, however, that some of these sites may be considerably older. Potentially the earliest sites, based on combinations of optically stimulated luminescence (OSL), thermo-luminescence (TL), electron spin resonance (ESR) and Uraniumseries dates, are the Mungo 3 skeleton in southeastern Australia, with an estimated age of 62,000 bp (Thorne et al. 1999), and Nauwalabila and Malakunanja rockshelters in northern Australia, with estimated ages for first occupation of between 53,000 and 60,000 bp (Roberts et al. 1993, 1994). Although these early dates for human occupation of Australia are by no means universally accepted, the evidence for occupation of Australia at least by 48,000 bp appears increasingly secure (Turney et al. 2001). The New Guinean north coast was settled at least by 40,000 bp, as established by the oldest site currently known on the mainland of New Guinea on the north coastal Huon Peninsula (Groube et al. 1986; Figure 1.4.1). Only one other Pleistocene site is known from a near coastal context; this is Lachitu Rockshelter (Gorecki et al. 1991; Gorecki 1993) in a tectonically active location on the north coast, with a basal age of 35,000 bp. It is likely that many Pleistocene sites on the southern side of Papua New Guinea were drowned by the rising sea level (Groube 1989: 295). At present, the oldest site in the main highlands region, dating around 30,000 bp, is Kuk Swamp, located near Mount Hagen. This site has yielded sparse evidence for human activity at this early date, in the form of human-transported fire-cracked rocks and some charcoal (Golson 1977a; White and O’Connell 1982). Overlying deposits at Kuk contain far more substantial evidence for human activity in the form of early drainage systems, the oldest phase dating to ca 9,000 bp (Golson 1977a; Denham et al. 2003). Other sites of Pleistocene age in New Guinea’s interior include an open site at Kosipe, situated high in the Owen Stanley Range, with stone tools including the distinctive ‘‘waisted’’ blades dating back to 26,000 bp (White et al. 1970); and Nombe Rockshelter, with a basal layer containing the bones of now-extinct animals in association with stone artifacts, probably dating to around 20–25,000 bp (Mountain 1983). Other sites in the New Guinea

T

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

121

................. 16157$

$CH4

02-08-07 10:22:09

PS

PAGE 121

122 / juliette pasveer

Figure 1.4.1. Location of some of the archeological and palynological sites mentioned in the text, with the earliest associated radiometric dates (uncalibrated 14 C or OSL) in years BP. The series of unnamed white dots in the Papua New Guinea Highlands all date to younger than 30,000 BP. The gray area that unites New Guinea and Australia represents the Pleistocene landmass of Sahul. Source: Pasveer (2004).

Highlands date to less than 18,000 bp (White 1972; Bulmer 1977, 1991; Watson and Cole 1977; White and O’Connell 1982). Island Melanesia was inhabited by 33,000 bp; the oldest site of this age is Matenkupkum located on New Ireland (Gosden and Robertson 1991). The site of Kilu on Buka, the northernmost island of the Solomon group, was occupied by 28,000 bp, proving that people were able to cross open sea distances of 130–180 km (Wickler and Spriggs 1988: 705). By at least 12,000 bp, this distance of open water crossing had increased to 200 km, as is shown by occupation of Pamwak Rockshelter on Manus Island by this time (Fredericksen et al. 1993). The prehistory of Island Melanesia also gives indications of the early domestication of plants (Yen 1995), translocation of animals (Flannery and White 1991), and the movement of obsidian over some 350 km, beginning in the Late Pleistocene and continuing into the Holocene (Gosden 1993). Interestingly, this region provides evidence of island exploitation at an earlier time than anywhere else in the world (Keegan and Diamond 1987). The area immediately to the west of New Guinea was until recently an archeological unknown. However, the area has long been of central interest as an arena of early and later human movements (Spriggs 2000). In recent years, archeological research has been conducted along two possible migration routes into New Guinea

................. 16157$

$CH4

02-08-07 10:22:35

PS

PAGE 122

Prehistoric Human Presence in Papua and Adjacent Areas / 123

(Birdsell 1977), one passing through northern Maluku (Bellwood et al. 1998), and another passing through the Aru Islands (Veth et al. 1998; O’Connor et al. 2002). In northern Maluku there is evidence for human occupation from 35,000 bp in Golo Cave, with indications of animal translocation from New Guinea around 8,000 bp (Bellwood et al. 1998: 261). Lemdubu Cave on the Aru Islands recently provided evidence for occupation dating back to around 27,000 bp (O’Connor et al. 2002, 2005).

Prehistoric Human Presence in Papua The first archeological excavation in western New Guinea was carried out as a ‘‘side activity’’ by members of the 1971–1973 Australian-Indonesian Carstenz Glaciers Expeditions (Hope et al. 1976). Hope and Hope (1976) reported that Mapala Rockshelter on Mt Jaya (ca 4,000 m; Figure 1.4.1) contained bones, ash, a few stone artifacts, and shell remains, dated to ca 5,440 bp, but most of the deposit was left in situ. Other indications of early human activity in the region came from research at Ijomba Bog at 3,600 m on Mt Jaya, where a pollen record suggests that firing may have led to an opening up of forest cover by 10,500 bp (Hope and Hope 1976). Similar evidence for deliberate or accidental firing of vegetation in the Baliem Valley dating to 32,000 bp is reported by Haberle et al. (1991). In 1995, excavations took place in two cave sites in the interior of the Vogelkop (Bird’s Head), the westernmost peninsula of Papua. These two sites form currently the only direct and detailed archeological evidence for Pleistocene to early Holocene occupation of Papua (Pasveer 2004), and provide a wealth of information on the circumstances under which the hunters and gatherers survived. The two sites are located on the Ayamaru Plateau, a limestone karst area located centrally on the Vogelkop (Figures 1.4.2 and 1.4.3). The plateau is situated ca 350 m above sea level and contains the three shallow connected Ayamaru Lakes. At least one of the lakes dates to the mid-Holocene, but it is possible that the other two have existed longer. Nowadays the population is concentrated around these lakes, although there are numerous villages and active and abandoned gardens in the wider region among the low, rainforested hills. Both sites are located at cave entrances with passages into a larger cave system: Kria Cave is located ca 3 km northeast of the easternmost lake, while Toe´ Cave is situated on a small peninsula jutting into the southwestern lake from its southern shore. Their distance from each other is ca 12 km. Kria Cave contains more than 2 meters of sediment and has a virtually undisturbed stratigraphy. Five occupation ‘‘units’’ were identified: the four bottom units, differing from each other only in quantities of material, contain mostly stone and bone artifacts, large quantities of faunal remains (the bulk of which come from a forest wallaby, Dorcopsis muelleri), and smaller amounts of molluscan shell, eggshell, ochre, charcoal, and a few botanic remains. The oldest radiocarbon date from the deepest unit (Unit V) is 6,900 bp (uncalibrated). However, not all of the sediment was excavated, and the date of first occupation of the cave can be

................. 16157$

$CH4

02-08-07 10:22:35

PS

PAGE 123

124 / juliette pasveer

Figure 1.4.2. Karst formation on the Ayamaru Plateau. Source: Courtesy J. Jelsma.

Figure 1.4.3. The Vogelkop Peninsula with location of the Ayamaru Plateau relative to areas of higher elevation. Source: Pasveer (2004).

................. 16157$

$CH4

02-08-07 10:23:12

PS

PAGE 124

Prehistoric Human Presence in Papua and Adjacent Areas / 125

extrapolated to ca 8,000 bp. The activities in the cave, which included consumption of the hunted wildlife, and manufacturing, use, and repair of stone and bone artifacts, remained virtually the same over 3,000 years. The youngest date in Unit II provides an age around 4,000 bp. The top unit in Kria Cave differs dramatically from the four underlying ones. This last occupation phase is characterized by the presence of only a few stone artifacts, no bone artifacts, a few pottery sherds, and a few bones of a single pig, which were directly dated to a minimum age of 1,840 bp. The quantities of faunal remains are considerably smaller and differ also in composition, with a conspicuous drop in wallaby remains. This unit contains a small shell midden and a human burial. It appears that the site was abandoned for some time after ca 4,000 bp, but occupation of the cave resumed by 1,840 bp at the latest, and was perhaps used for different purposes, including human interment. There were no clear signs or indications from the local Ayamaru people that the cave was still in use today, and it is uncertain when occupation of the site ceased. Toe´ Cave contains a maximum of 140 cm deposit and due to its sloping bedrock, its stratigraphy is very complex. However, detailed analyses of its contents and radiocarbon dates have led to the conclusion that the deposit is essentially intact, albeit with minor local slumping and infill. Two main ‘‘units’’ have been identified: the bottom one-third of the deposit is a Pleistocene unit with stone material that is more weathered, and faunal remains that are more fossilized than that from the upper unit; the top two-thirds is of Holocene age and is characterized by large quantities of stone artifacts and faunal remains (again dominated by the remains of Dorcopsis wallabies), and contain otherwise essentially the same kinds of material as were found in Kria Cave. Toe´ Cave differs from Kria Cave in the age of the occupation, the lack of bone artifacts, and the much larger (in size) and larger quantities of stone artifacts. However, there are clearly more subtle differences between the sites, which will be discussed below. Radiocarbon dating of Toe´ Cave was based predominantly on ratite eggshell fragments, which showed that occupation of this cave commenced around 26,000 uncalibrated years ago. With only 140 cm of sediment in this cave, this makes the sedimentation rate of the deposit remarkably low, suggesting that the site was not very frequently visited. Nevertheless, occupation continued until some time after ca 3,000 bp and possibly even until sub-Recent times. The Pleistocene unit in Toe´ Cave revealed some interesting climatic changes after the Last Glacial Maximum. Until now it has generally been assumed that this last glacial period did not have a noticeable influence on lowland tropical areas. However, a large proportion of the faunal remains from this part of Toe´ Cave’s deposit could be identified as animal species that today normally live above ca 1,000 m elevation (Figure 1.4.4); such animals would only be able to descend to lower altitudes if temperatures were lower. Mountainous areas above 1,000 meters are at least 50 km away from the site (Figure 1.4.3), well beyond a hunter’s range, even for New Guinean standards. This means that the Last Glacial Maximum must have pushed the vegetation zones further down the lowland mountain slopes than

................. 16157$

$CH4

02-08-07 10:23:13

PS

PAGE 125

126 / juliette pasveer

Figure 1.4.4. Toe´ Cave, proportional distribution (based on number of fragments) of vertebrate groups by habitat through the deposit. ‘‘pred.’’  predominantly. Source: after Pasveer (2004).

has always been assumed, and even at 350 m above sea level temperatures must have been some 5–6 degrees lower than today. When global climate reached its present state around 12–10,000 years ago, flora and fauna gradually receded to their current Holocene positions. Montane species left the area or became locally extinct, and the faunal composition became one typical of New Guinean lowland habitats. Interestingly, at this stage the fauna on the Ayamaru Plateau became dominated by the Brown Dorcopsis (Dorcopsis muelleri). This also became the focus of local hunters, presumably because this ground-dwelling species was a relatively easy catch, was abundant, and was one of the largest prey species in the area. The dominance of the Brown Dorcopsis during the Holocene is visible in the

................. 16157$

$CH4

02-08-07 10:24:06

PS

PAGE 126

Prehistoric Human Presence in Papua and Adjacent Areas / 127

assemblages from both caves. Some 75–80% of all animal bone is identified as this species. But does this mean that the Ayamaru hunters specifically targeted this animal? It is a well-known fact that tropical rainforests are very poor environments for humans and are not very productive in terms of food (see Rainforest Occupation before Agriculture, below), and most of the food resources are located high above ground. This makes it highly unlikely that New Guinean rainforest occupants could afford to be selective in their hunting efforts to the expense of other animals. This is supported by ethnographic information on New Guinean hunters.

RAINFOREST OCCUPATION BEFORE AGRICULTURE: THE HEADL AND AND BAILEY DEBATE A major point of debate in this region—and in fact for all tropical regions in the world—is whether tropical rainforest has ever been inhabited prior to the introduction of agriculture. This debate was started in the late 1980s by two American researchers, Robert Bailey and Thomas Headland, who independently came up with very similar hypotheses. These hypotheses are based on observations of contemporary hunter-gatherers who live in rainforests around the world. None of these groups of people live completely independently of their farming neighbors. The foragers exchange various products from the forest for agricultural products. Although rainforest habitats harbor a much higher diversity of species than other terrestrial ecosystems, most food resources are relatively inaccessible for ground-dwellers such as humans. Most animals live high up in the canopy of the trees, and are highly camouflaged and hard to find. Those animals that live on the forest floor are either very small and sedentary, or very large, solitary, highly mobile, and dispersed. Because the forest floor has little to offer in terms of food, this habitat can support only a relatively small number of large terrestrial mammals. Bailey et al. (1989) formulated the hypothesis that rainforests in general do not provide sufficient food resources for humans to be able to survive for long periods of time. Headland (1987) said essentially the same, but pointed out that it is the lack of carbohydrates in rainforests that really causes the problem, because edible plants are present in only very limited quantity due to the dense forest cover that limits the penetration of sunlight. Headland and Bailey (1991) therefore concluded that it would not have been possible for humans to live in rainforests until food resources were made more reliable through cultivation and domestication. They saw their theories supported by the apparent lack of archeological evidence for rainforest occupation before ca 9,000 bp. However, ample evidence has since been presented in response to their suggestions, for example, of sites on the Malay Peninsula (Endicott and Bellwood 1991) and in West New Britain (Pavlides and Gosden 1994). The evidence from the Vogelkop (Bird’s Head) of Papua can now be added to the available evidence of prehistoric rainforest occupation well before the advent of agriculture.

................. 16157$

$CH4

02-08-07 10:24:07

PS

PAGE 127

128 / juliette pasveer

So if the hunters were not specifically targeting the Dorcopsis wallaby, it must have been abundant in the area so that it was the one species most often encountered. However, rainforest environments cannot hold large numbers of relatively large ground-dwelling animals, for the same reason that humans have difficulty finding food. However, while solitary, mobile, and highly dispersed, forest wallabies were probably still the most abundant species in the area, and therefore the most common prey animal. The relative abundance of the Dorcopsis wallaby in the area during most of the Holocene is confirmed by the analysis of the population structure of the wallabies that were brought into the caves. Based on tooth eruption, tooth wear, and molar progression (the forward movement of the teeth), which are all related to age, it was established that the majority of animals were adult individuals; there were no pouch young and subadults were rare. According to general ecological principles, such an age structure is typical for a food-limited population that is living in high densities, relative to the carrying capacity of the environment that they inhabit (see The Effect of Random Culling, below), and this situation remained unchanged for thousands of years. That the wallaby population was living at its carrying capacity implies that it was not very often hunted. Although this appears to contradict its dominance in the excavated faunal assemblages, this can easily be explained if the bones in the caves accumulated at a very slow pace. The low hunting return is confirmed by the estimated minimum number of individuals (MNI; i.e., wallabies) brought to the sites, as represented in the faunal assemblages. This MNI can be calculated from the various body parts of Dorcopsis present in the caves, and this can be done for certain periods through time. It was calculated that in Kria Cave only one wallaby per 3 or 4 years was brought to the cave. In Toe´ Cave this was much less, perhaps one wallaby per 40 years. Although, these are very rough figures, it clearly shows that the sites were indeed rarely visited. As explained above, the age structure of the wallabies that were caught suggests that this infrequent visiting was not only true for these particular caves, but for most of the Ayamaru Plateau. The question, of course, is why did people visit the plateau so infrequently, if there was enough good-sized game available? If the Dorcopsis population remained dense and stable for thousands of years, then people were clearly not hunting them very much. Meat sources were not the problem, but perhaps, as Headland suggested, it was not so much the availability of protein, but more the lack of carbohydrates that limited human presence on the plateau. Rainforest is generally not very rich in edible plants, because of the limited amount of sunlight that can penetrate to the understory. It is very possible that humans did not stay long in the caves because they could not find sufficient plant foods to survive. Although this is somewhat speculative, it may be a plausible answer to this question. However, there was likely a combination of factors, including warfare and diseases, that caused a high mobility (and perhaps a high mortality) among the groups occupying the plateau. The living conditions described above for Pleistocene and early to mid-Holocene

................. 16157$

$CH4

02-08-07 10:24:07

PS

PAGE 128

Prehistoric Human Presence in Papua and Adjacent Areas / 129

THE EFFECT OF RANDOM CULLING ON POPUL ATION AGE STRUCTURE The ‘‘sustainable yield’’ of a population (i.e., the maximum proportion that can be randomly culled from a population without driving it to extinction) is related to both the population density (relative to the carrying capacity of its environment) and the population’s age structure, as has been well established in the fields of population ecology (Krebs 1978) and wildlife management (Caughley 1977; Getz and Haight 1989). A population age structure with a large proportion of very young animals (Figure 1.4.5; Structure A) is typical of low-density popula-

Figure 1.4.5. Relationship between sustainable yield and population density relative to the carrying capacity of a given environment. Source: Pasveer (2004).

tions; to save the population from extinction, random culling has to remain low because few animals have reached the breeding stage. A population that allows for higher levels of random culling is one that approaches a typical ‘‘pyramid’’ age structure (Odum 1971), with relatively large numbers of young animals, fewer adults, and even fewer aged individuals (Figure 1.4.5; Structure B). Hunting will

................. 16157$

$CH4

02-08-07 10:24:36

PS

PAGE 129

130 / juliette pasveer remove a large proportion of the breeding females but the high number of subadult individuals allow for a rapid recovery. An age structure dominated by mature to old individuals (Figure 1.4.5; Structure C) is characteristic of high density populations. The already saturated environment provides limited opportunities for recruitment of young animals and has correspondingly high natural mortality rates among young individuals. High levels of random culling will remove proportionally large numbers of breeding females, thereby having an immediate impact on the reproductive potential of the population as a whole; because, unlike the pyramid age structure, the low number of subadult individuals precludes a rapid recovery. Lower levels of culling, however, will result in a ‘‘thinning’’ of the population, increased recruitment of young individuals, and a shift towards a pyramid age structure with higher numbers of young and subadult individuals (Structure B). Thus, if a population were under hunting pressure, it would become visible in the age structure of the population over time. The rate at which a population can become ‘‘over-hunted’’ depends on a number of factors, including the species’ age at first breeding, litter size, interval between litters, and longevity (Caughley 1977; Alvard et al. 1997; Bodmer et al. 1997). The Brown Dorcopsis, with a relatively slow maturation, a litter size of one, an interval between litters of at least 6 months, and a likely maximum life span of around 10 years, qualifies as a relatively vulnerable species in these terms. Population densities approach carrying capacity only where population growth is limited solely by the productivity of the natural environment. For the Brown Dorcopsis, as one of the few relatively large mammals in the Ayamaru region and with no large predators prior to the arrival of humans, it is theoretically possible to attain carrying capacity.

Papuan people is very likely to be true for other lowland tropical areas in New Guinea, especially in limestone areas, which are famous for their poor soils. The situation was only to improve with the advent of plant cultivation. The faunal assemblage from Kria Cave shows a slight change in the age profile of the wallabies brought into the cave around 5,000 bp, which might indicate that hunting of Dorcopsis increased and/or its habitat was increasingly disturbed. It is likely that this was the result of an increase in human population densities in the area, together with efforts to open up the forest canopy to stimulate the growth of edible plants. This may indicate the onset of agriculture in this westernmost part of Papua.

Literature Cited Allen, J. 1994. Radiocarbon determinations, luminescence dating and Australian archaeology. Antiquity 68: 339–343. Allen, J., and S. Holdaway. 1995. The contamination of Pleistocene radiocarbon determinations in Australia. Antiquity 69: 101–112.

................. 16157$

$CH4

02-08-07 10:24:37

PS

PAGE 130

Prehistoric Human Presence in Papua and Adjacent Areas / 131 Alvard, M.S, J.G. Robinson, K.H. Redford, and H. Kaplan. 1997. The sustainability of subsistence hunting in the neotropics. Conservation Biology 11 (4): 977–982. Bailey, R.C., G. Head, M. Jenike, B. Owen, R. Rechtman, and E. Zechenter. 1989. Hunting and gathering in tropical rain forest: is it possible? American Anthropologist 91: 59–82. Bellwood, P., G. Nitihaminoto, G. Irwin, Gunadi, A. Waluyo, and D. Tanudirjo. 1998. 35,000 years of prehistory in the northern Moluccas. Pp. 233–275 in Bartstra, G.-J. (ed.) Bird’s Head Approaches. Irian Jaya Studies—A Programme for Interdisciplinary Research. Modern Quaternary Research in Southeast Asia 15. A.A. Balkema, Rotterdam. Birdsell, J.B. 1977. The recalibration of a paradigm for the first peopling of Greater Australia. Pp. 113–167 in Allen, J., J. Golson, and R. Jones (eds.) Sunda and Sahul: Prehistoric Studies in Southeast Asia, Melanesia and Australia. Academic Press, London. Bodmer, R.E., J.F. Eisenberg, and K.H. Redford. 1997. Hunting and the likelihood of extinction of Amazonian mammals. Conservation Biology 11: 460–466. Bulmer, S. 1977. Between the mountain and the plain: prehistoric settlement and environment in the Kaironk Valley. Pp. 61–73 in Winslow, J.H. (ed.) The Melanesian Environment. Australian National University Press, Canberra. Bulmer, S. 1991. Variation and change in stone tools in the Highlands of Papua New Guinea: the witness of Wan˜elek. Pp. 470–478 in Pawley, A. (ed.) Man and a Half: Essays in Pacific Anthropology and Ethnobiology in Honour of Ralph Bulmer. The Polynesian Society, Auckland. Caughley, G. 1977. Analysis of Vertebrate Populations. John Wiley and Sons, London. Denham, T.P., S.G. Haberle, C. Lentfer, R. Fullagar, J. Field, M. Therin, N. Porch, and B. Winsborough. 2003. Origins of agriculture at Kuk Swamp in the Highlands of New Guinea. Science 301: 189–193. Endicott, K., and P. Bellwood. 1991. The possibility of independent foraging in the rain forest of peninsular Malaysia. Human Ecology 19 (2): 151–185. Flannery, T.F., and J.P. White. 1991. Animal translocations. the zoogeography of New Ireland mammals. National Geographic Research and Exploration 7: 96–113. Fredericksen, C., M. Spriggs, and W. Ambrose. 1993. Pamwak Rockshelter: a Pleistocene site on Manus Island, Papua New Guinea. Pp. 144–152 in Smith, M.A., M. Spriggs, and B. Fankhauser (eds.) Sahul in Review. Pleistocene Archaeology in Australia, New Guinea and Island Melanesia. Occasional Papers in Prehistory 24. Department of Prehistory, Research School of Pacific Studies, Australian National University, Canberra. Getz, W.M., and R.G. Haight. 1989. Population Harvesting: Demographic Models of Fish, Forest, and Animal Resources. Princeton University Press, Princeton, New Jersey. Golson, J. 1977a. No room at the top: agricultural intensification in the New Guinea highlands. Pp. 601–638 in Allen, J., J. Golson, and R. Jones (eds.) Sunda and Sahul: Prehistoric Studies in Southeast Asia, Melanesia and Australia. Academic Press, London. Gorecki, P. 1993. Saltwater people of the southwest Kimberley coast. Pp. 154–159 in Burenhult, G. (ed.) The Illustrated History of Humankind. Vol. 2. Hunter-gatherers and Early Farmers. American Museum of National History, Harper, San Francisco, California. Gorecki, P., M. Mabin, and J. Campbell. 1991. Archaeology and geomorphology of the Vanimo Coast, Papua New Guinea: preliminary results. Archaeology in Oceania 26: 119–122. Gosden, C. 1993. Understanding the settlement of Pacific Islands in the Pleistocene. Pp. 131–136 in Smith, M.A., M. Spriggs, and B. Fankhauser (eds.) Sahul in Review: Pleistocene Archaeology in Australia, New Guinea and Island Melanesia. Occasional

................. 16157$

$CH4

02-08-07 10:24:37

PS

PAGE 131

132 / juliette pasveer Papers in Prehistory 24. Department of Prehistory, Research School of Pacific Studies, Australian National University, Canberra. Gosden, C., and N. Robertson. 1991. Models for Matenkupkum: interpreting a late Pleistocene site from southern New Ireland, Papua New Guinea. Pp. 20–45 in Allen, J., and C. Gosden (eds.) Report of the Lapita Homeland Project. Occasional Papers in Prehistory 20. Department of Prehistory, Research School of Pacific Studies, Australian National University, Canberra. Groube, L. 1989. The taming of the rain forests: a model for Late Pleistocene forest exploitation in New Guinea. Pp. 292–304 in Harris, D.R., and G.C. Hillman (eds.) Foraging and Farming. The Evolution of Plant Exploitation. Unwin Hyman, London. Groube, L., J. Chappell, J. Muke, and D.A. Price. 1986. 40,000 year old human occupation site at Huon Peninsula, Papua New Guinea. Nature 324: 453–455. Haberle, S.G., G.S. Hope, and Y. de Fretes. 1991. Environmental change in the Baliem Valley, montane Irian Jaya, Republic of Indonesia. Journal of Biogeography 18: 25–40. Haberle, S.G., G.S. Hope, and S. van der Kaars. 2001. Biomass burning in Indonesia and Papua New Guinea: natural and human induced fire events in the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 171: 259–268. Headland, T.N. 1987. The wild yam question: how well could independent huntergatherers live in a tropical rain forest ecosystem? Human Ecology 15: 463–491. Headland, T.N., and R.C. Bailey. 1991. Introduction: have hunter-gatherers ever lived in tropical rain forest independently of agriculture? Human Ecology 19 (2): 115–122. Hope, G.S. 1996. Quaternary change and the historical biogeography of Pacific islands. Pp. 165–190 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas. New Guinea to Eastern Polynesia: Patterns and Processes: 165–190. SPB Academic Publishing, Amsterdam. Hope, G.S., and J.H. Hope. 1976. Man on Mt. Jaya. Pp. 225–239 in Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.) The Equatorial Glaciers of New Guinea. Results of the 1971–1973 Australian Universities’ Expeditions to Irian Jaya: Survey, Glaciology, Meteorology, Biology and Palaeoenvironments. A.A. Balkema, Rotterdam. Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.). 1976. The Equatorial Glaciers of New Guinea: Results of the 1971–1973 Australian Universities’ Expeditions to Irian Jaya: Survey, Glaciology, Meteorology, Biology and Palaeoenvironments. A.A. Balkema, Rotterdam. Keegan, W.F., and J.M. Diamond. 1987. Colonization of islands by humans: a biogeographical perspective. Pp. 49–93 in Schiffer, M.B. (ed.) Advances in Archaeological Method and Theory 10. Academic Press, San Diego. Krebs, C.J. 1978. Ecology. The Experimental Analysis of Distribution and Abundance. 2nd ed. Harper and Row, New York. Mountain, M.-J. 1983. Preliminary report of excavations at Nombe rockshelter, Simbu Province, Papua New Guinea. Bulletin of the Indo-Pacific Prehistory Association 4: 84–99. O’Connell, J.F., and J. Allen. 1998. When did humans first arrive in Greater Australia and why is it important to know? Evolutionary Anthropology 6: 132–146. O’Connor, S., K.P. Aplin, M. Spriggs, P. Veth, and L.K. Ayliffe. 2002. From savanna to rainforest: changing environments and human occupation at Liang Lemdubu, Aru Islands, Maluku (Indonesia). Pp. 279–306 in Kershaw, P., B. David, N. Tapper, D. Penny, and J. Brown (eds.) Bridging Wallace’s Line: The Environmental and Cultural History and Dynamics of the SE Asian-Australian Region. Advances in Geoecology 34. Catena Verlag, Reiskirchen.

................. 16157$

$CH4

02-08-07 10:24:38

PS

PAGE 132

Prehistoric Human Presence in Papua and Adjacent Areas / 133 O’Connor, S., K. Aplin, K. Szabo´, J. Pasveer, P. Veth, and M. Spriggs. 2005. Liang Lemdubu: a Pleistocene cave site in the Aru Islands. Pp. 171–204 in O’Connor, S., M. Spriggs, and P. Veth (eds.) The Archaeology of the Aru Islands, Eastern Indonesia. Terra Australis 22. Pandanus Books, Canberra. Odum, E.P. 1971. Fundamentals of Ecology. 3rd ed. W.B. Saunders, Philadelphia. Pasveer, J.M. 2004. The Djief hunters: 26,000 Years of Rainforest Exploitation on the Bird’s Head of Papua, Indonesia. Modern Quaternary Research in Southeast Asia 17. A.A. Balkema, Leiden. Pavlides, C., and C. Gosden. 1994. 35,000-year-old sites in the rainforests of West New Britain, Papua New Guinea. Antiquity 68: 604–610. Roberts, R.G., R. Jones, and M.A. Smith. 1993. Optical dating at Deaf Adder Gorge, Northern Territory, indicates human occupation between 53,000 and 60,000 years ago. Australian Archaeology 37: 58–59. Roberts, R.G., R. Jones, N.A. Spooner, M.J. Head, A.S. Murray, and M.A. Smith. 1994. The human colonisation of Australia: optical dates of 53,000 and 60,000 years bracket human arrival at Deaf Adder Gorge, Northern Territory. Quaternary Geochronology (Quaternary Science Review) 13: 575–583. Spriggs, M. 2000. Out of Asia: the spread of Southeast Asian Pleistocene and Neolithic maritime cultures in island Southeast Asia and the western Pacific. Pp. 51–75 in O’Connor, S., and P. Veth (eds.) East of Wallace’s Line: Studies of Past and Present Maritime Cultures of the Indo-Pacific Region. Modern Quaternary Research in. Southeast Asia 16. A.A. Balkema, Rotterdam. Thorne, A., R. Gru¨n, G. Mortimer, N.A. Spooner, J.J. Simpson, M. McCulloch, L. Taylor, and D. Curnoe. 1999. Australia’s oldest human remains: age of the Lake Mungo 3 skeleton. Journal of Human Evolution 36: 591–612. Turney, C.S.M., M.I. Bird, L.K. Fifield, R.G. Roberts, M.A. Smith, C.E. Dortch, R. Gru¨n, E. Lawson, L.K. Ayliffe, G.H. Miller, J. Dortch, and R.G. Cresswell. 2001. Early human occupation at Devil’s Lair, southwestern Australia 50,000 years ago. Quaternary Research 55: 3–13. Veth, P., M. Spriggs, A. Jatmiko, and S. O’Connor. 1998. Bridging Sunda and Sahul: the archaeological significance of the Aru Islands, southern Moluccas. Pp. 157–177 in Bartstra, G.-J. (ed.) Bird’s Head Approaches. Irian Jaya Studies—A programme for Interdisciplinary Research. Modern Quaternary Research in Southeast Asia 15. A.A. Balkema, Rotterdam. Watson, V.D., and J.D. Cole. 1977. Prehistory of the Eastern Highlands of New Guinea. University of Washington Press, Seattle. White, J.P. 1972. Ol Tumbuna: Archaeological Excavations in the Eastern Central Highlands, Papua New Guinea. Terra Australis 2. Department of Prehistory, Research School of Pacific Studies, Australian National University, Canberra. White, J.P., K.A.W. Crook, and B.P. Buxton. 1970. Kosipe: a Late Pleistocene site in the Papuan Highlands. Proceedings of the Prehistoric Society 36: 152–170. White, J.P., and J.F. O’Connell 1982. A Prehistory of Australia, New Guinea and Sahul. Academic Press, Sydney. Wickler, S., and M. Spriggs. 1988. Pleistocene human occupation of the Solomon Islands, Melanesia. Antiquity 62: 703–706. Yen, D.E. 1995. The development of Sahul agriculture with Australia as bystander. Pp. 831–847 in Allen, J., and J.F. O’Connell (eds.) Transitions: Pleistocene to Holocene in Australia and Papua New Guinea. Antiquity 69, Special number 265.

................. 16157$

$CH4

02-08-07 10:24:39

PS

PAGE 133

................. 16157$

$CH4

02-08-07 10:24:39

PS

PAGE 134

section two 

The Physical Environment

................. 16157$

PRT2

04-13-07 13:50:04

PS

PAGE 135

................. 16157$

PRT2

02-08-07 10:21:51

PS

PAGE 136

2.1. Tectonic Geology of Papua dan a. polhemus h e is l a n d o f n e w g u i n ea can be thought of in simplest geological terms as the mountainous, tectonically deformed northern margin of Australia. Although we typically view New Guinea as a separate geographical entity, it is in fact separated from northern Australia by only a very shallow epicontinental sea less than 15 m deep, the Torres Strait/Arafura Sea, which did not even exist during most of the Pleistocene ice ages that spanned the last 20,000 years. Thus the existence of New Guinea as a discrete island is recent and transient product of the current warm interglacial period. Viewed from a global tectonic perspective, New Guinea has been formed by the collision of the very large, northward-moving Indo-Australian tectonic plate with the even larger, westward-moving Pacific Plate (Figure 2.1.1). This collision has been mediated to various extents by a series of smaller, intervening tectonic plates that lie in the region between the Philippines and the Solomon Islands. These smaller plates, specifically the Philippine Sea, Caroline, and Solomon plates, are

T

Figure 2.1.1. Major tectonic plates of the Indo-Australian region. Barbs at plate boundaries are on the overriding plates and indicate the direction of subduction beneath the plates. 137

................. 16157$

$CH5

02-08-07 10:22:30

PS

PAGE 137

138 / d a n a . p o l h e m u s

all bordered by subduction zone trenches, strike-slip faults, or island arcs, with assorted fragments of the island arcs having been swept against the northern margin of the Australian craton from the Cretaceous onward. As a result of these convergent plate motions, the northern half of New Guinea now consists of a composite of island arc terranes that have accreted to the Australian continental craton at various times over the last 75 million years. The exact sequence of these collisions is still to some degree uncertain, with varying scenarios having been proposed by Hamilton (1979), Kroenke (1984), Hill (2002), Hill and Hall (2003), and others. Most of these hypotheses recognize at least two episodes of terrane accretion: the first at some time between the latest Cretaceous and the Oligocene, involving an island arc that extended primarily to the west toward Sundaland; and the second in the Miocene to Pliocene, involving an arc that extended more to the east and perhaps north (see Figure 2.1.2 for a geological time chart). Because much of the onshore geology of New Guinea is overplated by Late Tertiary orogeny and hidden under dense forest, considerably more remains to be learned about the precise timing of such collisions and the particular sectors of the island attributable to them.

The Geological Dynamics of Island Arcs Because island arcs have been fundamental to the creation of New Guinea, it is useful to understand their formation, migration, and accretion. Island arcs are linear chains of volcanoes formed in mid-ocean settings above subduction zones that mark plate boundaries. In general, arcs display a predictable pattern of concentric features, beginning with a trench along the leading edge of the arc where one plate slides beneath the other. This is followed in turn by a fore-arc ridge, which represents the crest of an accretionary wedge of tectonic debris scraped up by, or stuffed beneath, the overriding plate in the system. Behind the fore-arc ridge is a shallow fore-arc basin, separating the ridge from a parallel volcanic arc that forms above the steeply dipping portion of the subducting plate, that has melted at depth to form rising magma (Figure 2.1.3). Finally, behind the volcanic arc is a back-arc basin, which frequently exhibits pull-apart extension similar to that seen at mid-ocean ridges. These features were reviewed in detail by Hamilton (1988), and are particularly well illustrated in the present day Sunda and Tonga arcs, which border the New Guinea region to the west and east, respectively. Modern examples of fore-arc ridge islands in these systems include the Mentawai Archipelago and Timor in the Sunda Arc and the Tonga Islands in the Tonga Arc; such islands are composed of chaotically jumbled oceanic sediments and frequently contain uplifted Quaternary reef limestones, sometimes underlain by blueschists and other metamorphics. The islands of the volcanic arcs behind these fore-arc ridges, such as Sumatra, Java, and the Lesser Sundas in the Sunda system, or the Lau Group in the Tonga system, have a contrastingly igneous composition, varying over time from tholeiitic basalts in young arcs to calc-alkalic basalts, andesites, and dacites in mature arcs (Hamilton 1988). It is important to distinguish island

................. 16157$

$CH5

02-08-07 10:22:30

PS

PAGE 138

Tectonic Geology of Papua / 139

ERA

PERIOD Quaternary Neogene

Cenozoic Tertiary

Paleogene Cretaceous

Mesozoic

Jurassic

Triassic

Permian

Pennsylvanian Carboniferous Mississippian Paleozoic

Devonian

Silurian

Ordovician

Devonian

EPOCH Holocene Pleistocene Pliocene Miocene Oligocene Eocene Paleocene Late Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early Late Middle Early

AGE ESTIMATES OF BOUNDARIES IN MILLION YEARS (MYA) 0.010 1.7 5.0 24 38 55 66 138

205

240

290

330

360

410

435

500

Figure 2.1.2. Geological time chart.

................. 16157$

$CH5

02-08-07 10:22:31

PS

PAGE 139

140 / d a n a . p o l h e m u s

Figure 2.1.3. Cross-sectional cartoon of an island arc system. The subducting plate is on the left side of the figure, and the overriding plate on the right. As the subducting plate slides under the overriding plate, sediments are scraped off of its upper surface and accumulate to form a fore-arc ridge. As the plate continues to subduct more deeply, it melts at depth to form magma, which rises upward and erupts through the overriding plate to create a volcanic arc. A submarine fore-arc basin lies between the fore-arc ridge and the volcanic arc. For further discussion of these features see the text.

arcs, which form at subduction zones, from progressional volcanic chains such as Hawai’i and Samoa that form above hot spots. Progressional volcanic chains consist of strings of islands with progressively increasing ages as one moves away from the hot spot plume, and do not exhibit the concentric structures or forward migration perpendicular to the longitudinal axis of the island chain that are typical of true island arcs. Island arcs are dynamic features that advance forward with time over the descending slabs of oceanic crust in the subduction zone, apparently due to a slow backward collapse of the crust on the subducted (i.e., descending) plate. This process is most pronounced in the center of an arc; thus migrating island arcs increase their curvature as they advance. At the same time, such advancing arcs produce new oceanic crust behind their advancing island fronts via crustal extension in the back-arc basin, lying behind the volcanic portion of the arc. Forward migration is generally continuous over time, though at varying rates along the length of the arc or in various sectors of it, and proceeds until subduction ceases, or until collision occurs with another arc or with a passive continental margin (the latter having been an important process in the formation of New Guinea). In the process of forward migration an arc may also split longitudinally along its length, with the front section nearest the trench migrating away from the remain-

................. 16157$

$CH5

02-08-07 10:22:57

PS

PAGE 140

Tectonic Geology of Papua / 141

der, as has occurred at least once in the Tonga Arc and on multiple occasions in the Mariana Arc. Such split arcs leave abandoned arc ridges in their wakes, the crests of which may form remnant islands, such as the Lau Islands in the Tonga system, or Yap and Palau in Micronesia (Karig 1971; Ewart 1988). Subducted slabs associated with senescent arcs can also survive for tens of millions of years at the bottom of the upper mantle, where they may be detected by recently refined seismic tomography techniques, thus indicating the positions of now-vanished subduction zones (Hall and Spakman 2003). Arcs typically form crescent-shaped oceanic archipelagos, examples of which are common along the western margin of the Pacific. Conspicuous examples easily picked out by examination of a globe or world map include the Aleutian Islands, Kamchatka and the Kurils, Japan, the Marianas, and Tonga. By contrast, arcs that have undergone collision with other land masses may exist as assemblages of multiply accreted tectonic units in oceanic settings, such as the Philippines, or as one or more layers of terranes laminated onto continental margins, as seen in northern New Guinea. Accreted arc terranes of the latter type frequently include exposures of ophiolites, which are sections of oceanic lithosphere that were uplifted and obducted during the arc collision. Following a collision, a new subduction zone of a polarity (i.e., dip) opposite to that which formerly lay ahead of the advancing arc may form behind the new composite terrane, usually in the extensionally thinned crust of the old back-arc basin. The strip of back-arc crust between the old arc and the new trench can thus become the fore-arc basement of a new arc system, which migrates away from the zone of collision. This type of sequence has been postulated for northern Papua New Guinea by various researchers (Dewey and Bird 1970; Karig 1972; Hamilton 1979), who interpreted the Miocene collision of a southward migrating arc with the Australian continental margin to have been followed by the formation of a new arc, the Schouten Archipelago, that is now beginning to migrate northward away from New Guinea. Subsequent studies have indicated that such a process does appear to be taking place, although it is in a far more incipient stage than previously believed (Cooper and Taylor 1987). Arc collisions with continents or other arcs are typically characterized by the emplacement of a distinct stratigraphic assemblage, which, as noted above, often includes ophiolites. Stratigraphically intact accreted arc complexes often exhibit sequential bands of tectonic melange (derived from the fore-arc ridge), limestone (derived from the floor of the fore-arc basin), basalts and other volcanics (derived from the volcanic arc), and ophiolite (derived from the back-arc basin). It is important to note that the ophiolitic section can be derived from either the fore-arc or the back-arc, and can therefore be either the first or last stratigraphic unit emplaced in an arc terrane collision, and both scenarios have been proposed for the ophiolites emplaced in the central mountains of New Guinea. Intact ophiolites are characterized by a highly distinctive sequence as one proceeds down-section of pillow basalt, massive gabbros, and serpentinized ultramafics. Sequential suturing of melange, limestone, volcanics, and ophiolites is well illustrated in the central highlands and northern coastal mountains of New Guinea, but may also be seen

................. 16157$

$CH5

02-08-07 10:22:58

PS

PAGE 141

142 / d a n a . p o l h e m u s

elsewhere around the Pacific Rim where arcs have fused to each other or accreted to continental margins, including western Luzon (Hamilton 1988), Kamchatka (Geist et al. 1994), southern Alaska (Plafker et al. 1989), the western face of the Klamath Mountains and Sierra Nevada in northern and central California (Wright and Wyld 1994), and the Cordillera Occidental of western Colombia and Ecuador (Van Thournout et al. 1992). Ophiolitic terranes and the soils derived from them are often high in nickel and relatively nutrient poor, and have been commonly referred to in the botanical literature as ‘‘ultramafic’’ or ‘‘ultrabasic’’ in regard to their association with distinctive plant communities.

Evolution of Regional Tectonic Models for the New Guinea Region Explaining the formation of New Guinea proved problematic until the general acceptance of plate tectonic theory and consequent understanding of island arc dynamics in the latter half of the twentieth century (Dewey and Bird 1970). Hamilton (1979) was one of the first modern geologists to attempt a regional tectonic synthesis that correctly interpreted the island as the product of a collision between a passive continental margin and a migrating island arc. He viewed the formation of the island as having resulted from collision with a single arc system that had formed above a northward-dipping subduction zone lying somewhere north of New Guinea during the Cretaceous and Early Tertiary. He hypothesized that this arc system had collided with New Guinea in the Miocene, forming the Central Ranges and Papuan Peninsula and their associated ophiolite belts. Following this collision, Hamilton proposed that the polarity of subduction had reversed, such that southward-dipping subduction was now occurring beneath the island in the vicinity of the Schouten Archipelago of northern Papua New Guinea. He also correctly noted that the Banda Arc was in the first stages of colliding with the southwestern margin of New Guinea in the Bomberai Peninsula region of Indonesian New Guinea. Kroenke (1984), in a synthesis covering the eastern half of New Guinea and areas eastward to the Solomons, Vanuatu, and Fiji, recognized four arc systems that he felt had been involved in the formation of the island: the Papuan, Trobriand, and Solomons arcs, which had already collided with New Guinea, and a fourth, the Bismarck Arc, that was in the process of doing so. He viewed these arcs as having been formed by alternating episodes of subduction along two major zones, one offshore of northern New Guinea, the other in the Solomon Islands. According to Kroenke’s (1984) model, both of these zones represented persistent and pervasive lines of weakness in the crust, so that each time subduction along one of zones became inactivated, it would reactivate along the other. In this regard, his model was to a large extent a more elaborate version of that proposed by Hamilton (1979), but incorporating more arcs and reversals of subduction. Kroenke hypothesized the following sequence of tectonic events in eastern New Guinea (for a CD-ROM video of these postulated plate and arc motions, consult Yan and Kroenke 1994). First, in the Cretaceous, rifting and basin formation oc-

................. 16157$

$CH5

02-08-07 10:22:58

PS

PAGE 142

Tectonic Geology of Papua / 143

curred along the passive northern Australian margin. This rifting, which took place in several phases, isolated small slivers of continental crust outboard of small, pull-apart ocean basins. These crustal slivers have been referred to as the ‘‘Inner Melanesian Arc’’ by many biogeographers, although this is a misnomer because they are not true arc systems in the sense discussed above. While there may have been some arc-related activity and back-arc spreading in these marginal basins during the final phases of this process, it was for the most part a continental rifting event, similar to the process seen in the current day Great Rift Valley of eastern Africa. Second, in the Middle Eocene north- and eastward-dipping subduction occurred below the Pacific Plate along the Aure-Moresby-Pocklington trench system, lying far north and east of the rifted Australian margin. This produced a southward-migrating arc (termed the ‘‘Papuan Arc’’ by Kroenke) in an oceanic setting. Third, in the Early Oligocene the eastward-dipping subduction below the Papuan Arc ceased as this arc collided with the Australian margin, causing overthrusting of arc terranes onto the Australian craton in both central New Guinea and Vogelkop. To accommodate continuing convergence between the Australian and Pacific plates, a new westward-dipping subduction zone developed far to the east, at the edge of the Australian Plate margin, forming the North Solomons Trench in what is now the Solomon Islands. This subduction resulted in the creation of an eastward-migrating arc (Kroenke’s ‘‘Solomons Arc,’’ the progenitor of the modern Solomons) in an isolated oceanic setting; western extensions of this arc, linked by transform faults, were speculated to have extended to the area north of modern Papua Province in Indonesian New Guinea. Fourth, in the Early Miocene the eastward-migrating Solomons Arc collided with the submarine flood basalts of the Ontong Java Plateau, which were too dense and massive to be subducted. This collision jammed the North Solomons Trench, ending this episode of subduction in the Solomons zone. Once again, continuing convergence between the Australian and Pacific plates shifted subduction to a new point of weakness, in this case reactivating subduction adjacent to New Guinea along the Wewak and Trobriand trenches (Hall 1997). This subduction was southor westward-dipping, in contrast to the previous east-dipping subduction in this region during the Eocene and Oligocene, and resulted in onshore volcanism on New Guinea itself, the formation of an island arc to the east (Kroenke’s ‘‘Trobriand Arc’’), and the gradual consumption of the sea floor separating the Solomons Arc islands from northern New Guinea. Fifth, in the Middle Miocene, as the intervening sea floor was eliminated via subduction, terranes associated with the Solomons Arc system collided obliquely with northern New Guinea from west to east (Davies et al. 1996; Hall 1997). Because these terranes could not be subducted into the trench, subduction in the New Guinea zone ended for a second time. This collision was a prolonged event that continued into the Pliocene, with the last unit to be sutured consisting of the Adelbert-Finisterre Terrane, north and northwest of modern Lae in Papua New Guinea. To accommodate continuing plate convergence, subduction was once

................. 16157$

$CH5

02-08-07 10:22:59

PS

PAGE 143

144 / d a n a . p o l h e m u s

again reactivated in the Solomons zone along the South Solomons Trench; this new subduction was eastward-dipping, in contrast to the previous westwarddipping polarity of the old North Solomons Trench. Far to the west, in Papua, the Vogelkop Peninsula was also sutured to the main body of New Guinea at this time. Sixth, in the Holocene, with the two zones of crustal weakness in the Solomons and New Guinea that had accommodated plate convergence throughout the Tertiary coming into ever closer juxtaposition, complex fracturing began to occur along the plate boundary zone northeast of New Guinea, creating many small arcs and subduction zones. Among these was the New Britain Arc, which began to advance southeastward over the Solomon Sea. At the western end of New Guinea, the accreted terranes of northern Vogelkop were dismembered by left lateral faulting, while the Banda Arc began the first stages of collision from the southwest. Kroenke’s model thus stressed the importance of subduction zones as persistent points of structural weakness in the crust, which could be reactivated by redistribution of stresses following arc collisions with non-subductable elements such as continental margins or submarine flood basalt plateaus. The model also proposed that two major arc collisions had contributed to the orogenies of New Guinea, the first being an Eocene–Oligocene collision that had formed the Central Ranges, and the second a Miocene collision that had formed the northern coastal ranges. Another important perspective in regard to the multiple arc hypothesis was provided by Pigram and Davies (1987). These authors agreed with Kroenke that several arc collisions had been involved in the formation of the island, and based on a large amount of field reconnaissance identified 32 tectonostratigraphic terranes lying outboard of the old Australian cratonic margin. Similar to Hamilton (1979) and Kroenke (1984), they accepted the existence of a southward-migrating arc that formed somewhere north of the Australian margin and subsequently collided with it, emplacing the ophiolites of the Central Ranges, although they postulated the timing of this collision to be Late Oligocene. They also proposed an offshore assembly for the Papuan Peninsula, their East Papua Composite Terrane, with this mega-terrane eventually suturing to the main body of New Guinea in the Miocene. In accord with Kroenke, Pigram and Davies also proposed that additional island arc collisions had occurred along the northern margin of New Guinea from the Miocene into the early Pliocene, but considered the sequence of events to have involved various microplates whose histories were too complicated and poorly understood for any mechanism to be proposed at the time. Most recently Hall (2002) and Hill and Hall (2003), using a wealth of new data, refined the preceding hypotheses in the context of a broader-scale regional tectonic model. The latter work stressed in particular the importance of the ‘‘Tasman Line,’’ which separates thick, strong, old Australian lithosphere in the west, underlying southern Papua Province in Indonesia, from thin, weak lithosphere in the east, underlying nearly all of Papua New Guinea. They ascribed the weakness of the latter lithosphere to a very old episode of terrane accretion that occurred along the eastern Australian continental margin from the Proterozoic to the Triassic

................. 16157$

$CH5

02-08-07 10:22:59

PS

PAGE 144

Tectonic Geology of Papua / 145

(600–250 mya), followed by an episode of extension that occurred along this same margin in the Cretaceous (130–75 mya). The latter event was the same rifting episode postulated by Kroenke (1984) as being responsible for opening the Tasman Sea, New Caledonia, and Coral Sea basins, and the consequent isolation of the Lord Howe Rise, Norfolk-New Caledonia Ridge, and Papuan Plateaus, respectively (Crawford et al. 2003), which now lie to the east and north of Australia as we know it today. Hall’s (2002) regional model recognized four major tectonic episodes that had shaped the geography of the New Guinea region, as follows: First, in the Early Eocene (50 mya), in general accord with Kroenke (1984), Hall’s model depicted the Central Ranges of New Guinea as being composed of accreted terranes from an arc that formed over a northward-dipping subduction zone, which may have been in existence by the Late Cretaceous. This arc system is still not well understood, but Hall suggests that it may have extended from Sundaland to New Caledonia. In Hall’s model the ophiolites derived from this arc were emplaced in medial New Guinea in the Eocene, in contrast to Kroenke (1984) who dated their emplacement to the Oligocene. Both models may be to some extent correct; the arc collision was oblique from west to east, and the collision may have continued gradually from the Eocene into the Oligocene, with the last suturing occurring in the east. Second, in the Middle Eocene (45 mya) major plate boundary shifts occurred across the entire Melanesian region, possibly due to the collision of India with southern Asia. Both the Hall and the Kroenke models agree that this tectonic rearrangement resulted in the initiation of an island arc (the ‘‘Melanesian Arc’’ of Hall, the ‘‘Solomons Arc’’ of Kroenke) over eastward-dipping subduction in what is now the Solomon Islands region. Hall’s model also suggests that this arc included the early cores of what would later become New Britain and the island of Viti Levu in Fiji. At the same time, westward-dipping subduction was initiated along the eastern margin of the Philippine Sea Plate (which lies outside the region modeled by Kroenke), with the eastern section of that plate adjacent to the subduction zone becoming detached to form the Caroline Plate. Continuing westward-dipping subduction along the eastern margin of this new plate created a north-south oriented arc lying northeast of New Guinea, termed by Hall (2002) the ‘‘East Caroline Arc.’’ Third, in the Late Oligocene (25 mya), once again, both the Hall and the Kroenke models agree that the Ontong Java Plateau collided with the subduction zone in the Solomons region, shutting down subduction there and shunting it to the south of the Solomon Sea, under New Guinea, in the process forming an arc to the east (the ‘‘Maramuni Arc’’ of Hall, or the ‘‘Trobriand Arc’’ of Kroenke). The existence of this postulated arc is, however, equivocal; despite a predicted 1,000 km of subduction in both tectonic models, Hall and Spakman (2003) could find no indication of the expected tomographic signature from such a subducted slab. To the north of New Guinea at this same time, the Caroline Plate was rotating clockwise, bringing the arc islands along its eastern flank into closer proximity to

................. 16157$

$CH5

02-08-07 10:23:00

PS

PAGE 145

146 / d a n a . p o l h e m u s

northern New Guinea (Figures 2.1.4, 2.1.5). These arc islands on the eastern Caroline Plate also moved past the western end of the Solomons, creating for a brief period during the Late Oligocene to Early Miocene an elongate, nearly continuous archipelago from the Philippine region eastward to Fiji. This reconstruction is similar in some respects to Kroenke’s (1984) concept of a Solomons Arc in the east connected by transform faults to other arc sectors somewhere north of New Guinea, but if such an extended archipelago did exist it was not a long-lived configuration. Fourth, in the Late Miocene (5 mya), Hall’s model postulates that continued

Figure 2.1.4. Tectonic reconstruction of the New Guinea region in the Middle Oligocene (30 MYA) following Hall (2002). By this time period the Early Tertiary accretion of island arc terranes from the first episode of collision in the mountains of central New Guinea was already complete. Vogelkop was still a detached microcontinent west of New Guinea proper and islands that would later become parts of Halmahera and Waigeo lay directly north of New Guinea at the southwest corner of the Caroline Plate. Barbs at plate boundaries are on the overriding plates and indicate the direction of subduction beneath the plates. For further discussion of these tectonic events see the text.

................. 16157$

$CH5

02-08-07 10:23:37

PS

PAGE 146

Tectonic Geology of Papua / 147

Figure 2.1.5. Tectonic reconstruction of the New Guinea region in the Middle Miocene (15 MYA) following Hall (2002). In this time period the Vogelkop microcontinent was in the process of being sutured onto the western side of New Guinea proper, creating the limestone anticlines of the Bird’s Neck region. Islands that would later become parts of Halmahera and Waigeo were in close proximity to the northwestern New Guinea coast and an island arc that had formed along the southeastern margin of the Caroline Plate in the Oligocene was beginning to collide obliquely with central and northern New Guinea, with this collision beginning on the central section of the northern coast and progressing slowly from west to east. Barbs at plate boundaries are on the overriding plates and indicate the direction of subduction beneath the plates. For further discussion of these tectonic events see the text. rotation and convergence of the Caroline Plate with northern New Guinea resulted in the emplacement of arc terranes from the Caroline Plate’s eastern margin in sequential fashion from west to east, forming the New Guinea north coastal ranges (Figure 2.1.6). By this time, the Solomon Sea had also began subducting to the northwest under southern New Britain, producing the impressive volcanism now seen on that island at Rabaul and elsewhere.

................. 16157$

$CH5

02-08-07 10:24:13

PS

PAGE 147

148 / d a n a . p o l h e m u s

Figure 2.1.6. Tectonic reconstruction of the New Guinea region in the Early Pliocene (5 MYA) following Hall (2002). By this time period the Vogelkop microcontinent had become sutured to the main body of New Guinea; Halmahera and Waigeo had attained some semblance of their present form and moved westward beyond New Guinea to near their current positions; and the island arc that had formed along the southeastern margin of the Caroline Plate was almost entirely sutured to northern New Guinea, with the Adelbert-Finisterre Terrane of the Huon Peninsula in the final stages of collision, and New Britain still a separate island (as it remains today). Strike-slip faulting along the Sorong Fault Zone had also begun to transport rifted slivers of New Guinea westward toward Sulawesi as direct collision between the Australian and Pacific plates was gradually replaced by a regime of left-lateral shear. Barbs at plate boundaries are on the overriding plates and indicate the direction of subduction beneath plates. For further discussion of these tectonic events see the text.

................. 16157$

$CH5

02-08-07 10:24:54

PS

PAGE 148

Tectonic Geology of Papua / 149

Hall’s model highlights the importance of the period from 52–48 mya, which marked the initiation of an extensive line of subduction along the entire margin of the western Pacific from the Marianas to Fiji, a distance of 3,000 km. The causes of this major tectonic rearrangement are not known, although as noted above some authors have proposed that it was linked to the collision of India with southern Asia, which appears to have altered plate trajectories in the entire Indo-Pacific region (Veevers et al. 2000a, 2000b). The most obvious manifestation of this change in tectonic regime was a sudden transition from the eastward-dipping subduction that had prevailed between 55–45 mya and caused arc systems to migrate westward toward the rifted slivers (such as New Caledonia) lying outboard of the eastern Australian margin, to a westward-dipping subduction that caused arcs to migrate eastward away from Australia. This is also the same time period during which the large ophiolite exposures are postulated to have been emplaced in the Central Ranges of New Guinea, on the Papuan Peninsula, and in New Caledonia, suggesting that in addition to global changes in plate circuits that might have been caused the by the India-Tibet collision far to the west, a trans-regional episode of nearly simultaneous trench jamming by large arc terranes may also have rendered continued east-directed subduction impossible across much of Melanesia, forcing the subduction flip (Kroenke 1984). By the early 2000s, then, tectonic models had evolved and been refined to the point that there was modest agreement about the broad tectonic outlines of New Guinea’s formation, at least for the central body of the island. Much of this work had centered on Papua New Guinea due to far better field reconnaissance and research university support in that country under the period of Australian administration, but with the development of the Grasberg mine in western New Guinea, increasingly detailed field mapping became available from the Indonesian half of the island as well. The information from Papua largely supported the conclusions drawn from the data obtained in Papua New Guinea, but also highlighted the fact that the Vogelkop sector had experienced a different history. Tectonic geologists are in general agreement that the core of the Vogelkop Peninsula is a section of the Australian craton (the Kemum Terrane of Pigram and Davies 1987) that became detached from the main continental mass sometime in the Mesozoic (Hamilton 1979). For instance, both Australia and Vogelkop share a Late Paleozoic to Early Mesozoic fossil Glossopteris flora, and have similar paleomagnetic polar wander paths (Giddings et al. 1985). Pigram and Davies date the separation of Vogelkop from northwestern Australia as Early Cretaceous, but the resulting terrane did not move far (Hill and Hall 2003), probably remaining slightly to the southwest of the main section of New Guinea, and possibly undergoing in-place clockwise rotation of up to 90 (Hamilton 1979). During the Early Tertiary the original core of Vogelkop was expanded by the fusion of the continentally-derived Misool terrane to its western margin and the arc-related Tamrau Terrane to its northern edge (Pigram and Davies 1987). This entire composite unit then moved eastward and was reintegrated into greater New Guinea in the Miocene. The Vogelkop suture zone is marked by faults bounding the Wandammen

................. 16157$

$CH5

02-08-07 10:24:54

PS

PAGE 149

150 / d a n a . p o l h e m u s

Peninsula, and by the stacked anticlines of continental shelf limestone in the Lengguru Fold Belt in the Bird’s Neck region. Although Vogelkop thus had a separate history from the remainder of New Guinea, this history was in many ways very similar to that of the main island, in that a cratonic core accumulated arc terranes along its northern margin. In addition, once integrated, both Vogelkop and central New Guinea experienced extensive left-lateral shearing of these accreted terranes by faults that developed in the Late Tertiary as collision with the Pacific Plate was increasingly accommodated by shear rather than direct collision.

Tectonic Provinces of New Guinea All of the geologists whose models were discussed above have recognized that due to its history of collision and accretion, New Guinea consists of several discrete tectonic provinces that run parallel to each other in a roughly east-west direction. These consist of a Stable Platform in the south, consisting of the old, undeformed Australian continental craton; the Fold Belt in the center, consisting of the deformed margin of this craton; and the Mobile Belt in the north, comprised of a complex mix of accreted arc terranes and tectonic slivers, many of which are now being displaced westward in a left-lateral fashion along various fault zones (Hamilton 1979; Pigram and Davies 1987; Charlton 1996; Hill and Hall 2003). A more detailed examination of each of these provinces is useful in understanding the geomorphology and petrology of the modern island.

the australian craton The Australian continental craton, also referred to as the ‘‘Stable Platform’’ by Hill and Hall (2003), comprises most of the southern half of New Guinea and consists of undeformed sedimentary rocks of predominantly Triassic age overlying a Precambrian basement (Hamilton 1979). The northern edge of this Australian craton, at the site of what would later become New Guinea, represented a passive margin throughout much of the Mesozoic, from the Triassic until the Middle to Late Cretaceous, when renewed rifting began along its eastern margin, followed subsequently by island arc collisions from the north. Although the Cretaceous coastline of proto-New Guinea lay in a roughly east-west orientation, an elevated ridge extended northward from it in the area of the current Papua New Guinea-Indonesia border, reaching as far as the Border Mountains west of the current Sepik Basin (Hamilton 1979; Davies 1990). This elevated ridge remained above water during the Mesozoic when most of the remaining platform was submerged (Davies 1990), and the reefs that formed along its eastern flank are now exposed as the limestone Darai Plateau of southern Papua New Guinea. Evidence for this ancient ridge can also still be seen in the drainage patterns of modern New Guinea, which flow away from it to both the east (Sepik and Fly rivers) and west (Mamberamo and Digul rivers). This structure has also had an important influence on the zoogeography of the island, because it represented a subaerial salient of Australia that projected

................. 16157$

$CH5

02-08-07 10:24:55

PS

PAGE 150

Tectonic Geology of Papua / 151

northward into the tropics and subtropics when the rest of the continent was still in cooler climatic zones. It also seems to have acted like a tectonic wedge, splitting the accreted arc terranes that arrived in the first episode of collision from the Late Cretaceous to the Oligocene. As noted above, the Vogelkop Peninsula for most of its history was a detached piece of this Australian continental craton that separated from the remainder of the platform in the Mesozoic. The Vogelkop Peninsula was never far away from the northwestern margin of the main Australian continent, however, because its post-Cretaceous tectonic history has been substantially similar to that of the main body of New Guinea, and it accumulated at least one layer of arc terrane laminations onto the northern margin of its continental core in generally the same historical sequence as the remainder of the island. In addition to the separation of Vogelkop in the west, several additional pieces of the Australian craton became detached along its eastern margin in the Late Mesozoic. This was apparently the result of the same extensional episode, discussed further below, in which continental slivers were pulled away northward or eastward, forming the Tasman Sea and New Caledonia Basin, bordered on their eastern margins by the Lord Howe Rise and Norfolk Ridge respectively. Subsequently, this same episode of extension led to the opening of the Coral Sea, which was also bounded on its northern margin by a narrow continental sliver. Although most of these events happened too far to the east to have a direct impact on the geological evolution of Papua, they would set in motion a series of later arc collisions and subduction reversals that would have an important influence on patterns of terrane accretion in western New Guinea. Unlike the situation in the west, where much remains to be learned about the rifting history of the Vogelkop and Misool terranes, a considerable amount of recent research has been devoted to the tectonics of the east Australian cratonic margin during the Cretaceous. Although there is uniform agreement among tectonic geologists regarding the general sequence of events involved in the rifting of this margin, the timing and the mechanisms that produced them are poorly understood. Crawford et al. (2003) considered the rifting to have begun 130–120 mya, and hypothesized that the extension was the result of the rollback of a rapidly retreating slab associated with a westward-dipping subduction zone that was active from 100–45 mya, possibly accelerated by a mantle plume. In their scenario, all of Australia east of the Tasman Line was created by arc accretion from systems migrating westward out of the Pacific from 600 mya to the Triassic. In the Late Cretaceous, this pattern then reversed itself and this composite margin was pulled apart. Gaina et al. (2003), by contrast, found little evidence for subduction and slab rollback, but did concur with the existence of a thermal anomaly under the east Australian margin in the Late Cretaceous between 100–60 mya, with the parallel sequence of continental slivers and intervening ocean basins being the result of ridge-plume interactions. Srolias et al. (2003) supported a similar timing for initiation of rifting along this margin, beginning at 100–90 mya, rather than at 130 mya

................. 16157$

$CH5

02-08-07 10:24:56

PS

PAGE 151

152 / d a n a . p o l h e m u s

as proposed by Crawford et al. (2003). All these recent studies do agree that the opening of the Coral Sea occurred subsequent to that of the Tasman Sea, beginning around 61 mya (Gaina et al. 1999) and ceasing by 52 mya (Gaina et al. 1999; Sdrolias et al. 2003). The Coral Sea Basin was the last of the marginal basins to open, and was also bounded by several narrow, elongate fragments of continental crust rifted from the northeastern margin of Australia. These slivers, referred to as the Eastern and Papuan Plateaus (Pigram and Davies 1987), were apparently never emergent above the ocean, but did constitute a topographic barricade against which southward migrating arc systems eventually collided, forming the Papuan Peninsula. In addition, another cratonic fragment, now present in the Jimi Terrane of Papua New Guinea (i.e., the Bismarck-Kubor block), rifted from the Australian margin and functioned as a separate island during the Late Cretaceous before being reintegrated into New Guinea at or near the time of the first arc collision episode (Pigram and Davies 1987; Davies 1990). Unlike the continental slivers north of the Coral Sea, this Bismarck-Kubor block does appear to have remained subaerial (i.e., above water) from the Cretaceous onward (Davies 1990). Other subaerial blocks of continental crust may also have been rifted from northern Australia and followed similar histories as small islands north of the main craton in the New Guinea region during the Late Cretaceous, but if so, then all of them were trapped and crushed between approaching island arcs and the Australian margin in the Eocene (Rogerson and Hilyard 1990).

the fold belt The exact mechanism by which the high central mountain ranges of New Guinea were uplifted, as well as the timing of this uplift, is still a matter of some debate. An island arc collision was probably involved, and as discussed above is considered to have occurred sometime between the Paleocene and Late Oligocene in western New Guinea (Hamilton 1979; Hill and Hall 2003), and perhaps extending through the Late Eocene and Early Oligocene in Papua New Guinea (Kroenke 1984; Pigram and Davies 1987; Davies 1990; Pigram and Symonds 1991; Davies et al. 1996). This collision progressed obliquely from west to east over a period of 5–10 million years. Hall (2002) depicts the arc system involved as forming over a northerly dipping subduction system that extended from Java to New Caledonia at 55 mya, with collision complete in central New Guinea by 45 mya; this same hypothesis was subsequently reiterated by Hill and Hall (2003). As noted by Davies et al. (1996), the orogeny related to an arc collision typically occurs millions to tens of millions of years following the collision itself, due to initial accommodation of the collision by crustal foreshortening. Thus estimates of timing for the subsequent uplift of the Central Ranges vary from Eocene (Davies et al. 1996) following Paleocene onset of collision, to Late Miocene (Hill and Hall 2003) following Oligocene collision. In either case, this initial episode of arc accretion is believed to have emplaced the overthrust ophiolite and metamorphic belts of central Papua Province (the Rouffaer Terrane of Pigram and Davies 1987),

................. 16157$

$CH5

02-08-07 10:24:56

PS

PAGE 152

Tectonic Geology of Papua / 153

along with correlative April Ultramafics, Marum Complex, and Papuan Ophiolite Belt in Papua New Guinea (the Sepik, Marum, and Bowutu terranes of Pigram and Davies 1987), and possibly the Tamrau Mountains of Vogelkop. As with the formation of the central mountains themselves, the ages and times of emplacement for of these various ophiolites in New Guinea are not well constrained. All of them seem to be products of an island arc or arcs that initially formed somewhere north of New Guinea in the Late Mesozoic before colliding with the Australian margin in the Early Tertiary (Davies and Jaques 1984; Davies et al. 1997; Hall 2002). By contrast, the ophiolites of the Cyclops Mountains, on the north coast near Jayapura, are only of Oligocene age and were emplaced considerably later (Monnier et al. 1999), probably by a completely different arc collision event. The mode of formation for the central Papuan ophiolites is also the subject of debate, with Monnier et al. (2000) considering them to have formed in a back-arc setting, while Hill and Hall (2003) interpreted them as being of fore-arc derivation. Further evidence for timing of the arc collision in the Central Ranges has come from field mapping done in Papua Province during the last decade by PT Freeport Indonesia, which indicates that there are various small exposures igneous rocks of Eocene to Oligocene age present along the northern margins of the ophiolite belt (Freeport geologists, pers. comm.). These are more numerous than shown on the geological map of Dow et al. (1986), and are in the proper position and juxtaposition to the ophiolites to potentially represent the remains of volcanic islands that accreted behind a more massive fore-arc ridge (similar to the situation seen today in the eastern limb of the modern Sunda-Banda arc system). In the middle reaches of the Wapoga River system, these igneous formations form the mountain foothills (Figure 2.1.7) and consist of undated deposits of andesite and pyroclastics (such as tufa) with intrusions of diorite, the latter flanked by veins of dark black hornfels caused hydrothermal alteration of the adjacent formations. The presence of these volcanic formations to the north of the ophiolites would seem to argue in favor of the Hill and Hall (2003) hypothesis that the Papuan ophiolites formed in a forearc setting. Once again, it is important to bear in mind that the accreted arc terranes of the New Guinea Central Ranges are tens of millions of years older than their hypothesized times of emplacement, while the uplift that followed their collision is by contrast tens of millions of years younger. It is also not uniformly agreed that this Early Tertiary arc collision actually formed the high Central Ranges that we see today (Figure 2.1.8). In central Papua, various authors have hypothesized that a subsequent episode of Miocene arc collision along the north coast caused a massive deformation of the Australian continental craton south of the overthrust ophiolitic and metamorphic terranes of the Irian Jaya Mobile Belt that had been emplaced in the Late Eocene or Early Oligocene, pushing the shallow water Miocene limestones that had formed on the southern flank of the Fold Belt to elevations of over 5,000 meters (Nash et al. 1993; Hill and Raza 1999; Hill and Hall 2003). Under this model, the resulting orogeny is hypothesized to have created a large, overthrust anticlinal structure, the Mapenduma Anticline, which forms the

................. 16157$

$CH5

02-08-07 10:24:57

PS

PAGE 153

154 / d a n a . p o l h e m u s

Figure 2.1.7. The northern face of the New Guinea central ranges in the Wapoga River basin of west-central New Guinea, looking toward the southwest. The northward-flowing Tirawiwa River in the foreground is exiting foothills composed of Tertiary volcanics; the higher ranges in the distance are composed of the ophiolites of the Rouffaer Terrane, emplaced in the Eocene–Oligocene period as the result of an island arc collision. Note the braided channel of the Tirawiwa River, which is overburdened with alluvium from these young mountains, and is typical of New Guinea rivers in areas subjected to rapid elevation driven by tectonic collisions. Photo: D. A. Polhemus.

core of the Central Ranges in Indonesian New Guinea and is suggested to be structurally related to the Muller Anticline of west central Papua New Guinea. The anticline hypothesis was also supported by Weiland and Cloos (1996), who proposed that the structure had been eroded in an asymmetrical fashion from the south, due to orographically induced rainfall, producing the huge escarpments that currently form the southern flank of the present day Central Ranges in Papua Province (Figure 2.1.9).

the mobile belt The Mobile Belt consists of various intermingled arc terranes that were laminated onto northern New Guinea by a series of arc collisions between the Oligocene and Pliocene (Figures 2.1.4–2.1.6). These terranes were unrelated to those involved in the formation of the Central Ranges, and instead had their genesis along the margins of the Caroline and Solomon Sea plates. They appear to include, from west to east, the Arfak Mountains, Biak, Yapen, the Van Rees, Foja, Cyclops, Torricelli, Prince Alexander, Adelbert, Finisterre, and Saruwaged mountains (corresponding

................. 16157$

$CH5

02-08-07 10:25:06

PS

PAGE 154

Tectonic Geology of Papua / 155

Figure 2.1.8. A view of the summit region of the Puncak Jaya massif, looking to the northeast. Note the overthrust limestone strata capped by glacial ice fields along the ridge crest, and the pronounced tilt of the limestone strata, which dip to the north (to the left of the picture) toward the basin of the Mamberamo River, and break away sharply to the south (to right of the picture) in spectacular escarpments, the summits of which lie at elevations in excess of 4,500 m. Photo: D. A. Polhemus.

to the Arfak, Gauttier, Cyclops, Torricelli, Prince Alexander, and Finisterre terranes of Pigram and Davies 1987), and possibly portions of the Papuan Peninsula. As with the arc terranes of the central mountains that preceded them, these terranes converged obliquely from west to east, with collision beginning in the Late Miocene in Indonesian New Guinea, but continuing until the Pliocene in Papua New Guinea (Cooper and Taylor 1987). The collision histories of the initially accreted terranes in this assemblage, which lie in Indonesian New Guinea, are poorly understood, while the histories of the most recently accreted terranes, such as the Adelbert-Finisterre Terrane of northern Papua New Guinea, are by contrast well studied. Nearly all tectonic reconstructions agree that the Adelbert-Finisterre Terrane accreted to New Guinea in the Late Miocene or Early Pliocene, and that it was the last of the Miocene terranes to arrive (but for a dissenting opinion see Findlay 2003). Kroenke (1984) provides a structural and tectonic analysis linking this terrane to the Solomons Arc system via a transform, and Pigram and Davies (1987) similarly suggest that it was previously part of a system that included the Bismarcks and Solomons, and formed in an isolated oceanic setting to the northeast of New Guinea. Given its late arrival in comparison to other Miocene terranes linked to Kroenke’s postulated Solomons

................. 16157$

$CH5

02-08-07 10:25:14

PS

PAGE 155

156 / d a n a . p o l h e m u s

Figure 2.1.9. A view of the southern face of the New Guinea Central Ranges near the Grasberg mine, looking west along the axis of the uplift. The exposed rocks of the Australian craton have been massively deformed by several episodes of island arc collision during the Tertiary. Photo: D. A. Polhemus.

Arc system, it is possible that this terrane occupied an isolated position on a sector of the arc lying between the Solomons to the east and the other Miocene terranes to the west. The arrival of these terranes on New Guinea’s northern margin may have had effects across a broad section of the island, as alluded to above. In Indonesian New Guinea this Miocene arc collision is hypothesized to have caused a massive deformation of the Australian continental craton south of the overthrust ophiolitic and metamorphic terranes of the Mobile Belt. The tectonic history of western New Guinea in the early to middle Miocene is, however, still uncertain, particularly in regard to the accretion of terranes in the northern coastal ranges and the formation of the Weyland Mountains in the Bird’s Neck region (Milsom 1991). For instance, a block of Miocene volcanic rocks is present along the southern coast of Cenderawasih Bay from Nabire westward (Dow et al. 1986), and may well represent the westernmost arc fragment to have accreted to the New Guinea mainland in this second episode of arc accretion, and prior to the reintegration of Vogelkop. But for the moment any assignment of such Miocene terranes to a westerly extension of the Solomons Arc system of Kroenke is tentative at best, pending more detailed geological and biogeographic analyses. It is also important to remember once again that the broad-scale collision between the Australian and Pacific plates in New Guinea region was not head-on

................. 16157$

$CH5

02-08-07 10:25:22

PS

PAGE 156

Tectonic Geology of Papua / 157

but rather oblique, occurring sequentially from west to east. Therefore, arc collision episodes in western New Guinea were generally initiated tens of millions of years earlier than related events in eastern New Guinea. Further, once convergence between the plates had been accommodated by initial collision and foreshortening, the continuing convergence between the plates was subsequently accommodated by left-lateral shearing along strike-slip faults (McCaffrey 1996). This change in collisional regime was initiated by 25 mya (Hall and Spakman 2003) and continues to the present day as the Pacific Plate slides westward relative to northern New Guinea at rates of up to 7 cm per year (McCaffrey 1996). As with the arc collisions that preceded it, this shearing commenced at earlier times in western New Guinea than in the eastern half of the island. Across the main body of northern New Guinea this zone of shear does not constitute a single narrow fault trace, but instead occupies a broad zone of strike-slip faults that runs through the entire Mobile Belt. As a result of this shear, fragments of various terranes that had been accreted into the island via earlier arc collisions have now been slivered apart and carried to the west, in some cases for considerable distances. For instance, the narrow, elongate island of Yapen appears to be a fault sliver that has been carried westward from the Van Rees Mountains across the top of Cenderawasih Bay. A few of the fault traces within the Mobile Belt extend well westward from New Guinea, and may have been involved in the westward transport of Papuan tectonic fragments into the Sulawesi region. For instance, the most prominent element of this left-lateral system, the Sorong fault zone, extends from near the mouth of the Mamberamo River, across Cenderawasih Bay and the top of Vogelkop, then between Halmahera and Obi in the Moluccas before terminating at the Sulu Archipelago immediately east of Sulawesi, a distance of over 1,500 km (Hamilton 1979; Baker and Malaihollo 1996; McCaffrey 1996). The Miocene accreted terranes of the Vogelkop Peninsula have been extensively sheared by this fault system; for instance, Pigram and Davies (1987) hypothesized that the Arfak Mountains, Yapen, and portions of Biak/Supiori all represented portions of a single original terrane now strung along this fault trace, while Hamilton (1979) considered Batanta to be a fault-displaced fragment of the Tamrau Mountains of northern Vogelkop, now shifted westward to a position north of Salawati and separated from that latter island by the deep, narrow Sagewin Strait that marks the axis of the fault itself. Similarly, the islands of Bacan, Obi, and the Banggi-Sula group in the Moluccan region are hypothesized to be fragments of New Guinea that have been carried westward along this same Sorong Fault system to positions far west from their original points of origin (Malaihollo and Hall 1996; Vroon et al. 1996; Charlton 1996). The net result of this Tertiary shearing has been to create a chaotic jumble of tectonic fragments across the entire northern third of New Guinea. Attempting to determine the original provenance of such fragments can be difficult, in that it requires reassociating them with source blocks that may lie hundreds of kilometers to the east. Such fault slivers have also had the potential to transport biota, leading to intermixed biogeographic patterns, with tectonic blocks that were once part of

................. 16157$

$CH5

02-08-07 10:25:23

PS

PAGE 157

158 / d a n a . p o l h e m u s

the main body of New Guinea now integrated into Vogelkop, and Vogelkop fragments now lying even further west, near Halmahera or even Sulawesi. Even with such mixing, the accretion of the Miocene arc terranes into northern New Guinea created a pronounced biogeographic signature, with many endemic species confined to the northern coastal ranges of the island and having their closest relationships to taxa in the Bismarcks or Solomons (Boer and Duffles 1996; Polhemus and Polhemus 1998).

present day convergent arc systems In addition to the systems discussed above, two other arcs, the New Britain Arc and the Banda Arc, are advancing toward New Guinea, and will collide with the island over the next ten million years if current plate motions continue. The Sunda Arc system (known as the Banda Arc at its eastern extremity), lying immediately to the west of New Guinea, is a textbook island arc formed at the boundary of the Australian and Eurasian plates above the well-defined Sunda subduction zone. It has yet to directly collide with New Guinea, but has been folded back north and westward into a fishhook shape by the northward motion of the Australian craton. The New Britain Arc is a very young system formed over a trench at the northern margin of the Solomon Sea, which is gradually closing from west to east as New Britain and the Papuan Peninsula converge on each other like a pair of scissor blades. Although these incipient collisions have not yet emplaced terranes into the main body of the island, they have provided dispersal corridors for biota, and produced buckling of the continental shelf limestones in the west (Polhemus 1996), creating young uplifts such as the Fakfak Peninsula.

Continuing Questions and Puzzles Given its tectonic complexity, and the large amount of field reconnaissance still to be undertaken in its rugged and heavily forested terrane, it is not surprising that much remains to be understood regarding the geological evolution of New Guinea. In Papua, for instance, our knowledge of local surface geology in many areas is still based on field surveys from the Dutch colonial era (Visser and Hermes 1962), while much of the more recent survey work has been undertaken by oil or mining companies and is thus proprietary information not readily available to the scientific community. As a result, although the tectonic models reviewed above now provide a broad picture of the island’s formation and evolution, many key areas of Papua continue to present geological questions. Waigeo Island, for instance, is of the correct age and composition to be part of the Eocene-Oligocene island arc that formed the Central Ranges (Charlton et al. 1991; Ling et al. 1991), and has had clear historical associations with greater New Guinea based on its biota (Polhemus and Polhemus 1998, 2002), but its high degree of species level endemism also implies an extended period of isolation by water gaps. The ophiolites of northern Waigeo appear correlated with those of eastern Halmahera (Hall and Nichols 1991), but the presence of thick and exten-

................. 16157$

$CH5

02-08-07 10:25:23

PS

PAGE 158

Tectonic Geology of Papua / 159

sively exposed limestone strata also indicate that much of the island was submerged during the Miocene (Charlton et al. 1991). The reconstructions of Hall (2002) and Hill and Hall (2003) show Waigeo and Halmahera forming in the Eocene along the southeastern margin of what would become the Caroline Plate and then moving westward just to the north of New Guinea and Vogelkop to their present positions (Figures 2.1.4–2.1.6). An alternate hypothesis is that Waigeo was a part of the early Tertiary arc that formed the Central Ranges, and accreted to northern Vogelkop in the area west of the Tamrau Mountains during the Oligocene, but was then sheared off and carried further westward along the Sorong Fault zone to its present position during the Pliocene. Many geological and tectonic questions also remain in regard to the Foja-Van Rees Mountains. For instance, why is this uplift bowed in a northward direction? Also, if it represents the remains of an accreted arc system, as most other north coast ranges in New Guinea do, then why is it, so far as currently known, devoid of ophiolites? Although there are indications of Late Tertiary igneous blocks within this uplift (Dow et al. 1986), the surface geology is primarily Pliocene shales and other mixed sediments, which are not typical of an accreted arc terrane. Clearly, the area is insufficiently surveyed geologically, but it would seem that the processes operating here are different than those that produced the other north coastal ranges of New Guinea. The Weyland Mountains also represent a problematic area at the western terminus of the Central Ranges. Although the core of these mountains consists of Miocene granodiorite and volcanics, they must have been in place by the Oligocene, because the geologic map of Dow et al. (1986) shows that the ophiolite belt of the Central Ranges is accreted to their northern margin (and extends even further westward, through the curve of the Bird’s Neck to the base of the Wandammen Peninsula). The overthrust metamorphic belts to the south of the Weyland Mountains are also similar to and apparently correlative with those of the remainder of the Central Ranges. Thus the Weylands are trapped between the ophiolites and the metamorphics, and perhaps represent a terrane (possibly associated with the Wandammen Peninsula) that was assembled offshore slightly to the north of the old western tip of New Guinea and then thrust southward when Vogelkop was sutured onto the island in the Miocene (Hill and Hall 2003). A final area that is poorly understood and also underwent significant tectonic rearrangement during the Miocene suturing of Vogelkop is the Wandammen Peninsula. It contains a small fragment of old continental cratonic rocks of Devonian and Silurian ages (Hamilton 1979; Pigram and Davies 1987) in fault contact across a major unconformity with much younger metamorphic rocks of Late Miocene to Late Pliocene age. The metamorphic rocks appear to have been rapidly uplifted in the last 3–4 million years, and in this respect the Wandammen Peninsula resembles the D’Entrecasteaux Islands lying immediately east of mainland Papua New Guinea. Both areas appear to represent metamorphic core complexes, and both seem to have formed in analogous settings, immediately offshore of the main body of New Guinea in areas subjected to recent faulting and rifting. By contrast, the

................. 16157$

$CH5

02-08-07 10:25:24

PS

PAGE 159

160 / d a n a . p o l h e m u s

suite of old Australian continental rocks amalgamated with this core complex may represent the remnants of a formerly separate island slivered from the northwestern Australian craton at some point in the late Cretaceous or early Tertiary, and then reintegrated during the suturing of Vogelkop. This proposed history would also account for the area’s anomalously high biotic endemism in groups such as cicadas (Boer 1986, 1994). New Guinea and the surrounding arcs and ocean basins remain one of the most tectonically complex and dynamic areas on the planet, and predicting its future is as difficult as deciphering its past. As noted by Hall (2002), ‘‘It is very difficult to predict what might be the form of the region in a few million years time. Simple logic indicates that Australia will continue to move northwards, and the Pacific will continue to move westwards relative to Eurasia. Therefore, within the Asian margin extending from Japan southwards through the Philippines to eastern Indonesia and northern New Guinea one might expect to see a relatively simple convergent mountain belt system produced as an interaction of these three plates. However, that has not been the pattern of development in this region for the last few million years, and probably much longer . . . The addition of arc crust and fragmentation of the margins of Eurasia, Australia and the Pacific has not been a simple, unidirectional process. Subduction has driven nearly all of this complex evolution.’’ Clearly, New Guinea and surrounding arcs present a fascinating set of problems for future geological study. Continuing research into the island’s tectonic history thus promises not only to refine our understanding of one of the world’s most geologically complicated and dynamic islands, but also to provide globally applicable insights into the dynamics of island arcs and continental margins in the Malay Archipelago, the Caribbean, and elsewhere around the world.

Acknowledgments The author wishes to thank Dr. Warren Hamilton of the U. S. Geological Survey, Denver; Dr. Loren W. Kroenke of the University of Hawai’i, Manoa; Dr. Kevin T. M. Johnson of the Bishop Museum, Honolulu; and Dr. Robert Hall of the Royal Holloway University of London, all of whom generously took time out of their busy schedules to educate him in the complexities of island arc geology. Any misinterpretations of their teachings and other studies cited herein are the author’s fault alone. This research was sponsored in part by a series of grants from the National Geographic Society, Washington, D.C., and by a grant from the National Science Foundation, Washington, D.C.

Literature Cited Baker, S., and J. Malaihollo. 1996. Dating of Neogene igneous rocks in the Halmahera region: arc initiation and development. In Hall, R., and D. Blundell (eds.) Tectonic

................. 16157$

$CH5

02-08-07 10:25:24

PS

PAGE 160

Tectonic Geology of Papua / 161 Evolution of Southeast Asia. Geological Society of London Special Publication 106: 499–509. Brown, B.J., R.D. Mu¨ller, C. Gaina, H.I.M. Stuckmeyer, H.M.J. Staff, and P.A. Symonds. 2003. Formation and evolution of Australian passive margins: implications for locating the boundary between continental and oceanic crust. Pp. 223–243 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Charlton, T.R. 1996. Correlation of the Salawati and Tomori Basins, eastern Indonesia: a constraint on left-lateral displacements of the Sorong Fault zone. In Hall, R., and D. Blundell (eds.) Tectonic Evolution of Southeast Asia. Geological Society of London Special Publication 106: 465–481. Charlton, T.R., R. Hall, and E. Partoyo. 1991. The geology and tectonic evolution of Waigeo Island, NE Indonesia. Journal of Southeast Asian Sciences 6: 289–297. Cooper, P., and B. Taylor. 1987. Seismotectonics of New Guinea: a model for arc reversal following arc-continent collision. Tectonics 6: 53–67. Crawfod, A.J., S. Meffre, and P.A. Symonds. 2003. 120 to 0 Ma tectonic evolution of the southwest Pacific and analogous geological evolution of the 600 to 200 Ma Tasman Fold Belt System. Pp. 383–403 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Davies, H.L. 1990. Structure and evolution of the border region of Papua New Guinea. In Carman, G.J., and Z. Carman (eds.) Petroleum Exploration in Papua New Guinea: Proceedings of the First PNG Petroleum Convention, Port Moresby, 12–14th February 1990. PNG Chamber of Mines and Petroleum, Papua New Guinea. Davies, H.L., and A.L. Jaques. 1984. Emplacement of ophiolite on Papua New Guinea. Pp. 341–349 in Gass, I.G., S.J. Lippard, and A.W. Shelton (eds.) Ophiolites and Oceanic Lithosphere. Geological Society of London Special Publication 14. Davies, H.L., R.C.B. Perembo, R.D. Winn, and P. KenGermar. 1997. Terranes of the New Guinea orogen. Pp. 61–66 in Hancock, G. (ed.) Proceedings of the Geology Exploration and Mining Conference, Madang. Australian Institute of Mining and Metallurgy, Melbourne. Davies, H.L., R.D. Winn, and P. KenGemar. 1996. Evolution of the Papuan Basin—a view from the orogen. In Buchanon, P.G. (ed.) Petroleum Exploration, Development and Production in Papua New Guinea: Proceedings of the 3rd PNG Petroleum Convention, Port Moresby, 9th–11th September 1996. PNG Chamber of Mines and Petroleum, Papua New Guinea. de Boer, A.J. 1986. Taxonomy and biogeography of the conviva group of the genus Baeturia Sta˚l 1866 (Homoptera: Tibicinidae). Beaufortia 36: 167–182. de Boer, A.J. 1994. Taxonomy and biogeography of the guttulinervis group of the genus Baeturia Sta˚l 1866 (Homoptera: Tibicinidae). Bijdragen tot de Dierkunde, 64: 87–100. de Boer, A.J., and J.P. Duffels. 1996. Biogeography of Indo-Pacific cicadas east of Wallace’s Line. Pp. 297–300 in Keast, A., and S. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Publishing, Amsterdam. Dewey, J.F., and J.M. Bird. 1970. Mountain belts and the new global tectonics. Journal of Geophysical Research 75: 2625–2647. Dow, D.B. 1977. A geological synthesis of Papua New Guinea. Bureau of Mineral Resources Australia, Bull. 201: 1–41.

................. 16157$

$CH5

02-08-07 10:25:25

PS

PAGE 161

162 / d a n a . p o l h e m u s Dow, D.B., G.P. Robinson, U. Hartono, and N. Ratman. 1986. Geologic map of Irian Jaya, Indonesia. Geological Research and Development Centre, Ministry of Mines and Energy, Bandung. 1:1,000,000 scale map. Ewart, A. 1988. Geological history of the Fiji-Tonga-Samoan region of the S.W. Pacific, and some paleogeographic and biogeographic implications. Pp. 15–23 in Duffels, J.P. (ed.) The Cicadas of the Fiji, Samoa and Tonga Islands, Their Taxonomy and Biogeography (Homoptera, Cicadoidea). Entomonograph 10. Findlay, R.H. 2003. Collision tectonics of northern Papua New Guinea: key field relationships demand a new model. Pp. 291–307 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Gaina, C., R.D. Mu¨ller, B.J. Brown, and T. Ishihara. 2003. Microcontinent formation around Australia. Pp. 405–416 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Gaina, C., R.D. Mu¨ller, J.-Y. Royer, and P. Symonds. 1999. Evolution of the Louisiade triple junction. Journal of Geophysical Research 104: 12927–12939. Geist, E.L., T.L. Vallier, and D.W. Scholl. 1994. Origin, transport, and emplacement of an exotic island-arc terrane exposed in eastern Kamchatka, Russia. Geological Society of America Bulletin 106: 1182–1194. Giddings, J., C. Klootwijk, W. Sunata, C. Loxton, C. Pigram, and H. Davies. 1985. Paleomagnetism of Australia’s active northern margin in New Guinea. Abstracts, Third Circum-Pacific Terranes Conference, Sydney. Hall, R. 1997. Cenozoic tectonics of SE Asia and Australia. In Howes, J.V.C., and R.A. Noble (eds.) Proceedings of the IPA, Petroleum Systems of S. E. Asia and Australasia. Indonesian Petroleum Association Conference, Jakarta. Hall, R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences 20 (4): 353–431. Hall, R., and G.J. Nichols. 1991. Terrane amalgamation in the Philippine Sea margin. Tectonophysics 181: 207–222. Hall, R., and W. Spakman. 2003. Mantle structure and tectonic evolution of the region north and east of Australia. Pp. 361–381 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Hamilton, W.B. 1979. Tectonics of the Indonesian region. U.S. Geol. Survey Professional Paper 1078. U.S. Government Printing Office, Washington, D.C. Hamilton, W.B. 1988. Plate tectonics and island arcs. Geological Society of America Bulletin 100: 1503–1527. Hill, K.C., and R. Hall. 2003. Mesozoic–Cenozoic evolution of Australia’s New Guinea margin in a west Pacific context. Pp. 265–290 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Hill, K.C., and A. Raza. 1999. Arc-continent collision in Papua New Guinea: constraints from fission track thermochronology. Tectonics 18: 950–966. Karig, D.E. 1971. Structural history of the Mariana Island arc system. Geological Society of America Bulletin 82: 323–344. Karig, D.E. 1972. Remnant arcs. Geological Society of America Bulletin 83: 1057–1068. Kroenke, L.W. 1984. Cenozoic development of the Southwest Pacific. United Nations

................. 16157$

$CH5

02-08-07 10:25:26

PS

PAGE 162

Tectonic Geology of Papua / 163 Economic and Social Commission for Asia and the Pacific, Committee for Coordination of Joint Prospecting for Mineral Resources in South Pacific Offshore Areas, Technical Bulletin 6. Ling, H.Y., R. Hall, and G.J. Nichols. 1991. Early Eocene radiolaria from Waigeo Island, eastern Indonesia. Journal of Southeast Asian Earth Sciences 6: 299–305. Malaihollo, J.F.A., and R. Hall. 1996. The geology and tectonic evolution of the Bacan region, east Indonesia. In Hall, R., and D. Blundell (eds.) Tectonic Evolution of Southeast Asia. Geological Society of London Special Publication 106: 483–497. McCaffrey, R. 1996. Slip partitioning at convergent plate boundaries of SE Asia. In Hall, R., and D. Blundell (eds.) Tectonic Evolution of Southeast Asia. Geological Society of London Special Publication 106: 3–18. Milsom, J. 1991. Gravity measurements and terrane tectonics in the New Guinea region. Journal of Southeast Asian Earth Sciences 6: 319–328. Monnier, C., J. Girardeau, M. Pubellier, and H. Permana. 2000. L’ophiolite de la chaıˆne centrale d’Irian Jaya (Indone´sie): e´vidence pe´trologiques et ge´ochemiques pour une origine dans un bassin arrie`re-arc. Comptes Rendus de l’Acade´mie des Sciences Paris 331: 691–699. Monnier, C., J. Girardeau, M. Pubellier, M. Polve, H. Permana, and H. Bellon. 1999. Petrology and geochemistry of the Cyclops ophiolites (Irian Jaya, East Indonesia): consequences for the Cainozoic evolution of the north Australian margin. Mineralogy and Petrology 65: 1–28. Nash, C.R., G. Artmont, M.L. Gillan, D. Lennie, G. O’Connor, and K.R. Parris. 1993. Structure of the Irian Jaya mobile belt, Irian Jaya, Indonesia. Tectonics 12: 519–535. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. Bureau of Mineral Resources, Journal of Australian Geology and Geophysics 10: 193–211. Pigram, C.J., and P.A. Symonds. 1991. A review of the timing of the major tectonic events in the New Guinea orogen. Journal of Southeast Asian Earth Sciences 6: 307–318. Plafker, G., W.J. Nokelberg, and J.S. Lull. 1989. Bedrock geology and tectonic evolution of the Wrangellia, Peninsular and Chugach Terranes along the Trans-Alaska Crustal Transect in the Chugach Mountains and southern Copper River Basin, Alaska. Journal of Geophysical Research 94: 4255–4295. Polhemus, D.A. 1996. Island arcs, and their influence on Indo-Pacific biogeography. Pp. 51–66 in Keast, A., and S. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Publishing, Amsterdam. Polhemus, D.A., and J.T. Polhemus. 1998. Assembling New Guinea: 40 million years of island arc accretion as indicated by the distributions of aquatic Heteroptera (Insecta). Pp. 327–340 in Hall, R., and Holloway, J.D. (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Polhemus, J.T., and D.A. Polhemus. 2002. The Trepobatinae (Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 6. Phylogeny, biogeography, world checklist, bibliography, and final taxonomic addenda. Insect Systematics and Evolution 33: 253–290. Rogerson, R.J., and D.B. Hilyard. 1990. Scrapland: a suspect composite terrane in Papua New Guinea. In Carman, G.J., and Z. Carman (eds.) Petroleum Exploration in Papua New Guinea: Proceedings of the First PNG Petroleum Convention, Port Moresby, 12–14th February 1990. PNG Chamber of Mines and Petroleum, Papua New Guinea. Srolias, M., R.D. Mu¨ller, and C. Gaina. 2003. Tectonic evolution of the southwest Pacific using constraints from back-arc basins. Pp. 343–359 in Hillis, R.R., and R.D. Mu¨ller

................. 16157$

$CH5

02-08-07 10:25:26

PS

PAGE 163

164 / d a n a . p o l h e m u s (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Van Thournout, F., J. Hertogen, and L. Quevedo. 1992. Allochthonous terranes in northwestern Ecuador. Tectonophysics 205: 205–221. Veevers, J.J. 2000a. Billion-year Earth History of Australia and Neighbors in Gondwanaland. GEMOC Press, Sydney. Veevers, J.J. 2000b. Change in tectono-stratigraphic regime in the Australian Plate during the 99 Ma (mid-Cretaceous) and 43 Ma (mid-Eocene) swerves of the Pacific. Geology 28: 47–50. Visser, W.A., and J.J. Hermes. 1962. Geological results of the exploration for oil in Netherlands New Guinea. Konikl. Nederlands Geol., Mijnbouw Genoot., Verh. Geol Ser. 20: 1–265. Vroon, P.Z., M.J. Van Bergen, and E.J. Forde. 1996. Pb and Nd isotope constraints on the provenance of tectonically dispersed continental fragments in east Indonesia. In Hall, R., and D. Blundell (eds.) Tectonic Evolution of Southeast Asia. Geological Society of London Special Pub. 106: 445–453. Weiland, R.J., and M. Cloos. 1996. Pliocene-Pleistocene asymmetric unroofing of the Irian Fold Belt, Irian Jaya, Indonesia: Apatite fission-track thermochronology. Geological Society of America Bulletin 108: 1438–1449. Wright, J.E., and S.J. Wyld. 1994. The Rattlesnake Creek terrane, Klamath Mountains, California: an early Mesozoic volcanic arc and its basement of tectonically disrupted oceanic crust. Geological Society of America Bulletin 106: 1033–1056. Yan, C.Y., and L.W. Kroenke. 1994. A plate tectonic reconstruction of the Southwest Pacific, 0–100 Ma (CD-ROM). Proceedings of the Ocean Drilling Program, Scientific Research 130, Chap. 43.

................. 16157$

$CH5

02-08-07 10:25:26

PS

PAGE 164

2.2. Soils of Papua geoffrey s. hope and alfred e. hartemink o i l i s t h e i n t e rf a c e between the biosphere, atmosphere, hydrosphere, and lithosphere. It is a complex medium and has biological, chemical, and physical properties. Soils support plant growth by providing nutrients, water, and anchorage. Soils are also highly diverse ecosystems with a complex but little understood biota of fungi, bacteria, arthropods, and annelids that live below the surface but may contain more biomass than the above-ground organisms. The soil stores and respires large amounts of carbon and is an important source and sink of greenhouse gases. There are five major soil-forming factors: climate, vegetation, relief, parent material, and time. Differences in the combination of these factors result in soils varying over relatively short distances. Most soils are derived from the weathering of the underlying rock (parent material). Such weathering is at its most rapid in the humid tropics and may result in very deep soils (⬎10 m). However, many soils owe part of their characteristics to continuing accessions of alluvium, windborne dust (loess), and organic matter (peat soils), colluvium, or volcanic debris. Thus parent material is often from multiple sources. Soils are always changing and will often not have reached a point of balance or maturity. Thus soil profiles (the description of a soil in cross-section) differ markedly between young and old soils. Soils are also significantly influenced by human activities, for example, by tillage, drainage, fertilization, mounding, or soil erosion. Soils are dynamic, with changes due to influx of sediment and decayed rock and removal of material by leaching and erosion in a changing climate. Hence the imprint of one soil over an older one (a palaeosol) is not uncommon. The range of rock types, climate, and relief in Papua provides a wide range of soil types. This brief chapter gives an overview of the main soils of Papua and is based on surveys and older work such as a soil map of New Guinea by Haantjens et al. (1967) with extrapolations from more intensively studied agricultural soils of Papua New Guinea (Wood 1982; Bleekers 1983; Hanson et al. 2001). Detailed land system and soil mapping has been undertaken in connection with transmigration (transmigrasi) settlements in Papua but unfortunately those data are not available publicly. Soils of Papua are somewhat similar to those occurring in western Papua New Guinea because of their shared geology. However, the soils of western New Guinea, including Papua, are less diverse than those of eastern New Guinea because Quaternary volcanic activity is not present.

S

Soil Classification Knowledge on the distribution of soils and their properties is essential for managing agricultural production and other soil services. In Papua, most villagers have Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

165

................. 16157$

$CH6

02-08-07 10:22:52

PS

PAGE 165

166 / geoffrey s. hope & a lfred e . hartemink

names for their soils and they are knowledgeable about soil properties. In textbooks and publications on soils in Papua New Guinea, the Soil Taxonomy classification scheme of the United States Department of Agriculture (USDA) is used (Hanson et al. 2001). Soil taxonomy uses measurable properties of a soil, including soil depth, color, texture (e.g., sand, sandy loam, loam, clay), structure (e.g., blocky, granular, crumb, columnar), consistency (e.g., sticky, porous), soil water and soil chemical properties such as pH (a measure of acidity), and the ability to retain nutrients. These properties are used to group soils into classes. A full classification, under the Soil Taxonomy, requires chemical and physical analyses (Soil Survey Staff 1998). Papua New Guinea uses the Soil Taxonomy because of the close links to the development of a comprehensive Resource Information System (PNGRIS) in PNG. The Soil Taxonomy is internationally recognized and should be applied to Papua as it has advantages when exchanging information. Some problems occur when the Soil Taxonomy is used to classify New Guinea soils because that taxonomy relies heavily on laboratory analysis, which is expensive. Many descriptions and analyses of Papuan soils do not have sufficient analytical data to allow a full Soil Taxonomy classification. In addition, soil moisture and soil temperature information is needed in the Soil Taxonomy, but only limited data are available for Papuan soils. In soil taxonomy, the highest level is the soil order, usually made up of several soil groups that reflect similar formational processes but different hydrological regimes or parent materials. The orders are usually arranged from least developed to most developed, as evidenced by horizonation, color (oxidation), and leaching. While there is a general correlation with time, some parent materials and topographic settings accelerate the development of weathering whereas cooler and drier conditions may hinder weathering. For example an iron-rich rock such as basalt on a mid-slope position where abundant water is running through the profile but which also has good aeration will weather to a bright red color. It may follow a path to a highly weathered soil more quickly than other rocks. The following soil orders occur in Papua with the more important soil groups indicated in italics. The descriptions are based on Bleekers (1983). The common terminology applied in Southeast Asia is from the Food and Agricultural Organization (FAO). The FAO term (Dudal 2005) is given in parentheses after the following descriptions according to the Soil Taxonomy, but cross-categories are common, as are a plethora of earlier terms, many adopted from European usage. Entisols are very young soils, with little or no profile except for a thin humic surface horizon. These soils occur mainly on recent alluvium or on steep slopes where soil erosion takes place, or on coastal deposits. Examples include some acidic mangrove clay soils, sulfaquents (cat clays) that are continuously waterlogged, waterlogged fluvial silt-clays with reducing conditions (aquents), silt-clay alluvium (fluvent), and riverine or beach sands (psamments). (FAO: gleysols, fluvisols, arenosols)

................. 16157$

$CH6

02-08-07 10:22:53

PS

PAGE 166

Soils of Papua / 167

Histosols are soils that contain very high levels of organic matter (peat soils). These soils are mostly dark brown to black in color, and occur in swampy areas. They are saturated with water for much of the year. Histosols are divided by the degree of breakdown of the plant material into fibrist, hemist, and saprist groups in waterlogged situations and folist on drained slopes. In some cases, these materials have been actively accumulating over thousands of years and can also be regarded as biogenic sediments. (FAO: Histosols, peats) Inceptisols are moderately weathered soils, with slightly developed horizons. Tropaquepts form on rock and alluvium on ridge crests and slight hollows under grassland, and are acidic. Humitropepts are deep clay soils formed under cool humid conditions with a deep organic rich upper layer. Eutropepts and dystropepts have a humic horizon over reddish or mottled lower layer that may contain stones, and are found on slopes or alluvium up to an altitude of 1,500 m. In alpine conditions above 3,500 m, cryochrepts are shallow alpine humus or skeletal soils with black peaty layer over a brown clayey layer. (FAO: cambisols, cryosols). Inceptisols also include a sub-order, andosols, which are productive soils derived from volcanic ash. These are absent in Papua, though very common in Papua New Guinea (New Britain) and some Moluccan islands such as Halmahera and Ternate. Vertisols are soils with high montmorillonite clay content that are sticky when wet and very hard when dry. These soils swell when wet and crack when dry but are generally of high fertility. They occur in seasonally wet depressions in drier regions near Merauke. (FAO: vertisols) Mollisols are soils in which there is accumulation and decomposition of organic matter. Biological activity (worms, ants, termites, roots, etc.) is high in these soils and contributes to soil turnover and weak horizonation. These soils generally have a high base (e.g., calcium, magnesium) content. Rendolls are shallow black soils common on limestones. Ustolls are found on fine grained rocks in savanna areas with a pronounced dry season while udolls are found in humid areas up to 1,500 m. They have a thick grayish or brown top horizon grading into a clayey brown lower horizon of limited permeability. (FAO: phaeozems) Alfisols are moderately weathered soils that have an argillic horizon (a layer within the soil profile with higher clay content due to movement of clay from the top to lower layers). These soils are sometimes very fertile. Variants (aqualfs, ustalfs, udalfs) occupy a range of waterlogged through seasonal to humid settings where the parent material has had a long time for horizonation to develop. Some contain plinthite, an iron-rich clay indicating poor drainage, which forms concretions if dried out. They include rhodudalfs, reddish clay soils formed on limestone, particularly raised reefs at low altitudes. The most common soil is the tropudalf, a deep brownish soil found on slopes on sedimentary rocks. (FAO: luvisols) Ultisols are strongly weathered and acid soils with an argillic horizon found in wet climates up to 3,000 m. These soils have a low base saturation. Plinthaquults occur on flat terrain around Merauke and have gray sandy-clay upper horizon over deep gray or brown mottled plastic clays. They are infertile and, once dried, form concretions (Schroo 1964). Tropohumults have high organic content

................. 16157$

$CH6

02-08-07 10:22:53

PS

PAGE 167

168 / geoffrey s. hope & a lfred e . hartemink

throughout the profile which is yellow to brown at the base and acid, and occur on rugged terrain on sedimentary and igneous rocks. Tropudults occur on gentle slopes below 1,000 m and have a red or yellow mottled horizon grading down to clay. They are acid and of low fertility. (FAO: acrisols, alisols, lixisols, plinthosols, podzols) Oxisols are very strongly weathered soils, with low fertility, which are unusual in Papua. These soils reflect tropical weathering of very long duration so they occur on old land surfaces with profiles several meters in depth. The upper profile has lost much of the clay and other silicate minerals through leaching, retaining limonite and sometimes bauxite or kaolinite. On ultramafic rocks they are often enriched in nickel, iron, and cobalt oxides. (FAO: ferralsols)

The Distribution of Soils in Papua The most common soils in Papua are entisols. These young soils cover more than a quarter of the total land area, reflecting the high geological and erosional activity. Entisols are common in the hills and foothills where alluvial deposits occur (Figure 2.2.1). In the highland basins, entisols occur on alluvial fans and are also found on sand sheets in sandstone country, such as north of Mt Trikora, and on alpine gravels (Figure 2.2.2). Inceptisols are common on more stable settings in montane areas and often have deep organic rich horizons overlaying clayey subsoils or shallow humic soils on limestone (Figure 2.2.3). Mollisols are deeper soils formed on limestone and are widespread in limestone terrains in the Central Range, Weyland, and Arfak mountains, as well as parts of Biak and Numfoor islands. Erosion occurs

Figure 2.2.1. Entisols on riverine silts, Baliem Valley near Wamena. These are formed on overbank alluvium and slopewash.

................. 16157$

$CH6

02-08-07 10:22:57

PS

PAGE 168

Soils of Papua / 169

Figure 2.2.2. Entisols at 4,500 m on Mt Jaya. The vegetated area on left has acid peats on old moraine (a cryochrept) while a thin litter layer has built up on fresh limestone rubble left by a glacier about 1880 C.E. in this area. Soil loss of a meter or more (Figure 2.2.4) can be readily seen on gardened limestone slopes because the boulders are etched up to the level of the soil that existed prior to clearance. This lowering may represent thousands of years of gardening. Alfisols are common in the northern slopes of the Central Range (Figure 2.2.5). Ultisols (strongly weathered soils) cover approximately 25% of the land area of Papua, and are dominant in the Merauke Regency, where they occupy more than half of the lowland area. They also are found on ultramafic rocks such as in the Cyclops Mts (Figure 2.2.6). Histosols are most prevalent in the Taritatu (Idenberg)-Mamberamo basin (Figure 2.2.7), around McCluer Gulf and the southern plain, where they occur on a third of the land area. They are also common above 3,000 m in the mountains (Bleeker 1980). Most areas are a mosaic of soil types often reflecting erosional history and gradients in slope processes, often giving rise to a soil succession (or catena). Palaeosols are also common (Figure 2.2.8). Thus soil mapping needs to be detailed. Table 2.2.1 estimates the importance of the Great Soil Groups in Papua and compares them to the figures derived from land system mapping in Papua New Guinea. Inceptisols are of less importance in Papua but histosols and oxisols, although still minor, occur more widely, reflecting the older landscapes and tectonic basins of the province.

................. 16157$

$CH6

02-08-07 10:23:25

PS

PAGE 169

170 / geoffrey s. hope & a lfred e . hartemink

Figure 2.2.3. Inceptisol and rock slope in the Daelah Valley west of Wamena. The downslope drainage ditches are designed to reduce soil moisture and the soil is coherent enough not to gully.

Soil Moisture The water balance (precipitation over evapotranspiration) of Papua is positive except in the Merauke Regency so that soil moisture rarely limits plant growth. Even in the driest stations soil moisture does not fall below 75 mm (50% of maximum storage) as was estimated from Western Province data from McAlpine and Keig (1983). In higher altitudes with more clouds, evaporation is low and soils are water-saturated during much of the year. These positive water balances result in reduced decomposition (and thus accumulation) of organic matter and possibly leaching of nutrients, together with landsliding, tunneling, and piping on steep sites and at-risk erodable soil types. However, drought occurs in both the highlands and lowlands with free drainage during long periods without rain, such as those experienced during the 1997–1998 El Nin˜o (Ballard 2000).

Soil Fertility In Papua, the highest density of settlement occurs in the highlands where entisols derived from alluvium are very important, together with alfisols and mollisols on

................. 16157$

$CH6

02-08-07 10:23:51

PS PAGE 170

Soils of Papua / 171

Figure 2.2.4. Mollisols eroding from gardened limestones, Baliem Valley near Tiom. Considerable soil loss has followed forest clearance here, and the area may soon be unusable. limestone. These soils are generally fertile. In contrast, 56% of the population of Papua New Guinea, almost three million people, live on inceptisols, many derived from volcanic ash, or mixed with volcanic ash from explosive eruptions that did not reach Papua. To compensate for the poor soil fertility, Papuans in the highlands have developed a system of returning organic matter to the soil to add nutrients and maintain favorable soil physical properties (Ploeg 2005; Chapter 6.3). Around 30% of the population in the seasonally dry region around Merauke cultivate ultisols. Attempts to develop these soils for rice agriculture have met great difficulties as large inputs of fertilizer and machinery are needed. The availability of nutrients for plants depends on several factors. Low levels of available nutrients in the soil may be natural, caused by low amounts of nutrients in the parent material from which the soil is derived; by fixation and immobilization of nutrients; or by losses, for instance, when high rainfall leaches nutrients from the soil. Nutrient imbalances (e.g., high calcium, low potassium) in the soil may also cause limited availability of a particular nutrient. Low nutrient levels may also result from cultivation, when agricultural crops and repeated burning

................. 16157$

$CH6

02-08-07 10:24:17

PS PAGE 171

172 / geoffrey s. hope & a lfred e . hartemink

Figure 2.2.5. Alfisol at Kwiyawagi, East Baliem Valley. The landforms are mature and more than 100,000 years old. remove nutrients that are not replaced by manure or inorganic fertilizers (Yaku and Widyastuti 2005). Nutrient deficiencies affect crop production in Papua. This problem is likely to increase in the future because of more intensive land use as the population rises, which results in reduced fallow periods in shifting cultivation system and higher land use intensities (Hartemink and Bourke 2000). Soil nitrogen availability is determined in part by the length and type of fallow, the introduction of organic matter (plant materials) into the soil, and climate (temperature and rainfall). Most plant-available nitrogen in Papuan soils is derived from organic matter. Soil nitrogen tends to be higher in soils of the highlands, where temperatures are lower and organic matter decomposes more slowly. In New Guinea soils the availability of phosphorus is dependent mostly on the organic matter content. A small part comes from the weathering of parent material or secondary minerals. Phosphorus is usually found in combination with calcium, magnesium, iron, and aluminum. Although relatively large amounts of total phosphorus may be present in the soil, little may be available to plants because phosphorus is held very tightly in the soil in organo-clay complexes. Phosphorus fixation can be severe in ultisols and oxisols and in soils derived from volcanic ejecta. The availability of soil potassium is related to rock type and the mineralogy and the stage of weathering of the soil. Potassium-deficient soils are usually highly

................. 16157$

$CH6

02-08-07 10:24:45

PS

PAGE 172

Soils of Papua / 173

Figure 2.2.6. Ultisol or oxisols on the Cyclops Mts west of Sentani, developed on ultramafic rocks. The clearing is very infertile and forest does not readily re-invade. In some places internal drainage has developed by solution of the rocks.

Figure 2.2.7. The Mamberamo Lakeplain, a tectonic basin crossed by the Tariku and Taritatu Rivers, is a complex of alluvial entisols and widespread histosols under swamp forest and floodplain grassland.

................. 16157$

$CH6

02-08-07 10:25:32

PS

PAGE 173

174 / geoffrey s. hope & a lfred e . hartemink

Figure 2.2.8. Slope profile of an inceptisol over a palaeosol at Supulah Hill, a sandstone hill north of Wamena. The humic layer has built up on sands that eroded and covered an earlier humic horizon about 30,000 years ago. weathered and leached with limited amounts of mineral reserves. Soils that develop on limestone and that have high levels of calcium and magnesium may have a potassium deficiency (nutrient imbalances). This is common, for example, on soils developed on Pleistocene coral terraces on Biak Island. Research on nutrient deficiencies of agricultural crops started in the 1950s, but only limited research is currently being carried out on soil nutrient deficiencies or on soil fertility management strategies in Papua (Yaku and Widyastuti 2005). Soil fertility problems exist in parts of the island. Further intensification of land use affects soil fertility, and nutrient deficiencies are therefore likely to increase, particularly in food crops where inorganic fertilizers are not being used. There is a need to monitor the development of nutrient deficiencies as well as to properly identify the deficiencies through trials and soil and foliar analysis. Papua differs from much of the rest of Indonesia in having low-fertility soils. This may be part of the reason why there seems to be more marked patterning in the forests (greater species diversity per unit area) compared to the monodominant stands of west Malesia. The soils may thus be fragile and prove difficult to return to forest once logged. Thus specific research into soil structure and nutrition within the province remains essential, as do active efforts to retain the soils already there.

................. 16157$

$CH6

02-08-07 10:25:58

PS

PAGE 174

................. 16157$

$CH6

02-08-07 10:25:58

PS

PAGE 175

A 120 26

PNG km x 1000

PNG % of area

2

8.6

C

C

D

B

A

B

D

B

D

Histosols

Inceptisols

48

219

A

A

C

C

D

B

B

B

B

Vertisols

Mollisols

34 7

⬍0.1

C

C

C

C

X

B

B

C

D

0.2

D

X

X

X

X

X

X

X

D

Alfisols

3

13.3

X

D

C

B

D

B

B

B

X

Ultisols

14

63

B

C

A

B

X

B

B

A

A

Oxisols

⬍0.1

0.007

C

C

D

X

X

D

X

X

X

Note: A: ⬎25%; B: 10–25%; C: 2–10%; D: ⬍2%; X: absent. The two PNG values for soil orders give the estimated area in thousands of km2 and the percentage land cover in PNG. B. J. Allen provided these unpublished data.

2

Vogelkop and Islands

B A

Coastal

Northern Ranges

Alpine and Subalpine A

C

Highland Valleys

Northern Trough

A B

Southern Slopes

A

Southern Lowlands

Entisols A

Merauke

Region

Table 2.2.1. Major soil orders distribution by physiographic area of Papua, compared with PNG

176 / geoffrey s. hope & a lfred e . hartemink

Literature Cited Ballard, C. 2000. Condemned to repeat history? ENSO-related drought and famine in Irian Jaya, Indonesia. Pp. 123–148 in Grove, R., and J.M.A. Chappell (eds.) El Nin˜o—History and Crisis. White Horse Press, Cambridge. Bleeker, P. 1980. The alpine soils of the New Guinea high mountains. Pp. 59–74 in van Royen, P. (ed.) Alpine Flora of New Guinea. Cramer-Verlag, Vaduz. Bleeker, P. 1983. Soils of Papua New Guinea. Australian National University Press, Canberra. Bourke, R.M. 1983. Crop micronutrient deficiencies in Papua New Guinea. Department of Primary Industry Technical Report 83/3. Department of Primary Industry, Port Moresby. Dearden, P.N., D.F. Freyne, and G.S. Humphreys. 1986. Soil and land resource surveys in Papua New Guinea. Soil Survey and Land Evaluation 6 (2): 43–50. Dudal, R. 2005. Soils of Southeast Asia. Pp. 94–104 in Gupta, A. (ed.) The Physical Geography of Southeast Asia. Oxford University Press, Oxford. Haantjens, H.A., J.J. Reijnders, W.L.P.J. Mouthaan, and F.A. van Baren. 1967. Major soil groups of New Guinea and their distribution. Communication No. 55, Royal Tropical Institute, Amsterdam. Hanson, L.W, B.J. Allen, R.M. Bourke, and T.J. McCarthy. 2001. Papua New Guinea Rural Development Handbook. Land Management Group, Australian National University, Canberra. Hartemink, A.E., and R.M. Bourke. 2000. Nutrient deficiencies of agricultural crops in Papua New Guinea. Outlook on Agriculture 29: 97–108 Hope, G.S. 1976. Vegetation. Pp. 113–172 in Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.) The Equatorial Glaciers of New Guinea. A.A. Balkema, Rotterdam. McAlpine, J.R., G. Keig, and R. Falls. 1983. Climate of Papua New Guinea. CSIRO/ Australian National University Press, Canberra. Ploeg, A. 2005. Sweet potato in the central highlands of west New Guinea. Pp. 149–161 in Ballard, C., P. Brown, R.M. Bourke, and T. Harwood (eds.) The Sweet Potato in Oceania: A Reappraisal. Oceania Monograph 56, Sydney. Schroo, H. 1964. An inventory of soils and soil suitabilities in West Irian: IIB. Netherlands Journal of Agricultural Science 12: 1–26. Soil Survey Staff. 1998. Soil taxonomy: a basic system of soil classification for making and interpreting soil surveys. Natural Resources Conservation Service, Agricultural Handbook 436. U.S. Department of Agriculture, Washington, D.C. Wood, A.W. 1982. The soils of New Guinea. Pp. 73–83 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Monographiae Biologicae 42, Dr. W. Junk Publishers, The Hague. Yaku, A., and C.A. Widyastuti. 2005. Sweet potato research and development in Papua, Indonesia: a review. Pp. 163–170 in Ballard, C., P. Brown, R.M. Bourke, and T. Harwood. The Sweet Potato in Oceania: A Reappraisal. Oceania Monograph 56, Sydney.

................. 16157$

$CH6

02-08-07 10:25:59

PS

PAGE 176

2.3. Climate of Papua michael l. prentice and geoffrey s. hope Climatic Setting of Papua a p ua i s t he w e s te r n h al f of an equatorial island that is the northern extension of the Australian continental plate, together forming a barrier that blocks the flow of surface water from the western Pacific to the Indian Ocean. As a result, surface waters transported across the Pacific and so the warmest on the planet pile up in the western Pacific north of Papua as the vast Western Pacific Warm Pool (WPWP) (Figure 2.3.1). The WPWP is the single largest heat source to the global atmospheric circulation on the planet. To the south of Papua is the shallow epicontinental Arafura Sea, which permits only slight water transfer between the oceans and also has high surface temperatures. Although no point in Papua is more than 250 km from the sea, the island is effectively divided in two by a ESE-WNW trending mountain cordillera that exceeds 3,500 m above sea level (asl) in Papua and reaches 5,000 m asl in the highest peak, Mt Jaya. This pattern is repeated in the west by a lower chain of mountains in the Vogelkop peninsula.

P

the atmosphere and seasonal changes The large-scale atmospheric and oceanic controls on the climate of Papua are discussed by MacAlpine et al. (1983), Nix and Kalma (1972), Hastenrath (1991),

Figure 2.3.1. Schematic cross-section of the zonal circulation on the equator during La Nin˜a (above) and El Nin˜o (below) phases of the El Nin˜o-Southern Oscillation. Source: after Webster and Lucas (1992).

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

177

................. 16157$

$CH7

02-08-07 10:25:28

PS

PAGE 177

178 / michael l . prentice & g eoffrey s. hope

and Philander (1990). Three major circulation systems control the weather of Papua. The meridional Hadley Circulation consists of equatorward air flow at the surface, rising air over the warmest sea-surface temperatures (SST), poleward flow in the free troposphere, and subsidence on the subtropical high-pressure centers. The zonal Walker Circulation is also thermally driven, but by the contrast in SST across the tropical Pacific (Figure 2.3.1). In the western Pacific, these circulations are abstractions, as ascents related to the Hadley and Walker Circulations are indistinguishable. Less well known are the oblique, large-scale circulation systems associated with the semi-permanent troughs in the mid-latitude westerlies, the socalled polar troughs. On the western sides of these troughs, surface air is steered equatorward while, on the eastern sides, equatorial air is driven poleward. These three circulation systems generate two important zones of surface air convergence. The northern and southern hemisphere cells of the Hadley Circulation meet at the Intertropical Convergence Zone (ITCZ), where deep convection is fueled by low-level convergence of moist air. Although narrow in the eastern Pacific, the ITCZ or equatorial trough is up to 1,200 km wide in the New Guinea region. The ITCZ migrates ⬃15 north and south over New Guinea annually following the warmest surface waters. The ITCZ is bordered by northern and southern monsoon shear lines (N/SMSL). The trough in the mid-latitude westerlies that most affects Papua is the South Pacific Convergence Zone (SPCZ). The surface wind systems affecting Papua are the trade winds, southeast and northeast, and the equatorial or monsoon westerlies. Depending on the direction of the surface pressure gradient between the two monsoon shear lines and their location with respect to the equator, trade winds over Papua are either southeasterly, northeasterly, or northwesterly and of variable depth. During the southernhemisphere winter, the SMSL shifts northward to coincide with northern New Guinea (equator), and the principal ITCZ convergence is to the north along the NMSL. The result is that New Guinea is completely embedded in the southeast trades, hence the southeast season. During southern summer, the NMSL shifts south to northern New Guinea and the principal ITCZ convergence is along the SMSL. In this case, the northeast trades veer at the equator and become the monsoon westerlies over New Guinea (the northwest season). In the transitional seasons, depending on the direction and strength of the surface pressure gradient in the equatorial trough, the predominant winds commonly differ between the northern and southern New Guinea lowlands, with the boundary between the two air masses in the highlands. The trade winds and monsoon westerlies are contained within the Atmospheric Boundary Layer (ABL), which is capped by a strong temperature inversion that is at ⬃2,000 m asl but disappears where convection breaks out during daytime along the ITCZ (Figure 2.3.2). This inversion marks the upper limit of the ABL cloud layer. Given their 2,000 m stable upper boundary, trade and monsoon winds are commonly blocked by the high Central Range, with the effect that highland New Guinea climates are dominated by local convection. During daytime, mountain heating generates intense convection and local circulations throughout the year.

................. 16157$

$CH7

02-08-07 10:25:28

PS

PAGE 178

Climate of Papua / 179

Figure 2.3.2. A cross section of major climate elements. Source: after MacAlpine, Keig, and Falls (1986).

the ocean and seasonal changes The WPWP exhibits temperatures above 27.5C, variable salinity (increasing downward), and an average depth of 150 m. It is effectively separated from the cold ocean by the thermocline, a relatively sharp cooling with depth (Figure 2.3.3). In the eastern tropical Pacific, the surface mixed-layer is cooler, on average 25C,

Figure 2.3.3. Changes in temperature and salinity with depth in the western Pacific. The profiles are from Cenderawasih Bay, a large bay on the north coast of Papua. Two soundings, CTD 6, at the mouth of the bay, and CTD 9, in the center of the bay, are typical.

................. 16157$

$CH7

02-08-07 10:25:50

PS

PAGE 179

180 / michael l . prentice & g eoffrey s. hope

with significant seasonal variability, and thinner, with an average thermocline depth of ⬍50 m (Figure 2.3.1). The lower SST reflects wind-driven upwelling of cool thermocline water along the equator and also deep cold water along the coast of South America. As a result, eastern Pacific surface waters form a cold tongue that narrows westward. The eastwest contrast in Pacific SST is largely responsible for the atmospheric pressure gradient across the Pacific that drives the Walker Circulation (Figure 2.3.1). SST in the central and eastern Pacific is also asymmetrical with respect to the ITCZ or thermal equator, which follows the zone of maximum SST between 3 and 10N. The Walker Circulation drives the westward-flowing and strong South Equatorial Current (SEC) and the weaker North Equatorial Current (NEC) across the tropical Pacific toward Papua (Figure 2.3.4). Between these two is the North Equatorial Counter Current (NECC), which flows eastward under the influence of equatorial westerlies. Because of the trade winds, the depth of the thermocline decreases from west to east across the tropical Pacific, as does the height of sea level (⬃50 cm). The zonal slope in the thermocline establishes a pressure gradient in the upper ocean that is directed from west to east and drives a strong subsurface current eastward along the equator within the ocean thermocline, the Equatorial Undercurrent (EUC).

Figure 2.3.4. Climatological schematic of western tropical Pacific currents. SEC: South Equatorial Current; NEC: North Equatorial Current; NECC: North Equatorial Counter Current (below 200 m); NGCC: New Guinea Coastal Current; NGCUC: New Guinea Coastal Undercurrent; MUC: Mindanao Undercurrent; MC: Mindanao Current; ME: Mindanao Eddy; HE: Halmahera Eddy; EUC: Equatorial Undercurrent.

................. 16157$

$CH7

02-08-07 10:26:04

PS

PAGE 180

Climate of Papua / 181

The surface current along the north coast of New Guinea, the New Guinea Coastal Current (NGCC), is strongly influenced by the semi-annual reversal in wind direction (Barmawidjaja et al. 1993; Wyrtki 1961). The NGCC flows to the west during the southern winter, the southeast season, and reverses during the northern winter. The NGCC joins the SEC around Halmahera and forms the Halmahera Eddy (HE) (Tomczak and Godfrey 1994; Lukas et al. 1996) (Figure 2.3.4). At a depth of about 200 meters, the New Guinea Coastal Undercurrent (NGCUC) flows to the west along the northern New Guinea coast and, after passing the equator, reverses direction and becomes the eastward-flowing EUC. In both hemispheres, surface waters flow away from the equator but recirculate back to it through the ocean thermocline and equatorial upwelling (Dietrich 1970). Along the northern coast of Papua in Cenderawasih Bay, the surface-mixed layer is 30C and features a salinity gradient from 33.3 psu (practical salinity units) at the northern end to 30.3–32.9 psu in the southern sector (Figure 2.3.3). The depth of this mixed layer is ⬃40 m in the southern bay and ⬃60 m in the north. The semi-restricted nature of the basin is clearly indicated by the oxygen content, which is low below 200 m depth compared to the open sea. The water mass below 150 m depth has hydrographic properties typical of western South Pacific central water. The upper water mass has lower salinity and reflects the positive water balance caused by high rainfall and river inflow. This water drains to the ocean and is replaced by cooler, saltier water crossing the 800 m deep sill of the bay. By contrast, the Arafura Sea has drying winds and a general circulation from east to west during southern winter. At this time, evaporation and clear skies cause high salinities and warming of the shallow water to temperatures over 32C toward the end of the year. The movement of the ITCZ over the region and development of the Australian Monsoon in December–March reverses the flow and SST falls. Papua is too equatorial to be affected by tropical cyclones (typhoons), which develop south of 6S, but associated rain can affect Papua. To the west of the Arafura shelf, upwelling of cool water from the Indian Ocean occurs off Kaimana, Misool, and the southwestern Vogelkop Peninsula. This leads to more seasonal and lower rainfall. The effect of the different water masses is evidenced by a dramatic vegetational change from a low monsoon forest, with winter deciduous elements on Misool Island, to tall evergreen rainforest on the northern coast of Salawati Island.

El Nin˜o–Southern Oscillation The Walker Circulation fluctuates in strength on interannual (3–7 year) timescales, resulting in oscillations in surface air pressure over the WPWP that are coherent but out of phase with surface pressure over the southeastern tropical Pacific (Philander 1990). This oscillation is known as the Southern Oscillation. When the circulation weakens, surface pressure in the eastern Pacific decreases while, in the western Pacific, surface pressure increases (Figure 2.3.1). The large convective zone over the WPWP shifts eastward, which reduces convection in the western Pacific, thereby causing surface pressure to increase. Fluctuations in the

................. 16157$

$CH7

02-08-07 10:26:05

PS

PAGE 181

182 / michael l . prentice & g eoffrey s. hope

Walker Circulation arise owing to two-way coupling with changes in tropical Pacific surface and near-surface waters. The relatively cool eastern tropical Pacific warms on an irregular, interannual period, and this causes the trades and, in fact, the entire Walker Circulation to slacken. This, in turn, permits the WPWP to drain eastward, further warming the eastern tropical Pacific Ocean and atmosphere and slowing the Walker Circulation. These processes lead to a basin-scale warm state referred to as El Nin˜o (Figure 2.3.1). When a sluggish wave in the depth of the ocean thermocline, launched westward when eastern Pacific SST was cool, returns, eastern Pacific SST starts to cool again, which increases the strength of the trades. This, in turn, causes the WPWP to retreat westward, increasing thermocline depth in the west. This relatively cool state of the tropical Pacific is referred to as La Nin˜a. The interannual fluctuations in the coupled Pacific ocean-atmosphere system are referred to as the El Nin˜o–Southern Oscillation or ENSO. Interestingly, ENSO arises because of internal processes without imposed external perturbations.

Radiation and Cloudiness Solar radiation at the top of the atmosphere is high all year round, but the high cloud cover reduces direct radiation at ground level. Areas with daily cycles of rain formation receive the bulk of their radiation in the morning. This leads to aspect differences that might not be expected given the high sun angle experienced through the year. West-facing slopes may receive significantly less radiation than east-facing ones, which can affect crop yields and altitudinal limits. Brookfield (1964) suggested that daily local circulations are responsible for agricultural success in the mountain valleys. Very high cloudiness and waterlogging hinder agriculture on the outer flanks of the mountains, but the intermontane valleys are drier and sunnier, supporting dense populations. Heating in the morning causes upslope winds to rise, creating cloud on the slopes. Where a valley is large enough, air descends in the center, giving clear skies and abundant sunlight (Figure 2.3.5). In the evening the process is reversed, as cold winds descend the slopes and air rises and supports evening shower activity in the valley center. Radiation at the top of the atmosphere varies by about 5–10% between November, when the sun is overhead, and July, when it is farthest away. However, cloudiness is lowest during southern winter and increases in the southern summer, so the radiation received at the ground varies less than at the top of the atmosphere. Values from Sentani, a relatively sunny location, suggest mean daily solar energy densities of 460–500 milliwatt hours/cm2 through the year (McAlpine et al. 1983). Maximum values for Merauke probably reach 625 milliwatt hours/cm2.

Temperature For New Guinea, meteorological observations, especially in the highlands, are sparse. Therefore, we regard the meteorological data collected at a transect of sites from sea level to 4,400 m asl on the south-facing slope of the Central Range at

................. 16157$

$CH7

02-08-07 10:26:05

PS PAGE 182

Climate of Papua / 183

Figure 2.3.5. Daytime circulation in a highland valley. The direction reverses at night. ⬃137 E and 4 S as representative of the southern half of Papua. This transect of sites is adjacent to the currently glaciated Mt Jaya massif and is hereafter referred to as the Mt Jaya transect. These stations show that there is a greater diurnal temperature range below 1,500 m asl than above it, particularly in January (Figure 2.3.6). For example, the difference between average January maximum and minimum temperatures is 8–10C below 700 m asl. However, above 2,500 m asl, the average January and July diurnal temperature difference is ⬃5C. These data also indicate that there is greater seasonality below 2,000 m asl than above it (Figure 2.3.6). This seems to be primarily due to a larger difference in average monthly maximum temperatures below 2,000 m asl than above. For instance, at 660 m asl, average January and July temperatures differ by 5C whereas, above 2,500 m asl, the difference is about 1C. January and July minimum temperatures differ by only 1C across all elevations. Atmospheric temperatures above Merauke recorded by radiosondes also exhibit increased seasonality below about 2,000 m asl (Figure 2.3.6). The decrease of temperature with altitude above 2,500 m asl on the Mt Jaya transect (surface lapse rate) averages 5.3C/km annually (Figure 2.3.6). This is about the same as the atmospheric lapse rate recorded by radiosondes at Merauke. Surface lapse rates below 2,500 m asl are higher, 7C/km, for maximum monthly temperatures reflecting relatively strong heating of air near the surface during daytime. Surface lapse rates for minimum temperatures are less than the annual average because of radiational cooling at night. Merauke radiosonde data show the same trends. Between 2,000 and 2,500 m asl, surface lapse rates for maximum monthly temperatures are relatively high. This feature likely reflects the average position of the top of the ABL. There is strong correspondence between the altitude and important biophysical boundaries. Megathermal lowland environments have temperatures of 25–35C

................. 16157$

$CH7

02-08-07 10:26:11

PS PAGE 183

184 / michael l . prentice & g eoffrey s. hope

Figure 2.3.6. Average maximum and minimum temperatures for January and July for surface stations on the Mt Jaya transect. Also shown are average January and July temperatures at standard pressure levels (height equivalent) above Merauke recorded by radiosondes launched at GMT(, o). Numbers are lapse rates in C/ km. throughout the year, with a modest diurnal range because of the high heat capacity of saturated air. At about 1,200–1,400 m, the mesothermal zone sees a reduction in tree leaf size and evapotranspiration is reduced. Above this zone, highland climates in intermontane valleys center around 18C but have a greater diurnal range, from 10–25C. Very rare frost can be experienced after periods of dry weather. Above 2,800 m asl, frost becomes more frequent, leading to microtherm climates and a change to subalpine forests with microphyll and nanophyll leaves. The treeline, at about 3,900–4,200 m asl, marks a mean annual isotherm of 6C and the tropic alpine zone (Barry 1978b, Hnatiuk et al. 1976). Daytime temperatures can reach 20C, but at night frosts of –2 to –5C are common. Because humidity remains high, the much colder nighttime temperatures of other tropical mountains (–20C) do not occur (Hastenrath 1991). Because of high snowfall, the snowline occurs at a mean annual temperature of about 1C, which was at around 4,650 m asl in 1972–73. Peaks above this altitude are in the nival zone, where plant life is reduced to snow algae (Peterson and Kol 1976). Snowfalls down to 3,800 m were common in 1971–73, but these melted rapidly, usually within a few hours.

................. 16157$

$CH7

02-08-07 10:26:21

PS

PAGE 184

Climate of Papua / 185

Humidity and Rainfall Water-vapor mixing ratios (MR) decrease with elevation on the Mt Jaya transect from 17–20 g/kg at sea level to about 6–10 g/kg at 4,400 m asl (Figure 2.3.7). The slope of the MR decrease with altitude on the surface transect increases significantly at ⬃2,000 m asl, which we interpret as reflecting the average elevation of the top of the ABL. This is consistent with water-vapor content in the atmosphere above Merauke, which shows a similar increase in slope between 1,500 and 3,000 m asl. On average, there is little seasonality in water-vapor content on the Mt Jaya transect, and what little there is is constant with altitude. Water vapor is at a maximum in January at all elevations. There is greater seasonality in ABL water vapor at Merauke than on the Mt Jaya transect. The ABL and lower free troposphere at Merauke become considerably drier during July than is the case on the Mt Jaya transect.

annual rainfall Papua is one of the wettest regions on earth. Much of Papua regularly receives 2,500–4,500 mm of rain per year, and a few areas receive over 7,000 mm of rain

Figure 2.3.7. Average January and July water-vapor mixing ratios for stations on the Mt Jaya transect (1994–2003, thin solid lines) and also for standard atmospheric pressure levels (height equivalent) above Merauke based on radiosonde observations (, o RAOB), 1994–2001. Water vapor profiles are also shown for July 1999 (dashes) and January 2001 (dots) on Mt Jaya.

................. 16157$

$CH7

02-08-07 10:26:29

PS

PAGE 185

186 / michael l . prentice & g eoffrey s. hope

every year. A map of the provincial pattern of annual rainfall (Figure 2.3.8; Brookfield and Hart 1966) shows that high rainfall occurs along both the north and south sides of the main ranges and around the eastern side of Cenderawasih Bay. High rainfall is also received on the south west of the Vogelkop. Very high annual rainfall, over 7,000 mm per year, is received from Ok Tedi in Papua New Guinea, west along the southern flank of the Merauke Range, past the Baliem Valley, to Timika. No records exist of the highest rainfall point in Papua, but it probably lies at ⬃800 m south of Mt Mandala and has an annual total of ⬃12,000 mm. Tembagapura at 1,950 m asl receives an average of 7,000 mm each year. A long-term record of 6,010 mm per year comes from 700 m asl on the southern slopes at Ninati, but the mid-altitude southern slopes of the main range are virtually uninhabited. It is not clear why the rainfall in the southeast season diminishes farther west because rainfall is also high at this time southwest of the Vogelkop. Within restricted regions, such as on the south side of the Central Range along the Mt Jaya transect, rainfall increases with increasing altitude above the surface (Figure 2.3.9). However, over the whole region, higher annual rainfall is not associated with increasing altitude. The highest alpine areas likely have lower annual rainfall than the mountain flanks. Areas where annual rainfall is below 2,000 mm occur in the Sentani area because of a rain shadow from the Cyclops Mts and the southern third of the Merauke District (Trans-Fly) where the seasonal dry period

Figure 2.3.8. Annual rainfall for Papua. Inset shows a rain shadow around Lake Sentani south of the Cyclops Mts. Source: Brookfield and Hart (1966).

................. 16157$

$CH7

02-08-07 10:26:40

PS

PAGE 186

Climate of Papua / 187

Figure 2.3.9. Average January and July rainfall for stations on the Mt Jaya transect. Rainfall for July 1999 and January 2001 are also shown. is longest (Figure 2.3.8). There is also a zone of lower rainfall (⬍2,500 mm) in the lowlands north of the Central Range, where a rain shadow is developed. Smaller intermontane basins north of the crest of the main range, such as the Ok Sibil, Baliem, Hitalipa, and Wissel lakes areas, are characterized by lower precipitation owing to a rain-shadow effect. At the Wissel Lakes, the Weyland Mts block moisture from the west while the Merauke Range provides a rain shadow from the east. These upland basins support dense populations. A similar structure provides a drier ‘‘neck’’ southeast of the Arfak Range and east of the Bomberai Peninsula. The driest place in Papua is at Barari in Arguni Bay with a mean annual rainfall of 1,020 mm (Brookfield and Hart 1966). This area is in a rain shadow from the ranges south of Cenderawasih Bay and the Bomberai peninsula.

rainfall seasonality Seasonality is not as marked in Papua as in parts of Papua New Guinea and Maluku because of its near-equatorial setting and position within the WPWP region. In many parts of Papua, most rain falls between January and April, the northwest season, with the least falling between May and August, the southeast season. In some parts of the country, this pattern is reversed and most rain is received during the southeast season, when the southeast trades strengthen. The southern slopes of the Central Range provide examples.

................. 16157$

$CH7

02-08-07 10:26:47

PS

PAGE 187

188 / michael l . prentice & g eoffrey s. hope

Along the Mt Jaya transect, the phase of the seasonal precipitation maximum changes at ⬃2,000 m asl, which is the average elevation of the top of the ABL and upper limit of the southeast trades (Figure 2.3.9). Below 2,000 m asl, the precipitation maximum is in the southeast season, whereas, above, the precipitation maximum is in the northwest season. Seasonality in precipitation is much reduced above 2,000 m asl. Measured as the relative difference in rainfall between the dry season and the wet season, the most seasonal parts of Papua are the southern half of Merauke regency (Figure 2.3.10). Merauke receives 1,513 mm, but no month is completely dry, rainfall is ⬎100 mm for every month except August. In many locations, rain is received all year round and there is no seasonal pattern to rainfall amounts. This simply reflects great variability in the sources and intensity of rain. Local, diurnal thunderstorms can provide punishing afternoon and evening showers in winter. Mesocale convective systems bring periods of rain and drizzle for days at a time when the ITCZ moves over the island in southern summer (Tokay and Short 1999).

interannual variability in rainfall Interannual variation in annual rainfall related to ENSO is low over much of Papua, especially in the highlands. Areas where rainfall variability is higher include the southeastern region near Merauke (MacAlpine et al. 1983). The amplitude of seasonal rainfall changes significantly with ENSO events. For instance, Papua experiences periods of uncharacteristically low rainfall during El Nin˜o warm events of ENSO. These events can seriously disrupt food production in Papua, as happened during the 1997–1998 El Nin˜o (Ballard 2000). That drought was associ-

Figure 2.3.10. Monthly temperature and rainfall at Merauke, 1998–2001. Source: Sukri et al. (2003).

................. 16157$

$CH7

02-08-07 10:26:55

PS

PAGE 188

Climate of Papua / 189

ated with heavy frosts at unusually low altitudes, and this killed sweet potato and other crops. Fires broke out across the island, burning normally wet forests. Sukri et al. (2003) present data for an extremely dry season at Merauke during the 2002 El Nin˜o, when almost no rain fell from June until late November. With the onset of rain, a severe dengue fever attack was experienced.

soil water surpluses and deficits The amount of water in the soil available to plants is critical to agriculture. Soil water is measured as ‘‘water balance.’’ Different types of soil can hold different amounts of water. Water can be absorbed or lost from different soils at different rates. The amount of water a soil can hold is known as the ‘‘field capacity’’ of a soil. The balance of water in the soil, measured in millimeters per day, is the difference between the amount of water entering the soil as rain and the amount of water lost from the soil through evaporation (from the surface), transpired by plants (lost from the leaves), or drained downward through the soil beyond the reach of agricultural plants. When a soil is capable of absorbing more water (that is, the soil is below field capacity), but no water is supplied by rainfall (or irrigation), the water balance is said to be in deficit. When water begins to run off the surface of a soil or is drained beyond the rooting zone of agricultural plants, the soil can absorb no more water and is said to be saturated and the soil water balance is said to be in surplus. Although different soils absorb and lose water at different rates, the most important determinant of soil water surpluses and deficits is rainfall. Five patterns of soil water balance can be observed in Papua: • Regular, seasonal, severe soil water deficits occur in the southern part of Merauke regency. • Irregular, moderate soil water deficits occur in the central part of Merauke regency, the Sentani-Genyem area, and Misool Island. • Infrequent, slight soil water deficits occur over much of the lowlands (below 1,200 m asl). • Rare deficits and moderate soil water surpluses occur on the New Guinea mainland at middle altitudes (1,200–1,500 m asl). • Rare deficits and large soil water surpluses occur in the highlands.

These patterns of soil water balance have a strong influence on agricultural systems. Where rainfall is high and regular and soils are usually saturated, the digging of drains to remove water from the soil and planting of certain crops in mounds to raise their roots above the saturated soil is critical for successful agricultural production. Where regular seasonal soil water deficits occur, a number of techniques are used to overcome the lack of soil water for part of the year. This is done primarily through the selection of crops planted. Irrigation in Papua is not common, although water management, such as stream diversion, occurs widely. Taro was once a major crop requiring irrigation, and the last remnants of such systems are still practiced in a few locations, such as the Baliem Valley.

................. 16157$

$CH7

02-08-07 10:26:55

PS

PAGE 189

190 / michael l . prentice & g eoffrey s. hope

Winds Winds are generally light and Papua lies north of the zone of destructive tropical storms, although some cyclone tracks approach the south coast with heavy rain reaching land. The southeast trade winds provide a constant light wind of 10–20 km/hr on the southern coast. Light westerly winds occur during the monsoon season. Strong wind is associated with local systems that meet mountain peaks, as ascending air is compressed and accelerated. Cold air drainage can also deliver bursts of cold air to valley floors at night. Powerful thunderstorms with very high rain intensity are common in Papua, because of the extreme height and size of some systems that may exceed 20,000 m and 50 km across. Extreme vertical wind shear occurs in these storms and is a major aviation hazard. Storm fronts are often associated with violent wind gusts that can cause local damage.

Climate and Glacier Change Climate modeling suggests that the tropics are experiencing temperature increases both on land and in the sea. This may lead to greater variability in the weather in the future. The subject in the New Guinea region is not well researched, but one clear climatic boundary, the extent of glaciers, is known. The Papuan glaciers are of critical importance because, as proven climate recorders, they are the only glaciers within the vast Indonesian low-pressure system. Glaciers on Mt Jaya and elsewhere in Papua (Hope et al. 1973) have been shrinking since they were first photographed in 1907 (Ballard et al. 2001) and 1936 (Colijn 1937). Known collectively as the Carstensz Glaciers, the principal Mt Jaya ice masses of the 1990s consisted of two valley glaciers, the Meren and Carstensz glaciers in the Meren and Yellow valleys, respectively, as well as two highelevation plateau glaciers, the West and East Northwall Firn (Figure 2.3.11). In the first survey of the glaciers in 1936 (Dozy 1938), the West and East Northwall Firn were continuous, and the eastern portion of the Northwall Firn provided ice-flow to the West Meren Glacier and so was part of the West Meren Glacier. The East Northwall Firn also fed an eastward-flowing lobe of the Meren Glacier referred to as the East Meren Glacier or the Harrer Glacier. The East Meren Glacier was substantial in the 1930s, according to geologic evidence. In the 1950s and 1960s, the Northwall Firn separated into western and eastern sections and, by 1987, the East Northwall Firn had effectively separated from the Meren Glacier. The magnitude of the recession of the Carstensz Glaciers, its causes, and its implications for local, regional, and global climate change are only qualitatively known (Prentice and Maryuani 2004). Other ice caps occurred at Mt Idenberg, 15 km west of Mt Jaya, on Mt Trikora, and on Mt Mandala. Ice disappeared from Mt Trikora in the 1960s (Hope, Peterson, and Mitton 1973) and from Mt Idenberg in 1978, leaving a small ice dome on Mt Mandala. These fluctuations have probably been common and the mountains may have been ice-free in the early Holocene, as no evidence for glaciation is known from this time (Prentice et al. 2005).

................. 16157$

$CH7

02-08-07 10:26:56

PS

PAGE 190

Climate of Papua / 191

Figure 2.3.11. Map view of the Carstensz Glaciers and changes in their extent since 1936. The 1972, 1942, and 1936 boundaries of the East Meren Glacier are schematic.

glacier area and length changes

The recession of the Carstensz Glaciers from ⬃11 km2 in 1942 to 2.4 km2 by 2000 represents about an 80% decrease in ice area (Figure 2.3.12). The Carstensz Glacier itself decreased slightly less, ⬃70%, over this interval. The West and East Meren Glaciers melted away completely between July 1997 and February 1999. Compari-

Figure 2.3.12. Changes in the area of the Mt Jaya glaciers between 1936 and 2000.

................. 16157$

$CH7

02-08-07 10:27:20

PS

PAGE 191

192 / michael l . prentice & g eoffrey s. hope

son of the Meren Glacier area in 1942 to the East Northwall Firn area in 2000 yields an 80% decrease in the area of this ice system. A longer record with more temporal detail is that of the recession of the Carstensz and West Meren Glacier fronts. Between 1936 and 2000, the Carstensz Glacier front receded about 1.2 km, a decrease in length of 46%. Over the same interval, the 2.6 km recession of the West Meren Glacier represents a 100% decrease in length. However, given the separation of the Meren from the East Northwall Firn, it is more reasonable to compare West Meren Glacier length in 1942 to East Northwall Firn length in 2000. The latter comparison yields a 62% decrease in length of the Meren-East Northwall Firn ice system. Recession rates are useful for resolving the causes of glacier recession. To isolate the effect of climate forcing, we look for the common signal in the recession-rate histories and attempt to factor out the influence of topographic complexity. The dominant pattern of the frontal recession-rate histories for the Carstensz and the West Meren glaciers appears (this is a low-resolution record) to be that the recession rate increased from the 1940s to a peak in 1995–1996, after which it decreased slightly. On the other hand, the rates of decrease in ice area exhibit considerable discrepancies and so do not support a single climate-forcing hypothesis. The rate of change in the Carstensz Glacier area indicates a progressive increase in climate forcing to a peak in 1995–1996. However, the rate in the reduction of area of the other ice masses deviates significantly from the Carstensz pattern.

glacier mass balance and ela changes For 1973–1994, the distribution of the total net snow balance (sum of snow gain and loss) with elevation on the Carstensz and West Meren glaciers indicates that, for the 22-year period, the balance was zero at ⬃4,780 m asl. Hence, the altitude of the equilibrium line (ELA) averaged 4,780 m asl between 1973 and 1994. This is consistent with the ELA that is apparent on the SPOT satellite imagery of July 1987, which is a transient ELA. The average ELA of 4,780 m represents a rise in the ELA of 120 m since 1972, when the ELA was determined to be 4,660 m asl, after correction of the Carstensz Glaciers Expedition topographic map (Hope et al. 1976) to the 1995 topographic map. For the two-year interval 1995–1996, the total net snow balances, Bn, for the Carstensz Glaciers were determined by photogrammetry to be all quite negative. The two largest glaciers, the Carstensz and East Northwall Firn, had similarly negative budgets that, on an annual basis, were –4,678  106 kg in water equivalent (average surface lowering of 3.3 m) and –4,739  106 kg (average surface lowering of 4.5 m), respectively. The distribution of Bn with altitude shows that the average ELA for the two years was above the highest glacier surfaces, which were at 4,940 m asl. This balance-based estimate for the ELA contrasts sharply with two different estimates of the ELA based on observation of the transient snow line. Those observations put the snow line at 4,650–4,750 m asl, which is well below our average ELA. For the period 1997–1999, Bn for the Carstensz Glacier and West Northwall Firn

................. 16157$

$CH7

02-08-07 10:27:21

PS

PAGE 192

Climate of Papua / 193

were slightly negative, while Bn for the East Northwall Firn was slightly positive. The West Meren Glacier melted away between 1997 and 2000 and so its Bn was negative but not quantifiable. Annual surface lowering of the Carstensz Glacier and the West Northwall Firn averaged 1.3 and 1.6 m/yr, respectively, whereas the surface of the East Northwall Firn gained altitude slightly. The general glacier retreat between 1972 and 2000 is broadly consistent with the warming trend in the lower free troposphere over western New Guinea during this interval. Additionally, there appears to be some correlation between interannual specific-humidity variations and glacier fluctuations. By piecing together scattered and discontinuous highland and radiosonde weather observations, we speculate that, at glacier altitudes, mean annual temperature increased ⬃1C over this 30-year period. This likely had some effect on the glaciers, but, at finer temporal resolution, the relationship is blurred for lack of contemporaneous data. Temperatures at sea level around western New Guinea increased significantly from a major decadal low in 1992–1993 to highs in 1995–1996 and 1996–1997 that were the highest since 1988–1989. These highs coincide with the estimates for highly negative mass balances and very high glacier ELA for 1995–1996. In addition to these effects, the frequent intervals of modeled low water vapor between 1991 and 1995 stand out as a significant series of events that preceded the 1995–1996 negative glacier mass balances. The prolonged near-drought conditions should have negatively effected the mass balance. From 1997 to 2000, the ELA was probably lowered because mass balances were more positive. This may relate to increased precipitation. Despite this reprieve, the ice is clearly marginal under current conditions of global warming and so may be lost from New Guinea within a few decades.

Discussion Papua has unusually wet climates in both an Indonesian and a global context, with large areas of the mountains too wet and cloudy to support viable agriculture. This moisture buffers the province against dry seasons and maintains thermal equability. The elevation of the high mountains in the late Tertiary set the main climate parameters, and the Papuan biota has adapted to the very great thermal range but low local variability since then. Even glacial shifts, while depressing snow-line and vegetational boundaries, did not remove the general moisture surpluses, except perhaps in the south. The impact of global warming may similarly be buffered in that rainfall in the highlands will probably be enhanced, although variability may also increase, with more frequent and severe El Nin˜o and La Nin˜a events possible. This will stress vegetation in the alpine zone and at boundary positions. The alpine area will be liable to invasion by shrubs and trees but, currently, fire maintains large tracts of open grasslands, so only alpine obligates will be under threat. Likewise, significant changes in alpine lake physical and chemical properties will threaten their flora and fauna. Some retreat of forests and change in forest composition might be

................. 16157$

$CH7

02-08-07 10:27:22

PS

PAGE 193

194 / michael l . prentice & g eoffrey s. hope

expected at the rainforest limit near Merauke. Severe frosts from future El Nin˜o events may kill off frost-sensitive species in upper montane forests and limit agriculture.

Acknowledgments We thank Khoiril Maryunani for permission to use unpublished results from the 2002 WEPAMA cruise of the RV Marion Dufresne and PT Freeport Indonesia for access to glacier data as well as weather data from stations in their lease area around Tembagapura. Michael Bourke kindly provided information about rainfall categories.

Literature Cited Ballard, C. 2000. Condemned to repeat history? ENSO-related drought and famine in Irian Jaya, Indonesia. Pp. 123–148 in Grove, R., and J.M.A. Chappell (eds.) El Nin˜oHistory and Crisis. White Horse Press, Cambridge. Ballard, C., S. Vink, and A. Ploeg (eds.). 2001. Race to the Snow: Photography of the Exploration of Dutch New Guinea, 1907–1936. KIT (Royal Tropical Institute), Amsterdam Barmawijaja, B.M., E.J. Rohling, W.A. van der Kaars, C. Vergnaud Grazzini, and W.J. Zachariasse. 1993. Glacial conditions in the northern Molluca Sea region (Indonesia). Palaeogeography, Palaeoclimatology and Palaeoecology 101: 147–167. Barry, R.G. 1978a. Aspects of the precipitation characteristics of the New Guinea mountains. Journal of Tropical Geography 47: 13–30. Barry, R.G. 1978b. Mountain climates of New Guinea. Pp. 75–110 in van Royen, P. (ed.) Alpine Flora of New Guinea. Cramer Verlag, Vaduz. Bellamy, J.A., and J.R. McAlpine. 1995. Papua New Guinea Inventory of Natural Resources, Population Distribution and Land Use Handbook. 2nd edition. PNGRIS Publication No. 6. Australian Agency for International Development, Canberra. Brookfield, H. 1964. The ecology of highland settlement: some suggestions. American Anthropologist 66: 20–38. Brookfield, H. 1989. Frost and drought through time and space: what were conditions like when the High Valleys were settled? Mountain Research and Development 9: 306–321. Brookfield, H., and B. Allen. 1989. High altitude occupation and environment. Mountain Research and Development 9: 201–209. Brookfield, H.C., and D. Hart. 1966. Rainfall in the tropical southwest pacific. Department of Geography Publ. G/3, Research School of Pacific Studies, Australian National University. Australian National University Press, Canberra. Colijn, A.H. 1937. Naar de Eeuwige Sneeuw van Tropisch Nederland. Scheltens and Giltay, Amsterdam. Fitzpatrick, E.A., D. Hart, and H.C. Brookfield. 1966. Rainfall seasonality in the tropical southwest Pacific. Erdkunde 20: 181–194. Hastenrath, S. 1991. Climate Dynamics of the Tropics. Kluwer Academic, Dordrecht. Hnatiuk R.J., J.M.B. Smith, and D.N. McVean. 1976. Mt. Wilhelm Studies, 2. The climate of Mt. Wilhelm. Research School of Pacific Studies, Department of Biogeography and Geomorphology Publications BG/4. Australian National University, Canberra.

................. 16157$

$CH7

02-08-07 10:27:22

PS

PAGE 194

Climate of Papua / 195 Hope, G.S., J.A. Peterson, and R. Mitton 1973. Recession of the minor ice fields of Irian Jaya. Zeitschrift fu¨r Glestcherkunde und Glazialgeologie IX: 73–87. Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.). 1976. The Equatorial Glaciers of New Guinea. Results of the 1971–1973 Australian Universities’ Expeditions to Irian Jaya: survey, glaciology, meteorology, biology and paleoenvironments. A.A. Balkema, Rotterdam. Kol, E., and J.A. Peterson. 1976. Cryobiology. Pp. 81–91 in Hope, G.S., J.A. Peterson, U. Radok, and I. Allison (eds.) The Equatorial Glaciers of New Guinea. A.A. Balkema, Rotterdam. Lam, H.J. (trans. L.M. Perry). 1945. Fragmenta Papuana 1–7. Sargentia V: 1–196. Lukas, R., J.P. McCreary, and T. Yamagata. 1996. Pacific low-latitude western boundary currents and the Indonesian Throughflow. J. Geophysical Res. 101: 12209–12216. McAlpine, J.R., G. Keig, and R. Falls. 1983. Climate of Papua New Guinea. CSIRO/ Australian National University Press, Canberra. McAlpine, J.R., and K. Short. 1974. Water balance estimates for Papua New Guinea. Technical Memorandum 74/9, CSIRO Division of Land Use Research, Canberra. Nix, H.A., and J.D. Kalma. 1972. Climate as a dominant control in the biogeography of northern Australia and New Guinea. Pp. 61–91 in Walker, D. (ed.) Bridge and Barrier: The Natural and Cultural Heritage of the Torres Strait. Department of Biogeography and Geomorphology, Research School of Pacific Studies, Australian National University, Canberra. Philander, S.G.H. 1990. El Nin˜o and the Southern Oscillation. Academic Press, London. Prentice, M.L., G.S. Hope, K. Maryunani, and J.A. Peterson. 2005. An evaluation of snowline data across New Guinea during the last major glaciation, and area-based glacier snowlines in the Mt. Jaya region of Papua, Indonesia, during the Last Glacial Maximum. Quaternary International 138–139: 93–117. Prentice, M.L., and K. Maryunani. 2002. The history of the Carstensz Glaciers 1936 to 1999 and relations to climate change. Report to PT Freeport Indonesia. Smith, J.M.B. 1977. Vegetation and microclimate of east and west facing slopes in the grasslands of Mt Wilhelm, Papua New Guinea. Journal of Ecology 65: 39–53. Sukri, N.C., K. Laras, T. Wandra, S. Didi, R.P. Larasati, J.R. Rachdyatmaka, S. Osok, P. Tjia, J. Saragih, M.S. Hartati, E. Listyaningsih, K.R. Porter, C.G. Beckett, I.S. Prawira, N. Punjabi, S.A. Suparmanto, H.J. Beecham, M.J. Bangs, and A.L. Corwin. 2003. Transmission of epidemic dengue hemorrhagic fever in easternmost Indonesia. American Journal of Tropical Medicine and Hygiene 68 (5): 529–535. Tokay, A., and D.A. Short. 1999. Tropical rainfall associated with convective and stratiform clouds: intercomparison of disdrometer and profiler measurements. Journal of Applied Meterology 38: 302–320. Tomczak, M., and J.S. Godfrey. 1994. Regional Oceanography: An Introduction. Pergamon, Oxford. Webster, P.J., and R. Lucas. 1992. TOGA COARE: the coupled ocean-atmosphere response experiment. Bulletin of the American Meterological Society 73: 1377–1416. Webster, P.J., and N.A. Streten. 1978. Late Quaternary Ice Age climates of tropical Australasia: interpretations and reconstructions. Quaternary Research 10: 279–309. Wyrtki, K. 1961. Scientific results of marine investigations of the South China Sea and the Gulf of Thailand 1959–1961. Naga Report, Vol. 2.

................. 16157$

$CH7

02-08-07 10:27:22

PS

PAGE 195

2.4. Papuan Terrestrial Biogeography, with Special Reference to Birds bruce m. beehler i o ge o g r ap h y i s t h e s tu d y of the natural distribution of plants and animals on earth. This chapter describes some general patterns in the distribution of terrestrial life across Papua. First, I summarize the geological evolution of Papua in the context of New Guinean plate tectonics. Second, I define a set of discrete Papuan physical regions (defined mainly by current physiography) and provide short but illustrative lists of species endemic to these geographic regions. Third, I illustrate geographic patterns of differentiation. The chapter concludes with a discussion of the possible processes that have generated these distinctive patterns. Once other major taxa of plants and animals are better surveyed, monographed, and mapped, I suspect that additional patterns will emerge that are the product of yet other, and in some cases more ancient, processes than those that influenced birds. The current (and past) distribution of plants and animals in Papua is astoundingly little-known, even when compared with that for poorly-known Papua New Guinea (Gressitt 1982; Beehler 1993; Supriatna 1999). This makes the task of providing a coherent overview of the subject difficult, and thus provides a compelling argument for the institution of a comprehensive biological survey of western New Guinea. Because of this, these discussions will be restricted primarily to patterns of distribution in birds (Beehler and Finch 1885; Beehler et al. 1986) and to a lesser extent mammals (Flannery 1995a,b; Helgen, pers. comm.), two of the betterdocumented major taxa (cf. Pratt 1982), but where possible the chapter provides some data on additional taxa (Allison 1993, 1996; Parsons 1999).

B

Tectonic History Given the comprehensive treatment provided in the chapter in this volume on tectonics (Chapter 2.1), and extensive discussion on the subject in additional chapters herein, I will here only briefly summarize the main geo-tectonic points of interest in order to delineate the major physical regions for Papua, about which I can pose biogeographic questions, and within which I can look for particular patterns of endemism. Described in simplest terms, present-day Papua includes a number of major landforms that can serve as subunits for delineating the current distribution of the biota. We now know that these present landforms are the product of a northwardmoving Australian Plate that, through collision with the westward-moving Pacific Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

196

................. 16157$

$CH8

02-08-07 10:26:14

PS

PAGE 196

Papuan Terrestrial Biogeography, with Special Reference to Birds / 197

Plate, has suffered considerable compression at its northern plate boundary as well as the accretion of a series of island arcs/terranes along its margin. As detailed by Polhemus (Chapter 2.1, the main source for the following brief narrative), the major resulting geographic components of this tectonic scenario include a stable Australian cratonic platform to the south, a New Guinea Fold Belt that constitutes the Central Cordillera, and a Mobile Belt to the north that includes the north coastal ranges and the various associated Papuan islands (Pigram and Davies 1987). Things appear more complex in the Bird’s Neck and Vogelkop regions, where contact with the Banda Arc seems to have produced shear and north-south movement in the Bird’s Neck and Wandammen Peninsulas, as well as the Onin and Bomberai peninsulas to the southwest. A close examination of Figure 2.4.1 shows that for the main body of Papua a majority of features trend northwestsoutheast, contrasting with the great curving bend of the Bird’s Neck that forms the current physical link between the isolated Vogelkop and Raja Ampat Islands to the west and northwest. The large oceanic gap created by the Cenderawasih Bay signals the dissection of western elements from the eastern (main) elements of geographic Papua. Thus the main body of Papua has tectonically mirrored what has taken place in central and western mainland Papua New Guinea, and the Vogelkop and islands components of the western half of Papua appear analogous to the more complex tectonic history of Papua New Guinea’s far southeast (Pa-

Figure 2.4.1. Papua Bird Regions.

................. 16157$

$CH8

02-08-07 10:26:20

PS

PAGE 197

198 / bruce m. beehler

puan Peninsula and Eastern Papuan Islands in association with the Bismarck Arc). In fact, the arc-shape of Seram, just to the west of Papua, is a reverse mirror image of New Britain to the north and east of Papua New Guinea. Both are evidence of the rotational interplay of plates and microplates in active and very complex tectonic segments of the Pacific along the great Ring of Fire. The Vogelkop includes a core of original Australian craton that separated from that main plate, plus accreted terrane components (North Tamrau, Arfak). The adjacent Raja Ampat Islands include a northern unit and southern unit, separated by the biogeographically significant Sagewin Strait. Some general patterns can be reviewed here. Those terranes that were initially accreted form the northernmost ridges of the Central Cordillera. The TarikuTaritatu (Mamberamo) Basin separates this fold belt from the north coastal ranges, uplifted Miocene island arcs that include the Cyclops, Foja (Gauttier/ Foya), and Van Rees (Rouffaer) ranges as well as mountainous Yapen Island. The various terrane-docking events produced uplift and rapid mountainbuilding, with associated downwarping of intermontane troughs/strike-slip faults (Tariku-Taritatu Trough, Sorong-Tamrau Fault). The northern front range of the Central Cordillera is quite impressive, with summits topping 3,700 m today. Further west, the Tamraus and Arfaks reach in excess of 2,800 m. All other northern ranges (Van Rees Range 1,500 m, Fojas 2,200 m, Cyclops 2,300 m) fail to reach high elevations, perhaps because of youth, or because of the counter-effects of rapid erosion. The greatest single tectonic feature in Papua is the western New Guinea orogen—the vast cordillera whose compressionary mountain-building has produced the highest summit in the Pacific (i.e., Puncak Jaya 4,884 m). Southward, this descends onto the stable and undeformed Australian craton, engulfed by a deep wedge of outwash sediments that have been eroded off the southern scarp of the main cordillera.

Natural Geography Plate movements over the last twenty-five million years have molded a Papua that today can be dissected into thirteen biogeographic subunits, which are briefly described below (Figure 2.4.1). Here I treat these in geographic fashion, from northwest to southeast. These are adapted from a set of ‘‘New Guinea Bird Regions’’ being developed for use in a revision of Beehler et al. (1986), so regional nomenclature is based on the whole island of New Guinea rather than Papua. Thus the region ‘‘southwest lowlands’’ in the Papuan context might be more appropriately ‘‘southern lowlands,’’ but the New Guinea-wide terminology is retained here for purposes of terminological stability and regional uniformity (New Guinea being a natural region; Papua being a politically-delineated subunit of the New Guinea region).

................. 16157$

$CH8

02-08-07 10:26:21

PS

PAGE 198

Papuan Terrestrial Biogeography, with Special Reference to Birds / 199

waigeo/batanta These two major islands of the Raja Ampat group, plus their associated islands, are mountainous and rugged and capped with young limestone, separated from each other by rather shallow straits. Kofiau can be allied with this unit for convenience, although it is apparently not tectonically a part of this terrane, as witnessed by Kofiau’s avian endemics Tanysiptera elliotti and Monarcha julianae. Faunally, Waigeo and Batanta share two prominent paradisaeid endemics, Paradisaea rubra and Cicinnurus respublica, a lowland and a montane bird of paradise, respectively. The bizarre brush-turkey (Aepypodius bruijnii) is known only from the hills of Waigeo, but there are some indications that it also inhabits the uplands of Batanta. The phalangerid Spilocuscus papuensis is a Waigeo/Batanta endemic also (Chapter 4.10).

misool/salawati Salawati and Misool islands lie on a shallow platform that is apparently fully docked to the western margin of the Australian craton. Misool is low but rugged with a cap of karstic limestone. Skeins of narrow parallel high karst islets stretch eastward from the east coast of Misool, evidence of some recent compression and uplift. Biotically, this unit appears to be most closely allied with the lowlands of the southern Vogelkop (for example, Cicinnurus magnificus and Paradisaea minor are shared with the Vogelkop and are absent from neighboring Waigeo/Batanta). Salawati’s northern coastal range faces the Sagewin Strait that separates Salawati from mountainous Batanta. The Sagewin Strait constitutes an important local biogeographic discontinuity that is allied to tectonic history (an extension of the Sorong fault). Fewer than five kilometers wide, the Strait constitutes a major vicariance break for the aforementioned species of Paradisaea (rubra vs. minor) and Cicinnurus (respublica vs. magnificus). In addition, the lowland paradisaeid species Cicinnurus regius and Seleucidis melanoleuca occurs south of the Strait but not on the northern side, as well as Casuarius bennetti (north side) and Casuarius unappendiculatus (south side). A thorough biogeographic analysis across this strait will no doubt produce a long list of plants and animals that have differentiated across this distributional barrier. It will be interesting to see if there is a similar pattern of discontinuity across this break on the Vogelkop, between montane biotas of the North Tamrau and the South Tamrau mountains.

vogelkop Otherwise known as the Bird’s Head, Kepala Burung, or Berau Peninsula, the mighty Vogelkop is, without doubt, the second most significant physical feature of the Papuan region after the Central Cordillera. The Vogelkop is separated from the main expanse of the New Guinea orogen, and composed of several accreted terranes that have been impacted by major mountain-building. Biologically, the Vogelkop is a very distinct Papuan subunit, with a wide range of birds and mammals endemic to it, although many of these species range into the highest summits

................. 16157$

$CH8

02-08-07 10:26:22

PS

PAGE 199

200 / bruce m. beehler

of the adjacent Bird’s Neck. Geographic species-pairs have formed on either side of the Bird’s Neck, with a Vogelkop form to the northwest and a central cordilleran form to the southeast. The strict Vogelkop endemics include the bird species Astrapia nigra and Lonchura vana, as well as the ringtail species Pseudochirops coronatus and Pseudochirulus schlegeli, a bandicoot Microperoryctes aplini, and a murid Stenomys arfakianus, among others (Chapter 4.10), plus several species of endemic herpetofauna (Chapter 4.5).

bird’s neck The Bird’s Neck is Papua’s most complex physiographic subunit, with a concatenation of terranes that show shear, extension, and north-south compression and uplift that relates to some rotational contact with the Banda Arc to the west and southwest. It is the only place in Papua where one can find deep fjordlands (Triton, Etna, and Arguni bays). Biotically, this unit is closely affiliated with the Vogelkop, and many of the regional avian endemics inhabit this pair of subunits (VogelkopBird’s Neck): Rallicula leucospila, Sericornis rufescens, Pachycephala meyeri, Melidectes leucostephes, Melipotes gymnops, Paradigalla carunculata, Parotia sefilata, and Amblyornis inornatus, plus the tree kangaroo Dendrolagus ursinus and the monotreme Zaglossus bruijnii. Also of note is that the Wandammen Peninsula supports the local endemic Dendrolagus mayri (Chapter 4.10) and several species of endemic herpetofauna (Chapter 4.5)

biak/numfoor These oceanic islands constitute the most distinct unit in the entire region (even more distinct than Kofiau Island west of Batanta). This oceanic unit shows no real affiliation with neighboring units. Nearby Yapen shows no biogeographic affinities with this unit, in spite of its proximity and insular nature (Yapen, in fact, is a continental land-bridge island). Many insular avian endemics were treated as races of more widespread forms by Mayr (1941), thus hiding the extent of this unit’s uniqueness. A list of species endemics includes the birds Eos cyanogenia, Megapodius geelvinkianus, Micropsitta geelvinkiana, Centropus chalybeus, Otus beccarii, Monarcha brehmii, Myiagra atra, Tanysiptera riedelii, Tanysiptera carolinae, Aplonis magna, Gerygone hypoxantha, and Zosterops mysorensis, and the mammals Dobsonia emersa, Spilocuscus wilsoni, Uromys boeadii, and Uromys emmae (Chapter 4.10).

yapen island Yapen Island is part of the Mobile Belt, and affiliated with the Sorong fault that we find dissecting the Tamrau Mountains to the west. Yapen apparently has been carried westward into Cenderawasih Bay from the allied Van Rees (Rouffaer) Range. Yapen shows affiliation with the northwestern lowlands and north coastal ranges, and supports no avian endemic species. Neither are there mammalian endemics to Yapen, although there are endemic frogs (Richards, pers. comm.).

................. 16157$

$CH8

02-08-07 10:26:22

PS

PAGE 200

Papuan Terrestrial Biogeography, with Special Reference to Birds / 201

Yapen is a land-bridge island, which explains its relative lack of endemism (in stark contrast to Biak/Numfoor).

northwestern lowlands The northwestern lowlands is a product of subduction of the northern edge of the Australian Plate beneath the Pacific Plate. Its affiliated endemics include two birds: the fig-parrot Psittaculirostris salvadorii and the friarbird Philemon brassii. There are no documented mammalian endemics, although there are a number of mammals restricted to the northern lowlands of New Guinea. Low endemism for this lowlands unit is expected, mainly because of weak physical isolation from adjacent lowland areas east and west.

north coastal ranges The north coastal ranges are a product of island arc collisions with New Guinea from Oligocene to Pliocene. The oldest contact and mountain-building occurred in the west and the youngest in the east. Three avian endemic are associated with this unit, all confined to the Foja Mountains—the Golden-maned Bowerbird, an as yet undescribed Melipotes honeyeater, and a six-wired bird of paradise, Parotia berlepschi (currently being restored as a full species subsequent to recent fieldwork in the Foja Mts). In addition, the recent survey has identified more than twenty endemic frogs, several endemic butterflies, and a number of endemic plants. The echidna Zaglossus attenboroughi is endemic to the Cyclops Range. Of course, there are many mammalian endemics that range though the northern coastal ranges of the island of New Guinea (e.g., Paraleptomys rufilatus, Dendrolagus pulcherrimus, D. scottae; cf. Helgen 2005).

sudirman range The Central Cordillera is divided into two units, separated by the Baliem Valley that flows northwest-southeast and forms a physical break between the highest summits of the west with those to the east. The western unit supports the highest mountains in all of the tropical Pacific (Sudirman Range), high-piled marine sediments that exceed 4,500 meters in a number of summits. The Sudirman Range supports a number of restricted range endemics, although many extend into the Jayawijaya/Star Mountains. Species endemics confined to the west include the birds Anurophasis monorthonyx, Petroica archboldi, Lichenostomus chrysogenys, and Lonchura teerinki, and many mammals (e.g., Coccymys albidens, Baiyankamys habbema, Dendrolagus mbaiso, Mallomys gunung, and Rattus omichlodes). More remarkable, among the mammals, is that the isolated Weyland Range, in the far west of this region, is home to a number of mammalian endemics (Chapter 4.10) and species of herpetofauna (Chapter 4.5). There is currently no evidence that there is comparable avian endemism in the Weylands.

jayawijaya/star mountains The Jayawijaya/Star Mountains region includes impressive high ranges only slightly less grand than those of the west, but is much less-well surveyed and rather

................. 16157$

$CH8

02-08-07 10:26:23

PS

PAGE 201

202 / bruce m. beehler

poorly-characterized biotically. There are no bird endemics, but several species of herpetofauna (Chapter 4.5) and at least one mammal endemic is known from the PNG sector of this region (e.g., Phalanger matanim).

aru islands The Aru Islands are land-bridge islands with close affinities to the southern lowlands and Trans-Fly. The Arus are the only place in the region where the rail Eulabeornis castaneoventris breeds (also found to breed in northern Australia). The Arus share the kingfisher Tanysiptera hydrocharis with the Trans-Fly, and the fruitdove Ptilinopus wallacii with the Southern Lowlands. The Arus possess no endemic bird species.

southwestern lowlands The southwestern lowlands is a triangular wedge of craton that widens strongly from west to east. It is Papua’s largest single expanse of lowland rainforest, having been built up from massive outwash of sediment-laden high-energy rivers that drain the steep and rain-drenched southern scarp of the Central Cordillera. The bird of paradise Paradisaea apoda and the mammal Uromys scaphax are nearendemics (Chapter 4.10). Endemism is low mainly because lowland zones have wide-ranging species and that there is a minimal the distributional barrier separating this southwest lowlands region from the Fly-Purari region to the east.

trans-fly The Trans-Fly is geologically allied to the southwestern lowlands, but distinct in being a highly seasonal savanna landscape that continues eastward into Papua New Guinea. The Trans-Fly has important relationships across the border and also with northern Australia, but is home to several near-endemics: the birds Megalurus albolimbatus, Lonchura stygia, and Lonchura nevermanni, and the mammal Dasyurus spartacus (Chapters 4.10, 5.12).

Patterns of Distribution

major patterns Again, based mainly on the avian distributional data, several general patterns of distribution can be seen in Papua. I focus on birds (as do Mack and Dumbacher, Chapter 4.9), and note that other contributors to this volume have considered biogeographic patterns in other taxa (e.g., freshwater fauna: Polhemus and Allen, Chapter 2.5; herpetofauna: Allison, Chapter 4.6; mammals: Helgen, Chapter 4.10; cicadas: Duffels and de Boer, Chapter 4.4). Lowland forest forms tend to be widespread but confined largely one of the three lowland zones: the northern or southern watersheds or the lowlands of the far west (VogelkopBird’s Neck). Lowland sister-species often meet and hybridize on the eastern verge of the Bird’s Neck where the Central Cordillera ends. This three-part pattern is found for the avian genera Talegalla, Goura, Paradisaea, Psittaculirostris, and Chalcopsitta. Thus, for

................. 16157$

$CH8

02-08-07 10:26:24

PS

PAGE 202

Papuan Terrestrial Biogeography, with Special Reference to Birds / 203

example, in the crowned pigeons, Goura cristata inhabits the lowlands of the Vogelkop and Bird’s Neck; Goura victoria inhabits the lowlands of the northern watershed of the main body of New Guinea (ranging into PNG); Goura scheepmakeri inhabits the southern lowlands (ranging into PNG). Montane distributions tend to range northwest to southeast, following the trend of the main New Guinean cordillera plus the highlands of the Vogelkop, a slightly offset and isolated part of Papua’s montane uplands. Montane species-groups break up variously, as outlined by the following examples. Paradigalla brevicauda ranges westward along the Central Cordillera from PNG to the western terminus of the Western Cordillera. It is replaced in the highlands of the Bird’s Neck and Vogelkop by Paradigalla carunculata. In a similar fashion, Astrapia splendidissima ranges along the Central Cordillera to its western terminus, to be replaced by Astrapia nigra in the uplands of the Vogelkop. In a slightly more complex pattern, the bowerbird genus Amblyornis includes a main cordilleran form (macgregoriae), a Vogelkop/Bird’s Neck form (inornatus), plus a North Coastal Range form (flavifrons). Additional examples of southeast-northwest cordilleran differentiation include Sericornis perspicillatus/rufescens and Melipotes fumigatus/gymnops.

minor patterns Other cordilleran forms sort out elevationally, as with the avian genus Epimachus, with meyeri inhabiting higher elevations throughout the length of the New Guinean cordillera, and fastuosus inhabiting slightly lower elevations and a range that is shifted to the west (from the Vogelkop eastward to central Papua New Guinea). There are several examples of montane speciation involving sister vicariants in the Central Cordillera and the north coastal ranges and include the genera Amblyornis, Melipotes, and Parotia. Apparently other major taxa speciate strongly in the northern ranges (e.g., frogs, mammals, plants). Clearly different processes may be operating to generate different patterns between the major taxa. In particular, plants and insect lineages may be considerably older than those of birds and may better track recent plate tectonics. This has been highlighted by Heads (2001a, 2002). More and better mapping of lineages is needed to elucidate these various patterns.

possible processes producing these detected patterns It is evident that most patterns are produced by allopatric speciation though various forms of vicariance. This means that sister populations that are geographically isolated by some physical or ecological barrier differentiate and are allowed to speciate because of this geographic isolation, which produces a break in gene flow. Diamond (1972, 1973) described his ‘‘dropout’’ phenomenon to explain the differentiation of east-west montane forms along the Central Cordillera of New Guinea. This invoked the creation of a break in the widespread range of a montane parent species, which subsequently permitted differentiation in isolation of an eastern and western form. With regard to differentiation of lowland forms, the Central Cordillera and the Bird’s Neck seem to offer suitable barriers to speciation. I thus posit that much of the speciation seen in Papuan birds is a product of classic

................. 16157$

$CH8

02-08-07 10:26:24

PS

PAGE 203

204 / bruce m. beehler

allopatric vicariance (Mayr 1963). It will be interesting to see whether such patterns also apply to the lesser-known major taxa (e.g., plants, arthropods) on the mainland of Papua. It must also be noted that there is also clear evidence of over-water dispersal leading to the evolution of island endemics. Witness the current presence of island-endemics in the kingfisher genus Tanysiptera on Biak, Numfoor, and Kofiau. The same process can be invoked to explain the whole range of avian endemics found on Biak Island. Note that none of these processes requires the movement of plates or terranes. For these relatively youthful avian lineages, it is apparent that species-level differentiation has taken place on a time frame that invokes primarily ecological barriers to dispersal to create vicariance events suitable for speciation. At higher levels (subfamily, family) current patterns of distribution have been examined by Heads (2001a,b, 2002), producing evidence (among plants and some animal groups) of lineage ‘‘massing’’ in relation to the plate tectonic scenarios described earlier in this chapter. For instance, New Guinea’s snakes and freshwater fishes are most abundant in southern New Guinea that remains a part of the Australian craton. The birds of paradise mass in New Guinea’s Central Cordillera. The flying foxes and Platymantis frogs mass on the volcanic islands north of New Guinea. These provide evidence of older processes that underlie the more modern events highlighted here. An in-depth analysis of the evolution and phylogeography of major regional plant and animal lineages is beyond the scope of this chapter, but would no doubt shed light on the biogeographic patterns observed on the island today.

anomalies Returning to birds and modern speciation events, there are, of course, anomalies that defy simple biogeographic explanation. For instance, the Cnemophiline ‘‘false bird of paradise’’ Cnemophilus macgregorii inhabits the high mountains of the Central Cordillera of PNG and extends west only partially into Papua’s Western Cordillera. Why it does not range westward throughout the Papuan cordillera to the Weyland Range is a bit of a mystery (when many other montane species like Cnemophilus loriae and many mammalian species range from the far west to the far east). Another peculiar anomaly is the presence of Otus beccarii on Biak Island. It is the only place Otus is found in the Papuan region, its nearest occurrence being in Wallacea to the west. Why is Otus not found on the intervening Vogelkop or Waigeo Island? One might consider the highly-restricted distribution of the Snow Robin (Petroica archboldi) to be anomalous. Is it a sister-form of the only other Petroica in the region, P. bivittata? If so, why is its tiny range (confined to the highest summits of the Western Cordillera) entirely contained in the much broader distribution of the montane P. bivittata? What process allowed P. archboldi to differentiate from P. bivittata? These are just a few trenchant examples of anomalies that will perhaps become more diverse when there are sufficient distributional data for plants and other animal groups. Needless to say, future

................. 16157$

$CH8

02-08-07 10:26:25

PS

PAGE 204

Papuan Terrestrial Biogeography, with Special Reference to Birds / 205

studies of Papuan plants and animals will generate both biogeographic oddities as well as clear and repeated patterns for decades to come.

The Way Forward: Future Studies Certainly, molecular phylogeographic studies as described by Mack and Dumbacher (Chapter 4.9) will continue to greatly clarify the natural distribution of specific clades as well as major lineages of the terrestrial biota of the better-known taxa (such as birds and mammals). In the instances of the less well surveyed taxa, long-term and in-depth field survey of key sub-lineages (focal genera) may pay great dividends, allowing one to get an idea of patterns of speciation or diversification (Heads 2001a,b, 2002; Welzen et al. 2001). These, too, will benefit from molecular systematic clarification of patterns of clade formation. Once done, it will then be possible to see if there are concordant patterns of distribution across the major taxa, which will allow possible resolution of general area cladograms that describe the overall patterns of biotic diversification on New Guinea.

Literature Cited Allison, A. 1993. Biodiversity of the fishes, amphibians, and reptiles of New Guinea. Pp. 57–226 in Beehler, B.M. (ed.) A Biodiversity Analysis for Papua New Guinea. Vol. 2 Conservation Needs Assessment. Biodiversity Support Program, Washington, D.C. Allison, A. 1996. Zoogeography of amphibians and reptiles of New Guinea and the Pacific region. Pp. 407–436 in Keast, A., and S. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Beehler, B.M. (ed.). 1993. A Biodiversity Analysis for Papua New Guinea. Vol. 2. Papua New Guinea Conservation Needs Assessment. Biodiversity Support Program, Washington, D.C. Beehler, B.M., and B.W. Finch. 1985. Species Check-List of Birds of New Guinea. Monograph No. 1. Royal Australian Ornith. Union, Melbourne, Victoria. Beehler, B.M., T.K. Pratt, and D.A. Zimmerman. 1986. Birds of New Guinea. Princeton University Press, Princeton, New Jersey. Diamond, J.M. 1972. Avifauna of the Eastern Highlands of New Guinea. Nuttall Ornith. Club 12. Cambridge, Massachusetts. Diamond, J.M. 1973. The distributional ecology of New Guinea birds. Science 179: 759–769. Flannery, T.F. 1995a. Mammals of New Guinea. Reed Books, Chatswood, NSW, Australia. Flannery, T.F. 1995b. Mammals of the Southwest Pacific and Moluccas. Reed Books, Chatswood, NSW, Australia. Frith, C.B., and B.M. Beehler. 1996. The Birds of Paradise. Oxford University Press, Oxford. Gressitt, J.L. (ed.). 1982. Biogeography and Ecology of New Guinea. W.R. Junk, The Hague. Heads, M. 2001a. Regional patterns of biodiversity in New Guinea plants. J. Linn. Soc. London. 136: 67–73. Heads, M. 2001b. Birds of paradise and bowerbirds: regional levels of biodiversity in New Guinea and correlations with terrane tectonics. J. of Zool., London 255: 331–339.

................. 16157$

$CH8

02-08-07 10:26:25

PS

PAGE 205

206 / bruce m. beehler Heads, M. 2002. Regional patterns of biodiversity in New Guinea animals. J. Biogeogr. 29: 285–294. Mayr, E. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Massachusetts. Parsons, M.J. 1999. The Butterflies of Papua New Guinea: Their Systematics and Biology. Academic Press, London. Pigram, C.J., and P.J. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. BMR J. Austr. J. Geology and Geophysics 10: 193–212. Pratt, T.K. 1982. Biogeography of birds in New Guinea. Pp. 815–837 in Gressitt, J.L. (ed.) Ecology and Biogeography of New Guinea. W.R. Junk, The Hague. Supriatna, J. (ed.). 1999. The Irian Jaya Biodiversity Conservation Priority-setting Workshop. Conservation International, Washington, D.C. van Welzen, P.C., H. Turner, and M.C. Roos. 2001. New Guinea: a correlation between accreting areas and dispersing Sapindaceae. Cladistics 17: 242–247.

................. 16157$

$CH8

02-08-07 10:26:25

PS

PAGE 206

2.5. Freshwater Biogeography of Papua dan a. polhemus and gerald r. allen e l an e s i an f r e sh w a t er b i o ta s appear to be characterized by foci of high endemicity clustered around tectonic provinces within individual large islands, such as New Guinea (D. Polhemus 1993; D. Polhemus and J. Polhemus 1998), or on geologically allied groups of smaller islands, such as the Louisiades and Solomons (D. Polhemus and J. Polhemus 2004). In lotic systems (see Definitions of Limnological Terms and Units, in Chapter 5.5, for definitions of terms), such as streams and rivers, this endemicity frequently displays a marked turnover in species elements along the length of individual catchments, linked to segregation of individual species by altitude, water temperature, substrate, bed profile, and terminal reach salinity gradients (D. Polhemus et al. 1992; J. Polhemus and D. Polhemus 1996; D. Polhemus and J. Polhemus 2001; Chapter 5.5). By contrast, lentic systems often harbor suites of localized endemic species centered around individual lakes or wetland complexes (Allen 1991). Polhemus (1993), in an analysis prepared for the Papua New Guinea Conservation Needs Assessment, defined 29 areas of freshwater endemism within New Guinea and closely adjacent islands, based on the distributions of certain aquatic insect groups, primarily aquatic true bugs (Heteroptera), damselflies (Zygoptera), and whirlygig beetles (Gyrinidae). These hypotheses of regional endemism were further refined for western New Guinea by D. A. Polhemus, G. A. Allen, and D. Wowor as part of the Irian Jaya Biodiversity Conservation Priority-setting Workshop, held in Biak, Indonesia in January 1997. By utilizing detailed vegetation maps that accurately depicted the extent of mangroves and lowland swamp forest, and by integrating additional data from freshwater fishes and crayfishes, this group was able to produce a detailed map of freshwater endemism for Irian Jaya (now Papua Province), with unit boundaries more precisely defined; this map was published in 1998 as an inset on the back of the larger conservation planning map resulting from that workshop. The results of this re-analysis indicated that the 1993 analysis of Polhemus was to a large degree accurate, but that certain units within New Guinea, particularly in the central mountain chain, had been too broadly defined and ought to be subdivided. In addition, as noted above, the precise boundaries of certain units were also modified based on additional faunal and vegetational data. Since that time, further freshwater faunal surveys in eastern New Guinea and offshore islands, funded in part by Conservation International, have provided a wealth of new data from both the Papuan Peninsula and the D’Entrecasteaux, Louisiade, and Marshall Bennett island groups. As with western New Guinea, these surveys have generally validated the areas of freshwater faunal endemism proposed by Polhemus (1993), but have also forced refinement of unit boundaries and dic-

M

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

207

................. 16157$

$CH9

04-02-07 11:57:20

PS

PAGE 207

208 / d a n a . p o l h e m u s & ge r a l d r. allen

tated recognition of certain new areas of endemism such as Rossel, Tagula, and Woodlark islands that were not treated in previous analyses due to a lack of information. As a result, the current analysis presented in this chapter recognizes 40 areas of freshwater endemism on New Guinea and nearby islands, grouped into 6 broad regions, with many of these endemic areas consisting of single islands or island groups (Table 2.5.1). In addition, 12 lacustrine subunits, comprising individual lakes or lake complexes with distinctive endemic biota, are recognized nested within the classification above (see Table 5.5.1). This analysis differs from the two previous analyses mentioned above in that it omits the Bismarck Archipelago, which will be dealt with subsequently after better freshwater data becomes available from surveys planned in the next several years. Areas of endemism as treated herein refer to regions within New Guinea which contain assemblages of endemic species that appear on the basis of current knowledge to display similarly circumscribed distributions. These areas of endemism are considered to be equivalent to nested sets, with larger areas often containing smaller distinctive subdivisions within them. These areas of endemism are outlined in Table 2.5.1, shown on Figures 2.5.1 and 2.5.2, and described in the text below; their defining faunal components are listed in Appendix 2.5.1. It must be stressed that the areas of endemism defined herein apply to freshwater aquatic organisms only, and may not be congruent with those exhibited by other groups of plants and animals. The approach of defining areas of endemism on the basis of congruent species distributions was used in previous reports dealing with Sulawesi (J. Polhemus and D. Polhemus 1988, 1990). In those studies it was found that the single island of Sulawesi could be viewed as at least five separate islands in terms of the distribution of its freshwater biota. The situation in New Guinea is similar but even more complex, and complicated to a degree by the island’s large size and complex history of geological assembly, coupled with a continuing absence of faunal survey data from many key regions. As a result it has been difficult to ascertain the definitive contact zones between many of the areas of endemism defined herein, and the boundaries depicted on Figure 2.5.1 in particular should be viewed as speculative in many cases, and open to further refinement as more detailed distributional data becomes available.

Groups Used in Defining Areas of Freshwater Biotic Endemism in New Guinea Three major groups of organisms have proven particularly useful for defining areas of freshwater endemism in Papua, and New Guinea as a whole: freshwater fishes, crayfishes, and aquatic insects. All have diverse and regionally distinctive species radiations within New Guinea. Because these groups have all had individual biogeographic histories in the region and have received differing degrees of attention by collectors, the distributions of their constituent species and the areas of endemism they occupy are not strictly congruent. None of the groups used in

................. 16157$

$CH9

02-08-07 10:26:35

PS

PAGE 208

Freshwater Biogeography of Papua / 209

Table 2.5.1. Areas of freshwater endemism as defined by freshwater fishes, crayfish, and selected aquatic insect groups Region 1: Raja Ampat Islands Area 1. Waigeo Area 2. Batanta Area 3. Misool Region 2: Vogelkop and Bomberai Peninsulas Area 4. Vogelkop Lowlands Area 5. Vogelkop Highlands Area 6. Vogelkop Anticlines Area 7. Fakfak and Kumawa Mountains Region 3: North Coast Ranges and Valleys and associated offshore islands Area 8. Biak-Supiori Area 9. Yapen Area 10. Northwest Papuan Coastal Lowlands Area 11. Van Rees and Foja Mountains Area 12. Cyclops Mountains Area 13. Bewani, Torricelli, and Prince Alexander Mountains Area 14. Adelbert, Finisterre, and Saruwaged Mountains Area 15. Mamberamo Basin Area 16. Sepik-Ramu-Markham Basin Region 4: Central Mountain Ranges Area 17. Mamberamo Foreland Area 18. Sepik-Ramu Foreland Area 19. Weyland Mountains Area 20. West Papuan Central Highlands Area 21. East Papuan Central Highlands Area 22. Morobe Highlands Area 23. Arafura Foreland Area 24. Trans-Fly Foreland Area 25. Papuan Gulf Foreland Region 5: Southern Coastal Lowlands and associated islands Area 26. Arafura Coastal Lowlands Area 27. Trans-Fly Coastal Lowlands Area 28. Papuan Gulf Coastal Lowlands Area 29. Aru Region 7: Papuan Peninsula and associated offshore islands Area 30. South Papuan Peninsula Foreland Area 31. Owen Stanley Mountains Area 32. Popondetta Foreland Area 33. Cape Nelson Peninsula Area 34. Cloudy Mountains Area 35. D’Entrecasteaux Islands Area 36. Basilaki Island Area 37. Misima Island Area 38. Tagula Island Area 39. Rossel Island Area 40. Woodlark Island

................. 16157$

$CH9

02-08-07 10:26:36

PS

PAGE 209

210 / d a n a . p o l h e m u s & ge r a l d r. allen

Figure 2.5.1. Areas of freshwater endemism on New Guinea and nearby island groups. For explanations of area definitions see text. Certain areas of endemism on offshore island groups occur beyond the boundaries of this figure and are depicted in Figure 2.5.2. Numbered areas are: 2. Batanta; 3. Misool; 4. Vogelkop Lowlands; 5. Vogelkop Highlands; 6. Vogelkop Anticlines; 7. Fakfak and Kumawa Mountains; 8. Biak-Supiori; 9. Yapen; 10. Northwest Papuan Coastal Lowlands; 11. Van Rees and Foja Mountains; 12. Cyclops Mountains; 13. Bewani, Torricelli, and Prince Alexander mountains; 14. Adelbert, Finisterre, and Saruwaged mountains; 15. Mamberamo Basin (Meervlakte); 16. Sepik-Ramu-Markham Basin; 17. Mamberamo Foreland; 18. Sepik-Ramu Foreland; 19. Weyland Mountains; 20. West Papuan Central Highlands; 21. East Papuan Central Highlands; 22. Morobe Highlands; 23. Arafura Foreland; 24. Trans-Fly Foreland; 25. Papuan Gulf Foreland; 26. Arafura Coastal Lowland; 27. Trans-Fly Coastal Lowland; 28. Papuan Gulf Coastal Lowlands; 29. Aru; 30. South Papuan Peninsula Foreland; 31. Owen Stanley Mountains; 32. Popondetta Foreland; 33. Cape Nelson Peninsula; 34. Cloudy Mountains; 35. D’Entrecasteaux Islands. this analysis contain species representatives in all of the areas of endemism defined, while in other cases a single regionally endemic species may occur across several areas of endemism. Even so, the observed patterns of distribution display remarkable similarity throughout the various groups studied, and indicate that the endemic areas they define are likely to pertain for the remaining elements of the island’s aquatic biota.

fishes Data for freshwater fishes has been derived primarily from Allen (1991, 1996, 2003a,b), Allen et al. (2000), and from additional unpublished collection records compiled by Allen. Four genera in particular proved to have large numbers of insular species with restricted distributions that were useful in defining areas of endemism: Melanotaenia, Mogurnda, Allomogurnda, and Hephaestus. These genera

................. 16157$

$CH9

02-08-07 10:26:53

PS

PAGE 210

Freshwater Biogeography of Papua / 211

Figure 2.5.2. Offshore areas of freshwater endemism in the New Guinea region. For explanation of area definitions and numbering see text. were particularly important because they contain species endemic to single lake basins, and therefore define areas of lacustrine endemism not well depicted by crayfish or aquatic insects (see Table 5.5.1).

crayfishes Data for New Guinea crayfishes were obtained from the publications of Holthuis (1939, 1950, 1956, 1958, 1982, 1986). The one group of particular interest was the genus Charax, which contains a set of 14 species that unambiguously define areas of endemism south of the island’s Central Divide, and contains certain lacustrine endemics, particularly in the Paniai Lakes and Lake Kutubu (see Table 5.5.1). In addition, Bott (1974) reported three genera of freshwater crabs containing 13 species from New Guinea, but the range of collection sites was relatively limited, therefore making it difficult to infer areas of endemism.

aquatic insects Although much additional work remains to be done, reliable distributional data is now emerging for certain groups of aquatic insects in New Guinea, including dragonflies and damselflies (order Odonata, suborders Anisoptera and Zygoptera, respectively), whirlygig beetles (order Coleoptera, family Gyrinidae) and water bugs (order Heteroptera). Information for Zygoptera was extracted from the publications of Lieftinck (1932, 1933, 1935, 1937, 1938, 1949a,b, 1955a,b, 1956a,b, 1957, 1958, 1959a,b, 1960, 1963); information for Gyrinidae from the works of Brinck (1976, 1981, 1983, 1984); and information for Heteroptera primarily from the works of Andersen (1975, 1989a,b), Lansbury (1966, 1968, 1972, 1973, 1993,

................. 16157$

$CH9

02-08-07 10:27:04

PS

PAGE 211

212 / d a n a . p o l h e m u s & ge r a l d r. allen

1996), D. Polhemus and J. Polhemus (1985, 1986a,b, 1989, 1997, 2000a,b,c, 2001, 2004), J. Polhemus and D. Polhemus (1987, 1991, 1993, 1994a,b, 1995, 1996, 2000, 2001, 2002), and Todd (1955, 1959). Species ranges were plotted for taxa in many genera within the above groups, searching for patterns of congruent circumscribed distribution. An attempt was made to include genera from as many different families as possible, so as to compare patterns among many lineages with separate evolutionary histories. On the basis of these range plots, areas of endemism were then defined that were at a minimum supported by species in several genera. The species occurring within each area of endemism are listed in Appendix 2.5.1. Certain areas on New Guinea and nearby islands have not been treated within the context of this analysis due to lack of information; these include Long Island, Karkar Island, the Trobriand Islands, and Numfoor Island.

Areas of Freshwater Endemism on New Guinea and Surrounding Islands Previous workers who have discussed the biogeography of aquatic organisms within New Guinea have identified four large endemic regions: Vogelkop, the north coast ranges, the central highlands, and the southern lowlands (Lieftinck 1938, 1949; Allen 1991). These areas generally correspond to the broad scale physiographic divisions within New Guinea as a whole and are relatively self-evident. The present study indicates that these four divisions are indeed valid, but contain many other smaller areas of endemism within them. These smaller areas are depicted in Figures 2.5.1 and 2.5.2, outlined in Table 2.5.1, and discussed in further detail below. The numbers in the following discussion match those assigned to units on Figures 2.5.1 and 2.5.2, in Table 2.5.1, and in Appendix 2.5.1; the area names reflect characteristic geographical or geomorphological features found within them. In the central section of New Guinea there is a clear altitudinal segregation of biota, and this has been accommodated by defining large lowland, foreland, and mountain faunal units. The general elevational ranges for these units are: lowland  0–50 m; foreland  50–1,200 m; highland  above 1,200 m. Although similar elevational segregation is also frequently seen in many of the smaller units defined herein for the New Guinea north coast ranges and offshore islands, such smaller mapping units have not been altitudinally subdivided for the sake of utility in the context of the current chapter.

region 1: raja ampat islands Area 1. Waigeo The large island of Waigeo, lying on the Papuan shallow water platform northwest of the Vogelkop, represents an ophiolite exposure and supports endemic taxa in nearly all groups examined.

................. 16157$

$CH9

02-08-07 10:27:06

PS

PAGE 212

Freshwater Biogeography of Papua / 213

Area 2. Batanta A small, narrow, but high island separated from Salawati along a narrow strait formed by the submarine trace of the Sorong Fault. It supports endemic taxa in several genera of aquatic Heteroptera. The mountains of northern Salawati Island may also be assignable to this unit, but given that the intervening Sagewin Strait is a major biogeographic boundary for birds, this is a tentative hypothesis that requires verification via future field surveys. Area 3. Misool The island of Misool lies on the Papuan continental shelf, and is primarily composed of upwardly deformed continental shelf limestones correlative to those of the Fakfak and Kumawa mountains on mainland New Guinea. Endemic taxa occur here in certain groups, while in other cases species are shared with the Vogelkop Lowlands to the east or with the central Moluccas (Ambon, Seram, Buru) to the west.

region 2: vogelkop and bomberai peninsulas Area 4. Vogelkop Lowlands The southern coastal lowlands of the Vogelkop Peninsula from Salawati Island southeastward to Etna Bay, including the low central portion of the Bomberai Peninsula. This is an area of endemism for Zygoptera and certain aquatic Heteroptera (Hydrometridae, Veliidae). Most of southern Salawati Island also falls within this unit; the Wagon Mountains of extreme northern Salawati appear, on the basis of limited aquatic insect surveys, to be provisionally allied to Batanta (Area 2), but are left unclassified in the current analysis. Area 5. Vogelkop Highlands This area is defined as including both the Arfak and Tamrau mountains in the northern half of the Vogelkop Peninsula. The Tamrau Mountains contain a core of Paleozoic basement and represent a detached fragment of the Australian craton. Endemic taxa occur in many groups in this highland area. This unit also contains the Ayamaru Lakes, which support at least four endemic fishes. Area 6. Vogelkop Anticlines This unit consists of the limestone anticlines of the Lengguru Fold Belt in the Bird’s Neck region, from the Jakati River southeastward to Etna Bay. These anticlines are steeply folded, forming sharp ridges with much karst terrain and poor integration of drainage. Several lakes of fluctuating size occur in the area behind Kaimana. The uplands of this unit form a transitional corridor between the Vogelkop Highlands and the mountains in the main body of New Guinea, and the lakes support numerous endemic fish species as well as an endemic genus (Pelangia). The Wandammen Peninsula has also been included in this unit for the present,

................. 16157$

$CH9

02-08-07 10:27:06

PS

PAGE 213

214 / d a n a . p o l h e m u s & ge r a l d r. allen

but this mountainous sliver of Australian craton will likely prove to be yet another discrete area of aquatic endemism once adequate faunal surveys are conducted Area 7. Fakfak and Kumawa Mountains These ranges are large limestone anticlines of upwardly deformed continental shelf limestone along the western margin of the Bomberai Peninsula. Both ranges are no older than Pliocene, and represent recently uplifted montane islands which have developed a limited assemblage of endemic aquatic insect species.

region 3: north coast ranges and valleys, and associated offshore islands Area 8. Biak-Supiori This nearly connected island pair lies off the Papuan continental platform, and was not connected to the main body of the island during the Pleistocene. Biak is mostly covered by Quaternary reef limestones, but Supiori, with greater elevation, contains exposures of andesitic island arc volcanics. Sampling for aquatic insects here has been relatively poor, but the presence of endemic species is known among the aquatic Heteroptera in the Gerridae and Notonectidae. Area 9. Yapen This island is a fault sliver rifted from the Van Rees Mountains on the main body of New Guinea. Its fauna is allied to the northern coastal ranges of Irian Jaya, but supports endemic species of Zygoptera and three endemic fishes. Area 10. Northwest Papuan Lowlands The Korime and Tami river basins at elevations below 400 meters appear on the basis of present sampling to be an area of faunal endemism. Future surveys are likely to reveal that this is an artificial picture created by a preponderance of sampling in the Jayapura area, and that the fauna characteristic of this area is in fact more widespread throughout the northern coastal lowlands of Indonesian New Guinea. Area 11. Van Rees and Foja Mountains This area covers the northern coastal ranges of Indonesian New Guinea, north of the Mamberamo Basin. It is poorly explored and very little collecting of its fishes or aquatic insects has been accomplished, but the few samples at hand indicate that the area supports endemic species in some groups, and is allied to the Bewani, Torricelli, and Prince Alexander mountains further to the east. This unit is bisected by the deep north-to-south gorge of the Mamberamo River, which may warrant its subsequent subdivision into two separate units. The unit as defined here includes the hill country south of Lake Sentani; this is the only part of the unit that has been surveyed to any degree. Certain regional endemics in this latter area are also shared with the Mamberamo Foreland (Area 17).

................. 16157$

$CH9

02-08-07 10:27:06

PS

PAGE 214

Freshwater Biogeography of Papua / 215

Area 12. Cyclops Mountains This is an extremely steep-sided mountain range immediately west of Jayapura, representing an accreted ophiolite terrane. Endemic species are known in many groups, including Heteroptera (Gerridae, Naucoridae) and Zygoptera. This is the only area in New Guinea from which Plecoptera have been collected. Area 13. Bewani, Torricelli, and Prince Alexander Mountains This area is the northern coastal mountain block lying between the Papua New Guinea border and the mouth of the Sepik River. This is a portion of an accreted island arc, and contains endemic species of Heteroptera in the families Naucoridae and Gerridae. As defined in this report, this unit also contains the Mount Bougainville Massif of the PNG-Indonesia border region. Area 14. Adelbert, Finisterre, and Saruwaged Mountains (Huon Peninsula) This northern coastal mountain block running from east of the Sepik River delta to the tip of the Huon Peninsula corresponds to the Adelbert-Finisterre Terrane of tectonic geologists, and is considered to be an accreted sector of a formerly separate island that was sutured to New Guinea in the Late Miocene to Pliocene (Hill and Hall 2003). As a result of their history as a separate land mass, these ranges contain many endemic species of Odonata and Heteroptera. Area 15. Mamberamo Basin This is the large structural basin drained by the Rouffaer and Idenberg (now Taritatu and Tariku) rivers, which are the upper branches of the Mamberamo. This area is very poorly surveyed for aquatic biota, but is known to support certain endemic fishes and damselflies. Area 16. Sepik-Ramu-Markham Basins These large structural basins lie behind the north coastal ranges in Papua New Guinea, and are drained by the Sepik, Ramu, and Markham rivers. The lowland fauna of this region includes endemic trepobatine Gerridae and at least 11 endemic fishes.

region 4: central mountain ranges Area 17. Mamberamo Foreland The northern foothills of the central mountains at elevations between 50 and 1,200 meters, from the Wamma River in the west to the Sepik-Mamberamo divide in the east. Many apparently endemic Zygoptera are known from this region due to collections made by the Third Archbold Expedition in 1939. Area 18. Sepik-Ramu Foreland The northern foothills of the central mountains at elevations between 50 and 1,200 meters, from the Sepik-Mamberamo divide in the west to the rim of the Bulolo

................. 16157$

$CH9

02-08-07 10:27:07

PS

PAGE 215

216 / d a n a . p o l h e m u s & ge r a l d r. allen

River basin in the east. This area has been surprisingly poorly surveyed for aquatic biota, but is known to harbor an endemic fish species. Area 19. Weyland Mountains This is a small, rugged mountain range lying slightly to the northwest of the western terminus of the central mountain ranges in Indonesian New Guinea, and having a separate geological origin. Limited surveys indicate that the aquatic insect biota of this range is different from that of the geographically proximate central ranges. Area 20. West Papuan Central Highlands This area includes the Central Dividing Range from the Wapoga River basin in the west to the headwaters of the Fly River in the east, at elevations above 1,200 meters. The boundary between this unit and the Vogelkop Anticlines has been plotted southward along the Wamma River and then across to Etna Bay. In the northwest the boundary between this unit and the Foja and Van Rees Mountains has been plotted along the upper course of the Owa River, but should be considered extremely uncertain, as this is basically unexplored terrain. This unit contains the highest mountains in New Guinea, and has extensive exposures of karst terrain. The continuous area of extremely high elevation running east to west down the center of the range may represent still another area of freshwater endemism, but present surveys are inadequate to establish this. Numerous scientific expeditions have traversed various parts of this unit, demonstrating marked endemism in Zygoptera and aquatic Heteroptera (Naucoridae, Corixidae, Veliidae). This unit includes the Paniai Lakes, which contain an endemic species of crab and fish, as well as an apparent crayfish species flock consisting of eight species. Area 21. East Papuan Central Highlands This area includes the Central Dividing Range between the upper Sepik and Fly rivers in the west and the Bulolo River in the east, containing the headwaters of the Kikori and Purari rivers at elevations above 1,200 meters. This is a complex uplift, with several well separated areas of extremely high terrain centered around peaks such as Mt Wilhelm and Mt Giluwe, and contains an extensive exposure of uplifted Paleozoic basement in the Kubor Anticline. Despite its topographic and geological diversity, the present surveys of aquatic insects indicate that it forms a single area of faunal endemism, and it is thus treated as an undivided unit in the current report. Survey work has been extensive in this region, perhaps accounting for the apparently widespread distribution of its fauna, which includes endemic Naucoridae and Veliidae. Area 22. Morobe Highlands This area includes the mountains east of the Bulolo River where the central ranges abut the western terminus of the Owen Stanley Range, including the headwaters of the Wampit and Watut rivers. This unit has also had extensive survey work due

................. 16157$

$CH9

02-08-07 10:27:07

PS

PAGE 216

Freshwater Biogeography of Papua / 217

to the presence of the Wau Ecology Institute. Its aquatic insect fauna shows notable differences from that of the East Papuan Central Highlands (Area 21), with endemic Gerridae and Naucoridae. As currently defined, this unit also includes the Herzog and Bowutu mountains, which are of disparate geological origin. Future survey work may show that this unit should be subdivided to accommodate endemism in these tectonic terranes. Area 23. Arafura Foreland This area is the southern foothills of the central mountains at elevations between 50 and 1,200 meters, between the Uta River in the west and the Lorentz River in the east. This unit, which is strongly indicated on the basis of both freshwater fish and aquatic insect data, includes the middle reaches of the Ajkwa and Lorentz Rivers, which have been subject to detailed biological surveys. Area 24. Trans-Fly Foreland This area includes the basins of the Digul, Fly, Aramia, and Turama rivers at elevations between 50 and 1,200 meters. Most surveys have concentrated in the vicinity of Kiunga, on the Fly, and have revealed a diverse aquatic biota with a different species composition from that seen in the Kikori and Purari basins to the east or the Arafura Foreland to the west. Area 25. Papuan Gulf Foreland This area includes the basins of the Kikori, Purari, Vailala, and Lakekamu rivers at elevations between 50 and 1,200 meters. This unit includes the elevated karst terrain of the Papuan Plateau, lying to the west of the Kikori River. Recent intensive surveys of this remote area have revealed a rich and distinctive fauna, with many endemic Heteroptera species in the families Gerridae and Veliidae. This unit also includes Lake Kutubu, with its diverse assemblage of endemic fishes.

region 5: southern coastal lowlands and associated continental shelf islands Area 26. Arafura Coastal Lowlands These are the southern coastal lowlands between Uta and the mouth of the Mappi River at elevations below 50 meters, which corresponds roughly to the head of tidal influence in these systems (the actual upper limit of tidal influence in fact lies somewhat lower, but the 50 m contour was chosen for practicality in mapping based on available charts). This area contains more areas of swamp than the slightly elevated Trans-Fly lowlands to the east, and on the basis of limited aquatic insect surveys appears to support a slightly different fauna. The eastern boundary of this unit has been set just west of the Mappi River, but this is a hypothetical alignment based on the mapped extent of palustrine habitats in the region.

................. 16157$

$CH9

02-08-07 10:27:07

PS

PAGE 217

218 / d a n a . p o l h e m u s & ge r a l d r. allen

Area 27. Trans-Fly Coastal Lowlands This area covers the extensive lowland basins of the Digul, Fly, Aramia, and Turama rivers lying between sea level and 50 meters. The eastern boundary of this unit is formed by the abrupt scarp of the Darai limestone plateau lying between the Turama and Kikori Rivers. The western boundary is poorly constrained, but has been mapped along the margin of the extensive palustrine habitats occurring immediately west of the Mappi River, which shares a common terminus with the Digul. This area of large, low gradient rivers and creeks supports a diverse and distinctive aquatic biota, including endemic fishes, crayfish, and aquatic insects. The area shares many faunal elements with northern Australia, indicative of a former land connection. Area 28. Papuan Gulf Coastal Lowlands This area is the southern coastal lowlands from the delta of the Kikori River eastward to Kerema. The boundaries of this unit are basically defined along the 50 meter contour in the lower Fly, Kikori, and Purari river basins, near the heads of the river deltas (see comments under Area 26). The area supports endemic species of Zygoptera, and endemic Heteroptera in the families Veliidae, Hydrometridae, and Gerridae (Trepobatinae). Area 29. Aru This is a set of low limestone islands lying on the Papuan continental platform. The fauna supports a few endemic species, but also shows strong alliances to the Arafura coastal lowlands and Vogelkop lowlands. Further collections in all these areas may show that the Aru biota is merely an isolated segregate of that occurring in the Arafura coastal lowlands (Area 26).

region 6: papuan peninsula and associated offshore islands Area 30. South Papuan Peninsula Foreland This area covers the submontane foreland from Kerema to Milne Bay. It supports endemic Zygoptera and Naucoridae, and several endemic fishes. It corresponds to the southern sector of the formerly separate East Papua Composite Terrane, which was sutured to the main body of New Guinea in the Oligocene (Davies et al. 1997). Area 31. Owen Stanley Range The central mountain chain of the Papuan Peninsula, comprised of uplifted ophiolites and subduction melange. An area of rich endemism for aquatic insects, with many endemic species in all groups. Area 32. Popondetta Foreland This is the large area of submontane foreland north of the Owen Stanley Range, from Cape Vogel to Manau. The area supports ten endemic fishes, and limited

................. 16157$

$CH9

02-08-07 10:27:08

PS

PAGE 218

Freshwater Biogeography of Papua / 219

entomological collections also indicate the presence of endemic Zygoptera and Heteroptera (Naucoridae). Area 33. Cape Nelson Peninsula This area of Late Tertiary volcanic peaks separated from the main body of the Owen Stanley Range by extensive lowlands is characterized by a distinctive suite of endemic aquatic Heteroptera. Area 34. Cloudy Mountains This unit, as defined by a suite of endemic aquatic Heteroptera and Odonata, includes the Cloudy Mountains, the extreme eastern extension of the Owen Stanley Range that forms the East Cape Peninsula, and Normanby Island in the D’Entrecasteaux group. It roughly corresponds to the geological Kutu Terrane of Pigram and Davies (1987). Area 35. D’Entrecasteaux Islands This is a chain of high islands with predominantly metamorphic geology, lying immediately north of the eastern Papuan Peninsula. Recent collections have demonstrated the presence of three endemic fishes, as well as endemic Heteroptera and Zygoptera, but have also shown that this endemism is confined to Goodenough and Fergusson islands, with Normanby Island forming part of the Cloudy Mountains area of endemism instead. Area 36. Basilaki Island This is a small, hilly, heavily forested island lying at the eastern terminus of the chain of islands that extends eastward from the China Strait. Recent surveys have demonstrated the presence of endemic aquatic Heteroptera. Adjacent Sideia Island may also prove to be assignable to this unit. Area 37. Misima Island Misima is a narrow island composed of a low limestone plateau on its eastern half and a very precipitous set of mountains formed from metamorphic rocks ringed by limestone terraces on its western half. The island harbors endemic species of aquatic Heteroptera and Odonata. Area 38. Tagula Island The largest island in the Louisiade Archipelago, hilly throughout with gentle slopes underlain by metamorphic bedrock. The island supports many endemic species of aquatic Heteroptera and Odonata. Area 39. Rossel Island A rugged, heavily forested island composed of metamorphic rocks similar to those of the Owen Stanley Range on New Guinea. The island supports a rich assemblage of endemic aquatic Heteroptera, and shares other regional endemics with Tagula.

................. 16157$

$CH9

02-08-07 10:27:09

PS

PAGE 219

220 / d a n a . p o l h e m u s & ge r a l d r. allen

Area 40. Woodlark Island An isolated island composed primarily of an elevated limestone surface with scattered higher ranges of hills formed from emergent metamorphic bedrock. No endemic fishes are known from the island, but it does support endemic species of aquatic Heteroptera and Odonata.

Tectonic Factors Correlating with Areas of Freshwater Endemism The areas of biotic endemism that we recognize on New Guinea are in many cases centered around Tertiary island arc fragments that have been incorporated into the northern half of the island. These arc fragments are represented by present day exposures of Paleogene volcanics, and Mesozoic to Paleogene ophiolites, which are the respective remnants of the former arc islands themselves, plus the oceanic crust and mantle that underlay them. Such exposures are well depicted on the 1:1,000,000 scale geological maps of Irian Jaya (Dow et al. 1986) and Papua New Guinea (Bain et al. 1972), and the 1:5,000,000 scale tectonic map of Hamilton (1978). Units interpreted as ophiolite include Mesozoic basic and ultrabasic (ultramafic) rocks (serpentinite, peridotite, pyroxenite, and gabbro), and Paleogene to Miocene gabbro, diorite and granodiorite; these ophiolite complexes are identified as such on the maps of Hamilton (1978) and Dow et al. (1986), but not on the Papua New Guinea sheets (although see discussion in Dow 1977). Units interpreted as island arc volcanics include Paleogene basaltic to andesitic lavas and more recent sedimentary formations derived from them. Also included in this assemblage are Paleogene volcanics capped by Quaternary reef limestones (such as those in the Manokwari area, Biak, and the northern Huon Peninsula). Congruence between rock units defined on the Irian Jaya and Papua New Guinea geologic maps has been implied, even though the authors in many cases used different symbols and somewhat different definitions for what appear to be similar stratigraphic units. The ages of the island arc fragments become progressively younger from west to east, being uniformly Paleogene in Irian Jaya but becoming as young as middle Miocene in the northern Papuan Peninsula. This is consistent with Hamilton’s (1979) interpretation that the collision front between the Australian continental margin (now buried beneath the Central Dividing Ranges) and the Tertiary volcanic arc was oblique, beginning first in the west and then proceeding sequentially toward the east (Chapter 2.1). It is now generally accepted that more than one arc was involved in these collisions (Hall 2002; Hill and Hall 2003). A broad band of ophiolite, relatively cohesive in Irian Jaya but much splintered by faulting in Papua New Guinea, borders the northern margin of the Central Dividing Ranges south of the Mamberamo, Sepik, and Ramu river valleys. This band appears to be correlative with the large ophiolite sheet that forms much of the Papuan Peninsula, and linked to a major episode of island arc collision in the Eocene and Oligocene. Other ophiolite exposures linked to a subsequent Miocene collision occur along

................. 16157$

$CH9

02-08-07 10:27:09

PS

PAGE 220

Freshwater Biogeography of Papua / 221

the northern coast, particularly in the Cyclops, Torricelli, and Prince Alexander mountains, which were formed by this separate episode of island collision and onramping. All of these accreted arc terranes support distinct assemblages of endemic aquatic species. Still another episode of island arc collision appears to have occurred in relation to the Adelbert-Finsterre Terrane, consisting of the Adelbert Range and the Huon Peninsula, which possesses a similar Middle Tertiary geology to the islands of the Bismarck Archipelago, most notably New Britain. This terrane most likely represents a former component of the Bismarck Arc that is now accreted onto the northern margin of New Guinea, although the exact paleoposition of this fragment is uncertain since tectonic motions in this region are exceptionally complex, with the crust fractured into numerous microplates (Hamilton 1979; Yan and Kroenke 1994; Hall 2002). The remaining islands of the Bismarck Arc are slowly being pushed onto the northern margin of New Guinea as the collision between the Bismarck Archipelago and the Papuan Peninsula continues, with the floor of the Solomon Sea being simultaneously subducted beneath the Papuan Peninsula on the south and the Bismarck Archipelago on the north. As a result, this system provides a perfect case study of the faunal interactions that occur during arc collision and accretion. Repetitive patterns of shared aquatic insect lineages among these areas are illustrated by Polhemus (1996). Finally, an older geologic feature that has direct bearing on how New Guinea acquired elements of its biota with Gondwanan origins is the basement high of Paleozoic granites that trends northward from the Cape York Peninsula into central Papua New Guinea (Davies 1990; Polhemus and Polhemus 1998). This high represents a northerly extension of the Great Dividing Range, and during the Mesozoic formed a long peninsula or chain of islands extending northward from Australia. The remnants of this basement high are exposed in modern New Guinea in only a few highly scattered locations along the Indonesia-Papua New Guinea border region. The remaining Paleozoic basement associated with this high is covered by the overthrust belts of the Central Dividing Ranges, and any evidence of faunal endemism associated with it has long since been lost. During the Mesozoic and early Tertiary, however, it is likely that the subaereal portions of this high provided tropical refugia for taxa that did not occur elsewhere in the main body of Australia, which lay in colder, more southerly latitudes. The case for such refugia in regard to the waterbug genus Rhagovelia was discussed by Polhemus and Polhemus (1988). Additional areas with extensive exposures of Paleozoic basement are found in western Papua, in the finger-like Wondiwoi Peninsula and in the central core of the Vogelkop Peninsula. These represent old fragments of Australian craton that have been uplifted, and in the case of Vogelkop broken away and rotated, relative to the remainder of the Australian continental platform. During portions of its history, the Vogelkop has been a separate island, and this is reflected by the high degree of species endemism in its biota (Polhemus and Polhemus 1998).

................. 16157$

$CH9

02-08-07 10:27:10

PS

PAGE 221

222 / d a n a . p o l h e m u s & ge r a l d r. allen

Petrological Factors Correlating with Areas of Freshwater Endemism The metamorphic and ophiolite belts of central New Guinea resulting from the early Tertiary island arc collision discussed above are large, relatively cohesive geological units, and this is reflected in their aquatic biotas. In Papua, for instance, the mid-montane aquatic Heteroptera assemblage typical of these exposures is distinctive but broadly distributed, occurring throughout the upper basin of the Mamberamo River. The local dispersal of this biota has no doubt been facilitated by the long river valleys that trend east to west along the strike of the Derewo Fault Zone. Interestingly, the aquatic insect biota of the ophiolite and metamorphic belts emplaced in the Oligocene seems to be considerably richer than that of the highland limestone belt lying immediately to the south, which was elevated later, in the Late Miocene and Pliocene. This is partly due to elevation; the limestones frequently lie at elevations ranging from 3,000 to 4,800 meters, and the jagged crest they form represents an insurmountable cold water barrier to many aquatic taxa, breached only by the Baliem Valley to the east and the Paniai basin to the west. Both of these gaps in the high crest drain to the south, and, as shown by the distribution of crayfishes (Holthuis 1982), have clearly acquired their aquatic biotas from that direction; as such, they have offered little opportunity for north-tosouth faunal interchange. In addition, the limestones produce stream basin characteristics that many aquatic insect species find unsuitable. Streams in these limestone regions usually have marly beds, basic water chemistries, and occupy catchments characterized by high gradients, frequent waterfalls, and poor integration of drainage; in numerous instances these streams disappear into caves in the karst terrain, then reappear great distances away as resurgences in the form of springs, often bursting as high waterfalls from sheer cliffs. Such habitats, though scenically spectacular, are not particularly easy for many aquatic species to colonize, because they present many significant physiographic barriers. In addition, during dry periods, much of the water retreats to underground conduits, and those permanent surface watercourses that do exist are often discontinuous. Stream basins in the ophiolite and metamorphic terrains, by contrast, show very different characteristics. They tend to have relatively pH neutral waters flowing in well sorted beds with high substrate heterogeneity, and occupying well integrated catchments with moderate gradients. This allows the development of a far richer aquatic biota than streams in limestone at similar elevations. The dichotomy is particularly well illustrated by a transect over the Central Ranges just west of Mt Jaya. At Tembagapura, on the south slope at an elevation of 2,000 meters, the mountain streams are high gradient, originate in limestone catchments, and support no aquatic Heteroptera. At Bilogai, on the north slope at 2,000 meters, the mountain streams are moderate gradient, originate in metamorphic catchments, and contain a diverse aquatic Heteroptera biota. This leads to the interesting possibility that, based on samples of aquatic insects, inferences about the petrology of

................. 16157$

$CH9

02-08-07 10:27:11

PS

PAGE 222

Freshwater Biogeography of Papua / 223

a catchment can be made a priori, even when the geological information is not precisely known. Not only are the aquatic insect biotas of ophiolitic and metamorphic terrains richer than those of other areas, they are also regionally distinctive. This is hypothesized to result from the fact that, in addition to providing suitable physical habitats, the ophiolites are in many instances markers for accreted terranes that have traveled considerable distances from their original point of formation, bringing isolated suites of aquatic species with them (Polhemus 1996; Polhemus and Polhemus 1998). Even after an arc terrane has been incorporated into a larger island such as New Guinea, its associated aquatic biota often appears to retain a high fidelity to the accreted block, probably because such accretions are frequently surrounded by topographic basins containing lakes and slow water streams that are barriers to the dispersal of upland species, or because the accreted ophiolitic terranes are often closely juxtaposed with adjacent limestone terranes also uplifted in the course of the accretion (as in the central mountain chain discussed above), which once again serve as effective barriers to dispersal of many aquatic species.

Literature Cited Allen, G.R. 1991. Field Guide to the Freshwater Fishes of New Guinea. Publication No. 9, Christensen Research Institute. Calendar Print Pte. Ltd., Singapore. Allen, G.R. 1995. Rainbowfishes in Nature and the Aquarium. Tetra Press, Melle, Germany. Allen, G.R. 1996. Freshwater fishes of Irian Jaya. Pp. 15–21 in Kitchner, D.J., and A. Suyanto (eds.) Proceedings of the first international conference on eastern IndonesianAustralian vertebrate fauna, Manado, Indonesia, November 22–26, 1994. Allen, G.R. 2003a. Unexplored islands of Papua New Guinea. Fishes of Sahul 17 (2): 926–948. Allen, G.R. 2003b. Allomogurnda, a new genus of gudgeon (Eleotridae) from fresh waters of New Guinea, with descriptions of 7 new species. Fishes of Sahul 17 (3–4): 978–997. Allen, G.R., K.G. Hortle, and S.J. Renyaan. 2000. Freshwater Fishes of the Timika Region, New Guinea. P. T. Freeport Indonesia, Timika. Andersen, N.M. 1975. The Limnogonus and Neogerris of the Old World. Entomologica Scandinavica, Supp. 7: 1–96. Andersen, N.M. 1989a. The coral bugs, genus Halovelia Bergroth (Hemiptera, Veliidae). I. History, classification and taxonomy of species except the H. malaya-group. Entomologica Scandinavica 20: 75–120. Andersen, N.M. 1989b. The coral bugs, genus Halovelia Bergroth (Hemiptera, Veliidae). II. Taxonomy of the H. malaya-group, cladistics, ecology, biology and biogeography. Entomologica Scandinavica 20: 179–227. Bain, J.H.C., H.L. Davies, P.D. Hohnen, R.J. Ryburn, I.E. Smith, R. Grainger, R.J. Tingey, and M.R. Moffat. 1972. Geology of Papua New Guinea. Bureau of Mineral Resources Australia, Canberra, Geology and Geophysics, Canberra. 1:1,000,000 scale map. Bott, R. 1974. Die su¨sswasserdrabben von Neu Guinea. Zoologische Verhandelingen 136: 1–36. Brinck, P. 1976. The Gyrinidae of the Bismarck Archipelago and the Solomon Islands (Coleoptera: Gyrinidae). Entomologica Scandivanica 7: 81–90.

................. 16157$

$CH9

02-08-07 10:27:11

PS

PAGE 223

224 / d a n a . p o l h e m u s & ge r a l d r. allen Brinck, P. 1981. Spinosodineutes (Coleoptera: Gyrinidae) in New Guinea and adjacent islands. Entomologica Scandinavica Supp. 15: 353–364. Brinck, P. 1983. A revision of Rhombodineutes Ochs in New Guinea (Coleoptera: Gyrinidae). Entomologica Scandinavica 14: 205–233. Brinck, P. 1984. Evolutionary trends and specific differentiation in Merodineutes (Coleoptera: Gyrinidae). International Journal of Entomology 26 (3): 175–189. Davies, H.L. 1990. Structure and evolution of the border region of Papua New Guinea. In Carman, G.J., and Z. Carman (eds.) Petroleum Exploration in Papua New Guinea: Proceedings of the First PNG Petroleum Convention, Port Moresby, 12–14th February 1990. PNG Chamber of Mines and Petroleum, Papua New Guinea. Davies, H.L., Perembo, R.C.B., Winn, R.D., and KenGemar, P. 1997. Terranes of the New Guinea orogen. Pp. 61–66 in Hancock, G. (ed.) Proceedings of the Geology Exploration and Mining Conference, Madang. Australian Institute of Mining and Metallurgy, Melbourne. Dow, D.B. 1977. A geological synthesis of Papua New Guinea. Bureau of Mineral Resources Australia, Bulletin 201: 1–41. Dow, D.B., G.P. Robinson, U. Hartono, and N. Ratman. 1986. Geologic map of Irian Jaya, Indonesia. Geological Research and Development Centre, Ministry of Mines and Energy, Bandung. 1:1,000,000 scale map. Hall, R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences 20 (4): 353–431. Hamilton, W. 1978. Tectonic map of the Indonesian region. U.S. Geol. Survey. Misc. Invest. Ser., Map 1-875-D. U.S. Government Printing Office, Washington, D.C. 1:5,000,000 scale map. Hamilton, W. 1979. Tectonics of the Indonesian region. U.S. Geol. Survey Prof. Paper 1078. U.S. Government Printing Office, Washington, D.C. Hill, K.C., and R. Hall. 2003. Mesozoic–Cenozoic evolution of Australia’s New Guinea margin in a west Pacific context. Pp. 265–290 in Hillis, R.R., and R.D. Mu¨ller (eds.) Evolution and Dynamics of the Australian Plate. Geological Society of Australia Special Publication 22 and Geological Society of America Special Paper 372. Holthuis, L.B. 1939. Decapoda Macrura, with a revision of the New Guinea Parastacidae. Zoological Results of the Dutch New Guinea Expedition 1939 3: 289–328. Holthuis, L.B. 1950. Results of the Archbold Expeditions. No. 63. The Crustacea Decapoda Macrura collected by the Archbold New Guinea Expeditions. American Museum Novitates 1461: 1–17. Holthuis, L.B. 1956. Contributions to New Guinea carcinology I. Nova Guinea (new series) 7: 123–137. Holthuis, L.B. 1958. Freshwater crayfish in the Netherlands New Guinea Mountains. SPC Quarterly Bulletin 8: 36–39. Holthuis, L.B. 1982. Freshwater Crustacea Decapoda of New Guinea. Pp. 603–619 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr. W. Junk, The Hague. Holthuis, L.B. 1986. The freshwater crayfish of New Guinea. Freshwater Crayfish 6: 48–58. Lansbury, I. 1966. Notes on the genus Aphelonecta (Hemiptera-Heteroptera: Notonectidae). Pacific Insects 8 (3): 629–632. Lansbury, I. 1968. The Enithares (Hemiptera-Heteroptera: Notonectidae) of the Oriental Region. Pacific Insects 10 (2): 353–442.

................. 16157$

$CH9

02-08-07 10:27:12

PS

PAGE 224

Freshwater Biogeography of Papua / 225 Lansbury, I. 1972. A review of the Oriental species of Ranatra Fabricius (HemipteraHeteroptera: Nepidae). Transactions of the Royal Entomological Society of London 124 (3): 287–341, 262 figs. Lansbury, I. 1973. A review of the genus Cercotmetus Amyot and Serville 1843 (Hemiptera-Heteroptera: Nepidae). Tijdschrift voor Entomologie 116 (5): 83–106, 91 figs. Lansbury, I. 1993. Rhagovelia of Papua New Guinea, Solomon Islands and Australia (Hemiptera-Veliidae). Tijdschrift voor Entomologie 136: 23–54. Lansbury, I. 1996. Two new species of Ciliometra Polhemus (Hem., Gerridae) from Papua New Guinea. Entomologist’s Monthly Magazine 132: 55–60. Lieftinck, M.A. 1932. The dragonflies (Odonata) of New Guinea and neighboring islands. Part I. Descriptions of new genera and species of the families Lestidae and Agrionidae. Nova Guinea 15, Zoology 5: 485–602. Lieftinck, M.A. 1933. The dragonflies (Odonata) of New Guinea and neighboring islands. Part II. Descriptions of a new genus and species of Playcneminae (Agrionidae) and of new Libellulidae. Nova Guinea 17, Zoology 1: 1–66. Lieftinck, M.A. 1935. The dragonflies (Odonata) of New Guinea and neighboring islands. Part III. Descriptions of new and little known species of the families Megapodagrionidae, Agrionidae, and Libellulidae (genera Podopteryx, Argiolestes, Papuagrion, Teinobasis, Huonia, Synthemis, and Protocordulia). Nova Guinea 17: 203–300. Lieftinck, M.A. 1937. The dragonflies (Odonata) of New Guinea and neighboring islands. Part IV. Descriptions of new and little known species of the families Agrionidae (sens. lat.), Libellulidae and Aeshnidae (genera Idiocnemis, Notoneura, Papuagrion, Teinobasis, Aciagrion, Bironides, Agyrtacantha, Plattycantha and Oreaeschna). Nova Guinea (new series) 1: 1–82. Lieftinck, M.A. 1938. The dragonflies (Odonata) of New Guinea and neighboring islands. Part V. Descriptions of new and little known species of the families Libellaginidae, Megapodagrionidae, Agrionidae (sens. lat.) and Libellulidae (genera Rhinocypha, Argiolestes, Drepanosticta, Notoneura, Palaiargia, Papuargia, Papuagrion, Teinobasis, Nannophlebia, Synthemis and Anacordulia). Nova Guinea (new series) 2: 47–128. Lieftinck, M.A. 1949a. Synopsis of the Odonate fauna of the Bismarck Archipelago and the Solomon Islands. Treubia 20, part 2: 319–374. Lieftinck, M.A. 1949b. The dragonflies (Odonata) of New Guinea and neighboring islands. Part VII. Results of the Third Archbold Expedition 1938–1939 and of the Le Roux Expedition 1939 to Netherlands New Guinea (II. Zygoptera). Nova Guinea (new series) 1: 1–82. Lieftinck, M.A. 1955a. Notes on Australasian species of Neurobasis Selys (Odonata, Argiidae). Nova Guinea (new series) 6 (1): 155–166. Lieftinck, M.A. 1955b. Notes on species of Nannophlebia Selys from the Moluccas and New Guinea (Odonata). Zoologische Mededelingen 33 (29): 301–318. Lieftinck, M.A. 1956a. Revision of the genus Argiolestes Selys (Odonata) in New Guinea and the Moluccas. Nova Guinea (new series) 7 (1): 59–121. Lieftinck, M.A. 1956b. Two new Platycnemididae (Odonata) from the Papuan region. Nova Guinea (new series) 7 (2): 249–258. Lieftinck, M.A. 1957. Notes on some argiine dragonflies (Odonata) with special reference to the genus Palaiargia Forster, and with descriptions of new species and larval forms. Nova Guinea (new series) 8 (1): 41–80, 5 pls. Lieftinck, M.A. 1958. A review of the genus Idiocnemis Selys in the Papuan region, with notes on some larval forms of the Platycnemididae. Nova Guinea (new series) 9 (2): 253–292.

................. 16157$

$CH9

02-08-07 10:27:12

PS

PAGE 225

226 / d a n a . p o l h e m u s & ge r a l d r. allen Lieftinck, M.A. 1959a. On the New Guinea species of Ischnura Charpentier and Oreagrion Ris, with special reference to the larval forms and notes on the species of adjacent regions (Odonata, Coenagrionidae). Nova Guinea (new series) 10 (2): 213–240. Lieftinck, M.A. 1959b. New and little known isostictine dragonflies from the Papuan region (Odonata, Protoneuridae). Nova Guinea (new series) 10 (2): 279–302. Lieftinck, M.A. 1960. Three new species of Notoneura Tillyard from western New Guinea. Nova Guinea (new series) 10 (7): 117–126. Lieftinck, M.A. 1963. New species and records of Libellulidae from the Papuan region. Nova Guinea, Zoology 25: 751–780. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. Bureau of Mineral Resources, Journal of Australian Geology and Geophysics 10: 193–211. Polhemus, D.A. 1993. Areas of biotic endemism in New Guinea, as indicated by the distributions of aquatic insect species. In Papua New Guinea Conservation Needs Assessment. U.S. Agency for International Development, Washington, D.C. Polhemus, D.A. 1996. Island arcs, and their influence on Indo-Pacific biogeography. In Keast, A., and S. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Publishing, Amsterdam. Polhemus, D.A., J. Maciolek, and J. Ford. 1992. An ecosystem classification of inland waters for the tropical Pacific islands. Micronesica 25 (2): 155–173. Polhemus, D.A., and J.T. Polhemus. 1985. Naucoridae of New Guinea. I. A review of the genus Nesocricos La Rivers (Hemiptera: Naucoridae) with descriptions of two new species. International Journal of Entomology 27 (3): 197–203. Polhemus, D.A., and J.T. Polhemus. 1986a. Naucoridae of New Guinea. II. A review of the genus Idiocarus Montandon (Hemiptera: Naucoridae) with descriptions of three new species. Journal of the New York Entomological Society 94 (1): 39–50. Polhemus, D.A., and J.T. Polhemus. 1986b. Naucoridae of New Guinea. III. A review of the genus Tanycricos with description of a new species. Journal of the New York Entomological Society 94: 163–173. Polhemus, D.A., and J.T. Polhemus. 1989. Naucoridae (Heteroptera) of New Guinea. IV. A revision of the genus Cavocoris La Rivers, with descriptions of four new species. Journal of the New York Entomological Society 97: 73–86. Polhemus, D.A., and J.T. Polhemus. 1997. A review of the genus Limnometra Mayr in New Guinea, with the description of a very large new species (Heteroptera: Gerridae). Journal of the New York Entomological Society 105: 24–39. Polhemus, D.A., and J.T. Polhemus. 1998. Assembling New Guinea—40 million years of island arc accretion as indicated by the distribution of aquatic Heteroptera (Insecta). Pp. 327–340 in Hall, R., and J. Holloway (eds.) Biogeographical and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Polhemus, D.A., and J.T. Polhemus. 2000a. Naucoridae (Heteroptera) of New Guinea. 6. A revision of the genera Sagocoris and Aptinocoris, with descriptions of new species. Journal of the New York Entomological Society 107 (4): 331–371. Polhemus, D.A., and J.T. Polhemus. 2000b. Additional new genera and species of Microveliinae (Heteroptera: Veliidae) from New Guinea and adjacent regions. Tijdschrift voor Entomologie 143: 91–123. Polhemus, D.A., and J.T. Polhemus. 2000c. New species of Microveliinae (Heteroptera: Veliidae) from the Raja Ampat Islands. Tijdschrift voor Entomologie 143: 279–289. Polhemus, D.A., and J.T. Polhemus. 2000d. A biodiversity survey of aquatic insects in the Ajkwa River basin and adjacent areas, Irian Jaya, Indonesia. Tropical Biodiversity 5: 197–216.

................. 16157$

$CH9

02-08-07 10:27:13

PS

PAGE 226

Freshwater Biogeography of Papua / 227 Polhemus, D.A., and J.T. Polhemus. 2001. A revision of the genus Ptilomera (Heteroptera: Gerridae) on New Guinea and nearby islands. Journal of the New York Entomological Society 109 (1): 81–166. Polhemus, D.A., and J.T. Polhemus. 2004. Two new genera and thirty new species of Microveliinae (Heteroptera: Veliidae) from the East Papua Composite Terrane, far eastern New Guinea. Tijdschrift voor Entomologie 147: 113–189. Polhemus, J.T., and D.A. Polhemus. 1987. The genus Valleriola Distant (Hemiptera: Leptopodidae) in Australia, New Caledonia, and Papua New Guinea with notes on zoogeography. Journal of the Australian Entomological Society 26: 209–214. Polhemus, J.T., and D.A. Polhemus. 1988. Zoogeography, ecology and systematics of the genus Rhagovelia Mayr (Heteroptera: Veliidae) in Borneo, Celebes and the Moluccas. Insecta Mundi 2 (3 and 4): 161–230. Polhemus, J.T., and D.A. Polhemus. 1990. Aquatic Heteroptera of Celebes: regional relationships versus insular endemism. Pp. 73–86 in Knight, W.J., and J.D. Holloway (eds.) Insects and the Rain Forests of South East Asia. Royal Entomological Society of London, London. Polhemus, J.T., and D.A. Polhemus. 1991. Three new species of marine water striders from the Australasian region, with notes on other species (Gerridae: Halobatinae, Trepobatinae). Raffles Bulletin of Zoology 39 (1): 1–13 Polhemus, J.T., and D.A. Polhemus. 1993. The Trepobatinae (Heteroptera: Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 1. Tribe Metrobatini. Entomologica Scandinavica 24 (3): 241–284. Polhemus, J.T., and D.A. Polhemus. 1994a. Four new genera of Microveliinae (Heteroptera) from New Guinea. Tidjschrift voor Entomologie 137: 57–74. Polhemus, J.T., and D.A. Polhemus. 1994b. The Trepobatinae (Heteroptera: Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 2. Tribe Naboandelini. Entomologica Scandinavica 25: 333–359. Polhemus, J.T., and D.A. Polhemus. 1995. The Trepobatinae (Heteroptera: Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 3. Tribe Trepobatini. Entomologica Scandinavica 26: 97–117. Polhemus, J.T., and D.A. Polhemus. 1996. The Trepobatinae (Heteroptera: Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 4. The marine tribe Stenobatini. Entomologica Scandinavica 27: 279–346. Polhemus, J.T., and D.A. Polhemus. 2000. The Trepobatinae (Heteroptera: Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 5. Taxonomic and distributional addenda. Insect Systematics and Evolution 31: 291–316. Polhemus, J.T., and D.A. Polhemus. 2001. The genus Mesovelia Mulsant and Rey in New Guinea (Mesoveliidae: Heteroptera). Journal of the New York Entomological Society 108 (3–4): 205–230. Polhemus, J.T., and D.A. Polhemus. 2002. The Trepobatinae (Gerridae) of New Guinea and surrounding regions, with a review of the world fauna. Part 6. Phylogeny, biogeography, world checklist, bibliography, and final taxonomic addenda. Insect Systematics and Evolution 33: 253–290. Todd, E.L. 1955. A taxonomic revision of the family Gelastocoridae (Hemiptera). University of Kansas Science Bulletin 37 (11): 277–475. Todd, E.L. 1959. The Gelastocoridae of Melanesia (Hemiptera). Nova Guinea (new series) 10: 61–94. Yan, C.Y., and Kroenke, L.W. 1994. A plate tectonic reconstruction of the Southwest Pacific, 0–100 Ma (CD-ROM). Proceedings of the Ocean Drilling Program, Scientific Research 130, Chap. 43.

................. 16157$

$CH9

02-08-07 10:27:13

PS

PAGE 227

228 / d a n a . p o l h e m u s & ge r a l d r. allen

Appendix 2.5.1: Taxa Defining Areas of Endemism Species occurring only in single lakes or lake systems are marked with an asterisk (*). Region 1: Raja Ampat Islands Area 1. Waigeo/Gam Heteroptera Gerridae Ciliometra waigeo Ptilomera waigeo Stygiobates rajana Notonectidae Enithares digitata Veliidae Neusterensifer gamensis Odonata Coenagrionidae Teinobasis prothoracica Platycnemididae Idiocnemis fissidens Protoneuridae Nososticta atrocyana Nososticta evelynae Nososticta erythroprocta Coenagrionidae Palaiargia nasisterna Megapodagrionidae Argiolestes coartans Argiolestes ochrostomus Perciformes Melanotaeniidae Melanotaenia catherinae Area 2. Batanta Heteroptera Veliidae Aegilipsicola insularis Neusterensifer batantana Tarsovelia rajana Perciformes Melanotaeniidae Melanotaenia batanta Melanotaenia sp.

................. 16157$

$CH9

02-08-07 10:27:13

PS

PAGE 228

Freshwater Biogeography of Papua / 229

Area 3. Misool Heteroptera Gerridae Ptilomera misoolensis Veliidae Neusterensifer misoolicus Odonata Protoneuridae Nososticta pyroprocta Coenagrionidae Palaiargia micropsitta Megapodagrionidae Argiolestes pyroprocta Perciformes Melanotaeniidae Melanotaenia misoolensis Region 2: Vogelkop and Bomberai Peninsulas Area 4. Vogelkop Lowlands Heteroptera Gerridae Iobates salawati Notonectidae Enithares n. sp. 1 Odonata Coenagrionidae Palaiargia eos Teinobasis micans Platystictidae Drepanosticta classeni Protoneuridae Nososticta xanthe Megapodagrionidae Argiolestes connectens Argiolestes fontinalis Perciformes Hemiramphidae Zenarchopterus ornithocephala Melanotaeniidae Melanotaenia irianjaya Terapontidae Hephaestus lineatus Gobiidae Glossogobius sp. 5

................. 16157$

$CH9

02-08-07 10:27:14

PS

PAGE 229

230 / d a n a . p o l h e m u s & ge r a l d r. allen

Area 5. Vogelkop Highlands Heteroptera Gerridae Ptilomera arfak Veliidae Tarsovelia arfak Notonectidae Enithares n. sp. 2 Odonata Platycnemididae Idiocnemis inornata Protoneuridae Nososticta dorsonigra Coenagrionidae Ischnura rhodosoma* Palaiargia arses Palaiargia flavovittata Megapodagrionidae Argiolestes convergens Argiolestes ornatus Argiolestes pallidistylus Argiolestes postnodalis Platystictidae Drepanosticta auriculata Drepanosticta inversa Coleoptera Gyrinidae Dineutes (Rhombodineutes) pectoralis avicularis Dineutes (Rhombodineutes) pectoralis pectoralis Perciformes Melanotaeniidae Melanotaenia ajamaruensis* Melanotaenia arfakensis Melanotaenia boesemani* Melanotaenia fredericki Pseudomugil reticulatus* Gobiidae Glossogobius hoesei* Area 6. Vogelkop Anticlines Heteroptera Veliidae Neusterensifer etna Odonata Coenagrionidae Palaiargia starreanum

................. 16157$

$CH9

02-08-07 10:27:14

PS

PAGE 230

Freshwater Biogeography of Papua / 231

Protoneuridae Nososticta lorentzi Nososticta silvicola Coleoptera Gyrinidae Dineutes (Rhombodineutes) silenus Perciformes Melanotaeniidae Melanotaenia angfa Melanotaenia parva* Melanotaenia pierucciae* Melanotaenia lakamora* Pelangia mbutaensis* Terapontidae Variichthys jamoerensis* Eleotridae Mogurnda aiwasoensis* Mogurnda kaifayama Mogurnda magna* Mogurnda mbuta Oxyelyotris altipinna* Area 7. Fakfak and Kumawa Mountains Odonata Coenagrionidae Palaiargia stellata Papuagrion flavipedum Region 3: North Coast Ranges and Valleys, and Associated Offshore Islands Area 8. Biak-Supiori Coleoptera Gyrinidae Dineutes (Rhombodineutes) pectoralis biakensis Heteroptera Gerridae Metrobatopsis insularis Notonectidae Enithares vulgaris Odonata Coenagrionidae Papuagrion insulare Perciformes Gobiidae Lentipes crittersius

................. 16157$

$CH9

02-08-07 10:27:15

PS

PAGE 231

232 / d a n a . p o l h e m u s & ge r a l d r. allen

Area 9. Yapen Heteroptera Gerridae Ptilomera yapenana Odonata Protoneuridae Nososticta wallacii Perciformes Melanotaeniidae Chilatherina pricei Melanotaenia japenensis Eleotridae Allomogurnda sampricei Area 10. Northwest Papuan Coastal Lowlands Odonata Coenagrionidae Teinobasis debeauforti Protoneuridae Nososticta callisphaena Nososticta cyaneura Nososticta rosea cruentata Perciformes Melanotaeniidae Chilatherina sentaniensis* Glossolepis dorityi Glossolepis incisus* Glossolepis pseudoincisus Melanotaenia corona Eleotridae Mogurnda wapoga Gobiidae Glossogobius sp. 10* Area 11. Van Rees and Foja Mountains (including Sentani Hills) Odonata Coenagrionidae Aciagrion tonsillare Palaiargia halcyon Papuagrion corruptum Papuagrion degeneratum Papuagrion laminatum Papuargia stuberi Teinobasis stigmatizans Platycnemididae Arrhenocnemis sinuatipennis

................. 16157$

$CH9

02-08-07 10:27:15

PS

PAGE 232

Freshwater Biogeography of Papua / 233

Perciformes Melanotaeniidae Chilatherina bleheri* Melanotaenia maylandi Area 12. Cyclops Mountains Heteroptera Gerridae Ptilomera cheesmanae Naucoridae Cavocoris bisulcus Veliidae Neusterensifer cyclops Odonata Coenagrionidae Palaiargia charmosyna cyclopica Papuagrion fraterculum Papuagrion rectangulare Papuagrion rufipedum Papuagrion spinicaudum Megapodagrionidae Argiolestes tristis Platycnemididae Idiocnemis nigriventris Coleoptera Gyrinidae Dineutes (Rhombodineutes) helleri stueberi Area 13. Bewani, Torricelli, and Prince Alexander Mountains Heteroptera Gerridae Metrobatoides genitalis Ptilomera wewak Veliidae Neusterensifer cyclops Rhagovelia thysanotos Odonata Coenagrionidae Teinobasis luciae Platycnemididae Idiocnemis chloropleura Coleoptera Gyrinidae Dineutes (Rhombodineutes) helleri helleri Perciformes Melanotaeniidae Chilatherina axelrodi

................. 16157$

$CH9

02-08-07 10:27:15

PS

PAGE 233

234 / d a n a . p o l h e m u s & ge r a l d r. allen

Terapontidae Hephaestus obtusifrons Eleotridae Mogurnda sp. 5 Area 14. Adelbert, Finisterre, and Saruwaged Mountains Heteroptera Gerridae Ptilomera biroi Veliidae Neusterensifer acuminata Odonata Platycnemididae Idiocnemis adelbertensis Idiocnemis huonensis Protoneuridae Nososticta astrolabica Coenagrionidae Palaiargia humida Megapodagrionidae Argiolestes kirbyi Argiolestes montivagans Platystictidae Drepanosticta dendrolagina Coleoptera Gyrinidae Dineutes (Rhombodineutes) pectoralis mesosternalis Dineutes (Rhombodineutes) tetracanthus buergersi Area 15. Mamberamo Basin (Meervlakte) Odonata Coenagrionidae Teinobasis argiocnemis Teinobasis olthofi Perciformes Melanotaeniidae Melanotaenia vanhuerni Gobiidae Eugnathogobius tigrellus Area 16. Sepik-Ramu-Markham Basin Heteroptera Gerridae Ciliometra sepik Perciformes Ariidae

................. 16157$

$CH9

02-08-07 10:27:16

PS

PAGE 234

Freshwater Biogeography of Papua / 235

Ariopsis coatesi Brustiarius nox Hemirhamphidae Zenarchopterus alleni Melanotaeniidae Chilatherina bulolo Glossolepis maculosus Glossolepis wanamensis Syngnathidae Microphis spinachioides Ambassidae Parambassis altipinnis Gobiidae Glossogobius coatesi Glossogobius sp. 14 Region 4: Central Mountain Ranges Area 17. Mamberamo Foreland Heteroptera Gerridae Metrobatoides bifurcatus Odonata Coenagrionidae Palaiargia alcedo Palaiargia charmosyna miniata Palaiargia ceyx ceyx Megapodagrionidae Argiolestes amphistylus Argiolestes aulicus Argiolestes lamprostomus Argiolestes simplex Argiolestes sponsus Podopteryx casuarina Platycnemididae Arrhenocnemis amphidactylus Cyanocnemis aureofrons Lochmaeocnems malacodora Platystictidae Drepanosticta eucera Drepanosticta lepyricollis Protoneuridae Selysioneura ranatra

................. 16157$

$CH9

02-08-07 10:27:16

PS

PAGE 235

236 / d a n a . p o l h e m u s & ge r a l d r. allen

Coleoptera Gyrinidae Dineutes (Rhombodineutes) heurni Dineutes (Rhombodineutes) sinuaticollis Dineutes (Rhombodineutes) tetracanthus tetracanthus Area 18. Sepik-Ramu Foreland Perciformes Terapontidae Hephaestus transmontanus Gobiidae Glossogobius sp. 14 Area 19. Weyland Mountains Heteroptera Gerridae Ptilomera nabire Veliidae Neusterensifer gladius Neusterensifer nabire Perciformes Melanotaeniidae Chilatherina alleni Glossolepis leggetti Melanotaenia rubripinnis Area 20. West Papuan Central Highlands Heteroptera Gelastocoridae Nerthra improcera Nerthra infecta Nerthra monticola Nerthra petila Veliidae Aegilipsicola iriana Neusterensifer iriana Tarsovelia reclusa Odonata Coenagrionidae Archiboldargia gloriosa Archiboldargia mirifica Ischnura ariel Ischnura isoetes Oreagrion armeniacum Oreagrion oreadum Oreagrion xanthocyane Palaiargia ceyx flammula

................. 16157$

$CH9

02-08-07 10:27:16

PS

PAGE 236

Freshwater Biogeography of Papua / 237

Palaiargia eclecta Palaiargia myzomela Papuagrion digitiferum Papuagrion ekari Papuagrion pandanicolum Papuagrion pesechem pesechem Papuagrion pesechem corniculatum Megapodagrionidae Argiolestes pectitus Platycnemididae Torrenticnemis filicornis Platystictidae Drepanosticta dorcadion Coleoptera Gyrinidae Dineutes (Merodineutes) archiboldianus Dineutes (Rhombodineutes) pectoralis occidentalis Perciformes Eleotridae Oxelyotris wisselensis* Decapoda Parastacidae Charax boschmai* Charax buitenijkae* Charax communis* Charax longipes* Charax murido* Charax pallidus* Charax paniaicus* Charax solus* Sundathelphusidae Rouxana roushdyi Area 21. East Papuan Central Highlands Heteroptera Gerridae Ptilomera jimi Notonectidae Enithares n. sp. 3 Odonata Coenagrionidae Ischnura acuticauda Platycnemididae Paramecocnemis stilla-cruoris

................. 16157$

$CH9

02-08-07 10:27:17

PS

PAGE 237

238 / d a n a . p o l h e m u s & ge r a l d r. allen

Coleoptera Gyrinidae Dineutes (Merodineutes) jocosus Dineutes (Merodineutes) priscus Dineutes (Rhombodineutes) pectoralis alticola Perciformes Eleotridae Allomogurnda hoesei Gobiidae Glossogobius brunnoides Area 22. Morobe Highlands Heteroptera Gelastocoridae Nerthra robusta Nerthra stevensi Gerridae Ptilomera morobe Naucoridae Cavocoris minor Cavocoris rotundatus Tanycricos froeschneri Veliidae Aegilipsicola rapida Neusterensifer bowutu Rhagovelia herzogensis Tanyvelia missim Coleoptera Gyrinidae Dineutes (Merodineutes) wauensis Dineutes (Rhombodineutes) pectoralis centralis Perciformes Eleotridae Allomogurnda flavimarginata Area 23. Arafura Foreland Heteroptera Gerridae Calyptobates kamoro Calyptobates kopi Ptilomera timika Stygiobates iweka Perciformes Atherinidae Craterocephalus nouhuysi

................. 16157$

$CH9

02-08-07 10:27:17

PS

PAGE 238

Freshwater Biogeography of Papua / 239

Melanotaeniidae Melanotaenia ogilbyi Plotosidae Oloplotosus mariae Pseudomugilidae Pseudomguil ivantsoffi Pseudomguil paskai Pseudomguil pellucidus Terapontidae Hephaestus habbemai Apogonidae Glossamia timika Eleotridae Bostrichthys strigogenys Mogurnda cingulata Area 24. Trans-Fly Foreland Heteroptera Gerridae Calyptobates simplex Ptilomera kiunga Veliidae Neusterensifer pseudocyclops Perciformes Ariidae Ariopsis taylori Melanotaeniidae Melanotaenia oktediensis Melanotaenia sexlineata Pseudomugilidae Kiunga ballochi Kiunga bleheri Gobiidae Glossogobius sp. 7 Glossogobius sp. 11 Area 25. Papuan Gulf Foreland Heteroptera Gerridae Ptilomera kutubu Ptilomera omo Stygiobates mubi Veliidae Neusterensifer lubu Tarsovelia kikori

................. 16157$

$CH9

02-08-07 10:27:17

PS

PAGE 239

240 / d a n a . p o l h e m u s & ge r a l d r. allen

Odonata Coenagrionidae Teinobasis debeauxi Coleoptera Gyrinidae Dineutes (Rhombodineutes) pectoralis papuanus Perciformes Atherinidae Craterocephalus lacustris Melanotaeniidae Melanotaenia caerulea Melanotaenia herbertaxelrodi* Melanotaenia iris Melanotaenia lacustris* Melanotaenia monticola Melanotaenia mubiensis Melanotaenia pimaensis Plotosidae Oloplotosis torobo* Terapontidae Hephaestus adamsoni* Hephaestus fuliginosus (also N. Australia) Hephaestus komaensis Eleotridae Allomogurnda landfordi Mogurnda furva* Mogurnda kutubuensis Mogurnda maccuneae Mogurnda malsmithi Mogurnda mosa Mogurnda pulchra Mogurnda spilota* Mogurnda variegata* Mogurnda vitta* Oxyeleotris caeca Gobiidae Glossogobius sp. 3 Glossogobius sp. 8 Glossogobius sp. 12 Glossogobius sp. 13 Decapoda Parastacidae Charax papuanus*

................. 16157$

$CH9

02-08-07 10:27:18

PS

PAGE 240

Freshwater Biogeography of Papua / 241

Region 5: Southern Coastal Lowlands, and Associated Continental Shelf Islands Area 26. Arafura Coastal Lowlands Heteroptera Gerridae Ciliometra minajerwi Odonata Coenagrionidae Papuagrion perameles Plagulibasis ciliata Teinobasis fulgens Teinobasis nitescens Perciformes Ariidae Tetranesodon conorhynchus Eleotridae Oxyeleotris stagnicola Terapontidae Hephaestus roemeri Area 27. Trans-Fly Coastal Lowlands Heteroptera Gerridae Ciliometra kiunga Mesoveliidae Mesovelia stysi Veliidae Phoreticovelia rotunda Odonata Coenagrionidae Austroagrion exclamationalis Perciformes Ariidae Arius taylori Eleotridae Mogurnda mogurnda(also N. Australia) Melanotaeniidae Iriatherina werneri (also N. Australia) Melanotaenia macullochi (also N. Australia) Ambassidae Ambassus macleayi (also N. Australia) Denariusa bandata (also N. Australia) Terapontidae Hephaestus raymondi Apogonidae Glossamia narindica

................. 16157$

$CH9

02-08-07 10:27:18

PS

PAGE 241

242 / d a n a . p o l h e m u s & ge r a l d r. allen

Toxotidae Toxotes lorentzi (also N. Australia) Gobiidae Gymnoamblyopus novaeguineae Area 28. Papuan Gulf Coastal Lowlands Odonata Gyrinidae Dineutes chalybeus Heteroptera Gerridae Ciliometra setosa Iobates ivimka Area 29. Aru Odonata Coenagrionidae Teinobasis buwaldi Region 6: Papuan Peninsula and Associated Offshore Islands Area 30. South Papuan Peninsula Foreland Heteroptera Gerridae Ciliometra femorata Ptilomera breddini Naucoridae Aptinocoris fenneri Aptinocoris sogeri Cavocoris ismayi Perciformes Ambassidae Tetracentrum apogonoides Melanotaeniidae Melanotaenia papuae Melanotaenia parkinsoni Melanotaenia sylvatica Terapontidae Hephaestus trimaculatus Gobiidae Lentipes watsoni Area 31. Owen Stanley Mountains Odonata Gyrinidae Dineutes loriae Dineutes macrochirus

................. 16157$

$CH9

02-08-07 10:27:18

PS

PAGE 242

Freshwater Biogeography of Papua / 243

Heteroptera Veliidae Neusterensifer goilala Rheovelia asymmetrica Rheovelia truncata Odonata Megapodagrionidae Argiolestes epihippiatus Argiolestes esuriens Argiolestes luteipes Argiolestes microstigma Argiolestes prothoracicalis Argiolestes saltator Argiolestes saltuarius Argiolestes tenuispina Area 32. Popondetta Foreland Heteroptera Gerridae Ciliometra hirsuta Naucoridae Sagocoris asymmetricus Veliidae Rhagovelia aureospicata Rhagovelia caesius Rhagovelia hirsuta Perciformes Hemirhamphidae Zenarchopterus robertsi Atherinidae Craterocephalus kailolae Pseudomugilidae Pseudomugil connieae Pseudomugil furcatus Chandidae Tetracentrum caudovittatus Tetracentrum honessi Eleotridae Mogurnda lineata Mogurnda orientalis Tateurndina ocellicauda Gobiidae Glossogobius sp. 4

................. 16157$

$CH9

02-08-07 10:27:18

PS

PAGE 243

244 / d a n a . p o l h e m u s & ge r a l d r. allen

Area 33. Cape Nelson Peninsula Heteroptera Veliidae Brechyvelia tufi Neusterensifer tufi Rheovelia fonticola Area 34. Cloudy Mountains Heteroptera Gerridae Ciliometra priori Veliidae Aegilipsicola peninsularis Neusterensifer femoralis Neusterensifer sagarai Rheovelia petrophila Tanyvelia minima Odonata Protoneuridae Selysioneura rangifera Selysioneura rhaphia Area 35. D’Entrecasteaux Islands Heteroptera Veliidae Neusterensifer kula Rheovelia robinae Odonata Megapodagrionidae Argiolestes annulipes Argiolestes armeniacus Platycnemididae Rhyacocnemis sufficiens Protoneuridae Selysioneura arboricola Perciformes Eleotridae Allomogurnda insularis Allomogurnda montana Gobiidae Lentipes venustus Area 36. Basilaki Island Heteroptera Veliidae Rheovelia basilaki

................. 16157$

$CH9

02-08-07 10:27:19

PS

PAGE 244

Freshwater Biogeography of Papua / 245

Odonata Megapodagrionidae Argiolestes n. sp. 1 Area 37. Misima Island Heteroptera Veliidae Neusterensifer misima Rheovelia anomala Odonata Protoneuridae Selysioneura drymobia Area 38. Tagula Island Heteroptera Veliidae Neusterensifer sulcata Tanyvelia tagulana Odonata Megapodagrionidae Argiolestes n. sp. 2 Platycnemididae Idiocnemis leonardi Area 39. Rossel Island Heteroptera Veliidae Neusterensifer yela Odonata Megapodagrionidae Argiolestes n. sp. 3 Area 40. Woodlark Island Heteroptera Veliidae Neusterensifer muyuw Odonata Megapodagrionidae Argiolestes n. sp. 4 Protoneuridae Selysioneura virgula

................. 16157$

$CH9

02-08-07 10:27:19

PS

PAGE 245

2.6. Paleontology of Papua geoffrey s. hope and ken p. aplin o s si l s r ec o r d the former animals and plants that lived in the sea and on land in Papua as it changed from a shallow temperate sea north of Australia to shallow tropical seas and islands that finally coalesced into mountain ranges and accreted isolated land areas such as the Vogelkop (Bird’s Head) Peninsula (Metcalfe 2001; Hall 2001; Pigram and Davies 1997; Quarles van Ufford and Cloos 2005; Chapter 2.1). Fossil plant leaf assemblages have been found in the early Permian Aiduna Formation, which outcrops on the southern flank of the Central Ranges south of Mt Jaya, Waghete, and the Weyland Mountains (Jongmans 1940 Rigby 2001). Ferns (Osmundaceae), Glossopteris, and other seed ferns and gymnosperms (Gigantonoclea) provide a flora which has both Gondwanan and Cathaysian elements suggesting connections from both northern and southern directions. This is the earliest indication of terrestrial habitats in New Guinea. Marine fossils are well known from the late Palaeozoic, Mesozoic, and Tertiary rocks of Papua. Shells of brachiopods occur in Permian rocks (Archbold 1991, 1992, 2001; Table 2.6.1) and show some affinities with northern taxa as well as southern, in agreement with the plant fossils. In the early Permian the South Pole lay on the present location of South Australia and the sea where Papua would form had a paleolatitude of 50–60S. There was considerable interchange and fluctuations from temperate to tropical through the Permian. By the mid-Jurassic, perhaps 160 million years ago, some islands were present north of Gondwana in central Papua New Guinea and the Vogelkop Peninsula. Mollusks, including ammonites, an extinct group of cephalopods resembling Nautilus, are common in the Jurassic limestones and siltstones of old reefs now forming the northern ranges of the Central Highlands (Westermann 1995; Figure 2.6.1). The main range, formed from Miocene and younger limestones, has eroded

F

Table 2.6.1. Early Permian Brachiopod genera of Papua Aktastinian Artinskian Aulosteges Baigendzhinian Callispirina Cancrinella Chonetinella Cleiothyridina Cruricella

Echinalosia Heteralosia Hustedia Linoproductus Neochonetes Neospirifer Rhipidomella Sommeriella Spiriferellina

Stenoscisma Stereochia Stictozoster Stictozoster Streptorhynchus Sulcataria Syringothyris Taeniothaerus

Source: Archbold (1991a).

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

246

................. 16157$

CH10

02-08-07 10:26:43

PS

PAGE 246

Paleontology of Papua / 247

Figure 2.6.1. The Jurassic ammonite Satoceras satoi from the Homejo area. The type specimen is in Japan. Source: Westermann and Callomon (1988).

to expose the older Mesozoic, and ammonites occur, for example near Wamena, Tiom, and Homejo, and at many other locations on the flanks of the mountains (Gerth 1965). The same or related taxa are found in Papua New Guinea and from the Sula Islands (Westermann and Callomon 1988). Carbonate platforms were widespread in the Tertiary and consist of reef complexes, oolitic marls, and deepwater black lime oozes. Hence limestones are widespread, and these now form the highest peaks along the main range. They contain abundant fossils of mollusks, crinoids, bryozoans, and corals. Parts of present day New Guinea emerged in the Oligocene (ca 35 million years ago) but much of this area resubmerged under shallow seas in the early Miocene (20 million years ago), perhaps leaving a few small islands above water (Hall 2001).

Vegetation Plant fossils, mainly pollen and spores, have been studied in cores taken from oil exploration holes in order to provide ages based on the age ranges of recognizable

................. 16157$

CH10

02-08-07 10:27:28

PS

PAGE 247

248 / geoffrey s. h o p e a n d k e n p. aplin

taxa. However, not much information has been published. Morley (2000) has described Eocene pollen from the Waripi Formation on the Vogelkop Peninsula. This includes Australian taxa such as Casuarina (including the fossil genus Triorites harrisii), Myrtaceae, and Euphorbiaceae and no typically Asian taxa, suggesting that a substantial sea barrier still separated New Guinea and Asia at this time. The flora, and the presence of evaporites in the rocks, suggests a semi-arid seasonal climate as might be expected to have prevailed north of Australia in the subtropical high zone. At this time, sites may have been 20 degrees further south than at present (Frakes 1997). While rare Nothofagus grains occur in the Waripi Formation in the Eocene it only becomes common in the mid-Miocene, in good accord with its appearance in Papua New Guinea (Khan 1976). Nothofagus originated in southern Australia and Antarctica and one group, the brassospora group, was dominant across Australia in the Eocene. Its expansion into New Guinea during the Miocene may reflect conditions cooler than present that allowed it to occupy lowland sites (Truswell 1992). The brassospora group became extinct in Australia in the late Pleistocene but five species still remain in New Caledonia while there are eight species in Papua and a further six species in Papua New Guinea (Read et al. 2005). The New Guinea species occur at mean annual temperatures ranging from 10.6–23.5C, with annual precipitation of 1,762–7,733 mm. The genus thus heralds the presence of these cool and humid climates in the region from the late Cenozoic. Nothofagus was unable to disperse further west although gymnosperms such as Agathis, Phyllocladus, and Dacrycarpus did reach Maluku.

Vertebrate Fossils In the late Cenozoic New Guinea was a series of low tropical islands with possible intermittent connections to each other and to Australia. They were also a destination for migrant Asian plants and animals able to cross increasingly narrow water gaps. Although extremely diverse rainforest faunas were present in northern Australia through the Miocene (Archer et al. 1999), only a few lineages of marsupials made it to, or else survived in, New Guinea. There are no Tertiary mammal fossils so far from Papua and our knowledge of this early period is dependent on a small collection of fossils from the Otibanda Formation near Wau, Papua New Guinea, most of them collected during sluicing operations in the Bululo goldfield. The Otibanda fossils are late Pliocene in age, about 2.5–3.1mya (Flannery 1994), and the paleo-environment is thought to be rainforest near sea level. Nearly all of the fossils are from large marsupials, although they include a few small mammal bones and one murid rodent tooth. Four marsupial families are represented: a thylacine (marsupial wolf) quite similar to the modern species Thylacinus cynocephalus; a dasyurid belonging to the extant New Guinean genus Myoictis; three species of browsing kangaroos (family Macropodidae); and three species of the extinct family Diprotodontidae. Among the kangaroos, the two species of Protemnodon are related to Pliocene Australian forms but the third species, Watutia novaeguineae,

................. 16157$

CH10

02-08-07 10:27:28

PS

PAGE 248

Paleontology of Papua / 249

is most similar to Miocene kangaroos from northern Australia. The three diprotodontids were goat- to cow-sized animals and were also browsers. One species, Kolopsoides cultridens, is quite different from the other two and may represent a more ancient New Guinean heritage. This fauna thus seems to have originated from northern Australia at a time when Australia had some rainforest cover. Tentatively, we can identify two Australian waves of migration by the Pliocene and one from Asia as the ancestors of New Guinea’s murine rodents. Pleistocene sea level oscillations provided repeated opportunities for elements of the modern Australian fauna to immigrate across the Arafura land bridge. Unfortunately, the known fossil record of this period remains poor and there is little direct evidence of faunal mixing. Whereas Australian Pleistocene deposits produce the bones of numerous giant marsupials (belonging to four families) and also of giant birds (extinct family Dromornithidae), only one of these lineages is recorded in New Guinea, the diprotodontid subfamily Zygomaturinae. A single fossil dentary fragment, described as Zygomaturus nimborensia, is known from lowland New Guinea (Hardjamasita 1985). The discovery site of this bone, handed in from Nimboran west of Sentani, has not been located so no dating is yet possible. The animal was about cow-sized and may have lived in forest or slightly more open woodlands. Given the extensive karst landforms in the area it is likely that more material may be discovered. At montane altitudes diprotodontids were found in the Pureni peatlands near Tari, Papua New Guinea and named Hulitherium thomasettii (Flannery and Plane 1986). In addition to a small species of cassowary, Casuarius lydekkeri (Rich et al. 1988), crocodilian fossils also occur in this deposit which, at 1,400 m, is above the altitudinal limit for crocodiles today. Uranium series dating of this deposit suggests that it may be around 80,000 years old (Haberle 1998). Several large, montane, browsing kangaroo species (Protemnodon niobe and P. tumbuna) are also known from cave sediments dated to around 25,000 years old in Nombe Cave near Chuave (Flannery et al. 1981); P. tumbuna was probably descended from the Pliocene Protemnodon otibandus. Bones from a kangaroo handed by Yot Murip to a missionary at Kwiyawagi, on the West Baliem River in the central ranges led to the investigation of a cave at an altitude of about 2,900 m at Kelangur (Figure 2.6.2). The cave contained abundant fossilized remains of small mammals alongside bones of two extinct species, a small diprotodontid, Maokopia ronaldii (Figure 2.6.3), and a stout kangaroo, Protemnodon hopeii (Flannery 1992; Hope et al. 1993). Compared to their midmontane relatives, both of the Kelangur species have relatively high-crowned molars suggestive of adaptation to grazing in subalpine grasslands. Glacial conditions at this time would have extended the subalpine grasslands to below the level of the cave, so the fauna would have been living in shrubby grasslands. Gravels in the banks of the West Baliem River also contain Maokopia bone. A carbon date from a peat band in the gravels suggests that this fauna was still flourishing about 40,000 years ago. Small mammal remains from Late Pleistocene sites are usually assumed to be-

................. 16157$

CH10

02-08-07 10:27:29

PS

PAGE 249

250 / geoffrey s. h o p e a n d k e n p. aplin

Figure 2.6.2. Skull of the small diprotodontid marsupial Maokopia ronaldii from the West Baliem River, Kwiyawagi. long to extant species. However, one collection from archeological sites on the Ayamaru Plateau of the Vogelkop Peninsula of Papua contained the bones and teeth of several previously unknown small mammals, including a ringtail possum (Petauroides ayamaruensis) and a tiny striped possum (Dactylopsila kambuayai; Aplin et al. 1999). Another recent study of late Pleistocene archeological remains from the Aru Islands (which were connected to New Guinea during periods of low sea level) led to the discovery of a new kind of peroryctid bandicoot which has yet to be named (Aplin and Pasveer 2005). Numerous Holocene records of faunal change have been obtained from rockshelter and cave sites investigated for archeology in Papua New Guinea but such records are rare so far in Papua. Treeline rockshelters around 4,000 m altitude at Mt Jaya and south of Kwiyawagi yielded bones of a small wallaby, Thylogale christensenii, which seems to have inhabited the scrub and grasslands (Hope 1981; Figure 2.6.4). Radiocarbon dates from hearths at both sites suggest that the animal died out about 3,000 years ago (Hope et al. 1993). Other fauna known to have died out in the Holocene include the thylacine or marsupial wolf, the bones of which occur in several Pleistocene cave sites in Papua New Guinea. Archeological faunas from caves on the Vogelkop Peninsula and on the Aru Islands also give

................. 16157$

CH10

02-08-07 10:27:40

PS

PAGE 250

Paleontology of Papua / 251

Figure 2.6.3. Kelangur Cave, 8 km west of Kwiyawagi, has Late Pleistocene skulls and teeth on a former stream channel.

Figure 2.6.4. Mapala rockshelter, at 4,150 m on the northern slope of Mt Jaya, is the type locality for an extinct Holocene wallaby, Thylogale christensenii. clues about past environments. They show a pronounced altitudinal lowering of montane faunal species during late glacial times on the Vogelkop, and the former presence of open savanna habitats on the Aru Islands at times when they were connected to the main island. The Aru Islands today support lowland rainforest. In both areas, the transition from glacial to post-glacial environmental conditions was accompanied by faunal turnover involving numerous local extinctions, some of which might have been exacerbated by human hunting.

................. 16157$

CH10

02-08-07 10:28:12

PS

PAGE 251

252 / geoffrey s. h o p e a n d k e n p. aplin

Evolutional Directions A curious effect of mountain building in New Guinea, is that while much of the Australian portion of a northward drifting Meganesia went through a drying trend that culminated in Pleistocene aridity, New Guinea’s high mountains and blocking of the equatorial ocean circulation served to buffer the island against aridification. This enabled it to act as a refuge, particularly at altitude, for Gondwanan species with a preference for cool, moist subtropical and temperate environments. Thus a relatively geologically young New Guinea was able to recreate Gondwanan niches which preserve Gondwanan plant taxa such as Dacrydium, Dacrycarpus, and Nothofagus. Pliocene faunas from southern Australia often overlap with modern mountain faunas in New Guinea, for example the 4.6 million year old site at Grange Burn, in western Victoria. Here, the mammal fossils include cuscus-like phalangerid possums, and kangaroos resembling tree-kangaroo and the modern New Guinean genus Dorcopsis, probably living in a habitat of relatively open forest (Rich and Thompson 1982; Flannery et al. 1996). Similarly, the slightly younger Pliocene Otibanda fauna from New Guinea’s Wau Valley has some features in common with late Miocene faunas of Australia (Rich and Thompson 1982). With the exception of the marine and bat faunas, New Guinea has clearly acquired the founders of its present biota with some difficulty, suggesting that barriers and filters have almost always been operating. Thus the New Guinea marsupial fauna, while distinctly related to and derived from that of Australia, is comparatively depauperate due to many marsupial families failing to reach or successfully establish in New Guinea (Chapter 4.10). The highly diverse possum faunas of the Australian Miocene did not gain a foothold presumably because there were marine barriers. The semi-permanent connection to northern Australia through the Pleistocene continued to act as a filter because it supported semi-arid habitats and hence was difficult for rainforest species to traverse. On the other hand, the low founder diversity allowed rapid speciation and adaptive radiations of both the marsupials and the Asian newcomers, the murid rodents. Species and generic diversities in the two major groups of terrestrial mammals are approximately equal in New Guinea, in contrast to the forest regions of Australia where marsupials are overwhelmingly dominant. This contrast suggests that both marsupials and placentals effectively entered New Guinea into empty niches.

Literature Cited Anderson, C. 1937. Palaeontological notes no. 4: fossil marsupials from New Guinea. Records of the Australian Museum 20: 73–76. Aplin, K.P., and J.M. Pasveer. 2005. Mammals and other vertebrates from late Quaternary archaeological sites on Pulau Kobroor, Aru Islands, eastern Indonesia. Pp. 41–62 in O’Connor, S., P. Veth, and M. Spriggs (eds.) Archaeology of the Aru Islands, Terra Australis 22. Pandanus Press, Canberra. Aplin, K.P., J.M. Pasveer, and W.E. Boles. 1999. Late Quaternary vertebrates from the Bird’s Head Peninsula, Irian Jaya, Indonesia, including descriptions of two previously

................. 16157$

CH10

02-08-07 10:28:13

PS

PAGE 252

Paleontology of Papua / 253 unknown marsupial species. Records of the Western Australian Museum Supplement 57: 351–387. Archbold, N.W. 1981a. Permian brachiopods from Western Irian Jaya, Indonesia. Geological Research & Development Centre, Paleontology Series 2: 1–25. Archbold, N.W. 1981b. Quinquenella magnifica sp. nov. (Chonetidina, Brachiopoda) from the Permian of Irian Jaya, Indonesia: a study of the ontogeny of a chonetid brachiopod. Geological Research & Development Centre, Paleontology Series 2: 27–34. Archbold, N.W. 1991a. Early Permian Brachiopoda from Irian Jaya. Bureau of Mineral Resources (BMR) Journal of Australian Geology and Geophysics 12 (4): 287–296. Archbold, N.W. 1991b. Late Paleozoic brachiopod faunas from Irian Jaya, Indonesia. Pp. 347–353 in MacKinnon, D.I., D.E. Lee, and J.D. Campbell (eds). Brachiopods through Time. A.A. Balkema, Rotterdam. Archer, M., R. Arena, M. Bassarova, K. Black, J. Brammall, B.N. Cooke, K. Crosby, H. Godthelp, M. Gott, S.J. Hand, B. Kear, A. Krikmann, B. Mackness, J. Muirhead, A. Musser, T.J. Myers, N. Pledge, Y. Wang, and S. Wroe. 1999. The evolutionary history and diversity of Australia’s mammals. Australian Mammalogy 21: 1–45. Flannery, T.F. 1992. New Pleistocene marsupials (Macropodidae, Diprotodontidae) from subalpine habitats in Irian Jaya. Alcheringa 16: 321–331. Flannery, T.F. 1994. The fossil land mammal record of New Guinea: a review. Science in New Guinea 20: 39–48. Flannery, T.F. 1995. Mammals of New Guinea. Australian Museum/Reed Books, Sydney. Flannery, T.F., M-J. Mountain, and K.P. Aplin. 1983. Quaternary kangaroos (Macropodidae; Marsupialia) from Nombe Rockshelter, Papua New Guinea, with comments on megafaunal extinction in the New Guinea Highlands. Proceedings of the Linnean Society of New South Wales 107: 77–99. Frakes, L.A. 1997. Grossplots: a method for estimating the temperature state of the Earth and of Australia, Cretaceous to Middle Miocene. Australian Journal of Botany 45: 359–372. Gerth, H. 1965. Ammoniten des mittleren und oberen Jura und der a¨ltesten Kreide vom Nordabhang des Schneegebirges in Neu Guinea. Neues Jahrbuch der Geologie und Paleontologie Abhandlungen 121: 209–218. Haberle, S.G. 1998. Late Quaternary vegetation change in the Tari Basin, Papua New Guinea. Palaeogeography, Palaeoclimatology, Palaeoecology 137: 1–24. Hall, R. 2001. Cenozoic reconstructions of SE Asia and the SW Pacific: changing patterns of land and sea. Pp. 35–56 in Metcalfe, I., J.M.B. Smith, M. Morwood, and I. Davidson (eds.) Faunal and Floral Migrations and Evolution in SE Asia-Australasia. Balkema, Lisse. Hardjasasmita, H.S. 1985. Fossil diprotodontid: Zygomaturus Owen 1859 dari Nimboran, Irian Jaya. Pertemnan Ilumah Arkeologi 3: 999–1004. Hope, G.S., T.F. Flannery, and Boeardi. 1993. A preliminary report of changing Quaternary mammal faunas in subalpine New Guinea. Quaternary Research 40: 117–126. Hope, J.H. 1981. A new species of Thylogale (Marsupialia: Macropodidae ) from Mapala rockshelter, Jaya (Carstensz) Mountains, Irian Jaya (Western New Guinea). Records of the Australian Museum 33: 369–387. Jongmans, W. 1940. Beitrage zur kenntnis der Karbonflora von Niederlandsch NeuGuinea. Mededeelingen Geologische Stichting 1938–1939: 263–274. Khan, A.H. 1976. Palynology of Neogene sediments from Papua (New Guinea) stratigraphic boundaries. Pollen et Spores 16: 265–284. Menzies, J.I., and C. Ballard. 1994. Some new records of Pleistocene megafauna from New Guinea. Science in New Guinea 20: 113–139.

................. 16157$

CH10

02-08-07 10:28:13

PS

PAGE 253

254 / geoffrey s. h o p e a n d k e n p. aplin Metcalfe, I. 2001. Palaeozoic and Mesozoic tectonic evolution and biogeography of SE Asia-Australasia. Pp. 15–34 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Morley, R.J. 2000. Origin and Evolution of Tropical Rain Forests. London: Wiley. Oloriz, F., and G.E.G. Westermann. 1998. The perisphinctid ammonite Sulaites n. gen. from the Upper Jurassic of the Indo-Southwest Pacific. Alcheringa 22: 231–240. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. Bureau of Mineral Resources Journal of Australian Geology and Geophysics 10: 193–211. Prasad, M.N.V. 1981. New species of fossil wood Planoxylon from the late Paleozoic of Irian Jaya, Indonesia. Bulletin of the Geological Research and Development Centre 5: 37–40. Quarles van Ufford, A., and M. Cloos. 2005. Cenozoic tectonics of New Guinea. American Association of Petroleum Geologists Bulletin 89: 119–140. Rich, P.V., M.D. Plane, and N. Schroeder. 1988. A pygmy cassowary (Casuarius lydekkeri) from late Pleistocene bog deposits at Pureni, Papua New Guinea. BMR Journal of Australian Geology and Geophysics 10: 377–389. Rich, P.V., and G.F. van Tets. 1982. Fossil birds of Australia and New Guinea: their biogeographic, phylogenetic, and biostratigraphic input. Pp. 235–384 in Rich, P.V., and E.M. Thompson (eds.) The Fossil Vertebrate Record of Australasia. Monash University, Clayton. Rigby, J.F. 1997. The significance of a Permian flora from Irian Jaya (West New Guinea) containing elements related to coeval floras of Gondwanaland and Cathaysialand. Palaeobotanist 45: 295–302. Tenison-Woods, J.E. 1878. On some Tertiary fossils, from New Guinea. Proceedings of the Linnean Society of NSW (1) 2: 267–268. Truswell, E.M. 1993. Vegetation changes in the Australian Tertiary in response to climatic and phytogeographic forcing factors. Australian Systematic Botany 6: 553–557. Westermann, G.E.G. 1995. Mid-Jurassic Ammonitina from the Central Ranges of Irian Jaya and the origin of stephanoceratids. Hantkeniana 1: 105–118. Westermann, G.E.G., and J.H. Callomon. 1988. The Macrocephalitidae and associated Bathonian and early Callovian (Jurassic) Ammonitina of the Sula Islands and New Guinea. Palaeontographica A 203: 1–90.

................. 16157$

CH10

02-08-07 10:28:14

PS

PAGE 254

2.7. Paleoecology and Paleoenvironments of Papua geoffrey s. hope h e n w e l o ok at the landscapes of Papua we can ask how they came into being and how long they have been as they are now. Because New Guinea has risen from the sea only in the last few million years and is an actively evolving landmass with tectonic activity and very high rates of erosion and accumulation, these landscapes are relatively young. However, some surfaces have existed for 100,000 years or longer. Such landscapes often preserve records of climate and environmental change in land forms and deposits such as glacial moraines, lake bog and cave sediments, alluvium and colluvial mantles, and marine deposits (Williams et al. 1998). Information about the changes experienced by biomes comes from the study of biological and geomorphological records supported by dating. Here we consider what is known of the changes over the past 60,000 years or so, based on studies of pollen and sediments in swamps and small lakes in a range of Papuan environments from the lower montane to high alpine zones. There are also marine records of land pollen from southeast of Papua in the Banda Sea. While no other proxy has been studied in Papua there is scope for work on paleofaunas, opaline phytoliths, diatoms, and possibly tree rings. There is now a considerable body of knowledge about the Quaternary, covering the last two million years, and during which time the present land mass of New Guinea was effectively in place. The Quaternary marks the time when global ice caps expanded periodically and the mass of deep ocean was chilled to 4C. It is a glacial epoch during which the climate is relatively sensitive to variations in the seasonality of sunlight, which induced periods of ice build-up called glacials and periods of relatively reduced ice, called interglacials. In general terms, glacials have lasted about 100,000 years and interglacials about 10,000 years over the past million years or so. (Ages less than 25,000 are given as calibrated ages before present (bp) unless stated otherwise). At present we are near the end of an interglacial known as the Holocene (Hope 2005). Growth of ice sheets during the glacial maxima lock up twice as much ice on land in the northern hemisphere as is contained in Antarctica today. This causes sea level to fall and exposes large areas of dry land (Figure 2.7.1). The sea level 22,000 years ago lay at about 110 m, but for most of the last 100,000 years sea level has been 20–60 m lower than present. Paleo-shorelines for the last 22,000 years for southeastern Papua have been modeled by Yokoyama et al. (2001), allowing for tectonic sinking associated with loading the shelf with water. The Arafura Shelf extended west to Aru Islands, and was a broad flat plain that connected Papua to the Aru Islands and Australia for perhaps 85% of Quaternary time (Figure 2.7.2). Much of the remainder of Papua was little changed, as the coast is very steep. However, Bintuni Gulf was dry land

W

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

255

................. 16157$

CH11

02-08-07 10:27:39

PS

PAGE 255

256 / geoffrey s. h o p e

Figure 2.7.1. Sea level change for the past 300,000 years.

Figure 2.7.2. Sea levels at glacial maximum.

................. 16157$

CH11

02-08-07 10:27:52

PS

PAGE 256

Paleoecology and Paleoenvironments of Papua / 257

and Misool and Salawati islands were joined to the Vogelkop. From 100,000 years to about 11,000 years ago, apart from a refreshing 3 km swim across the Sagewin Channel from Batanta Island to then Salawati peninsula near Sorong, it would have been possible to trudge the 5,000 km from Kawe´ Island on the equator, to the hills of Maatsuyker, then a rocky peninsula of southernmost Tasmania at 44S. Biak Island expanded but remained distinct from Yapen Island and the mainland. The major landscape units of Papua are the southern plain, the central ranges, the intermontane trough, and the northern ranges. The same structural elements are found at a smaller scale on the Vogelkop Peninsula. Because the climate and geomorphology is different in each zone we can examine environmental change in each.

Glaciation and Deglaciation in the High Mountains Glaciation has affected only a tiny area of tropical Papua, but it is the clearest expression of climate change and the effects of changing temperatures are also seen in changes of vegetation boundaries. The 3 km2 of glacial ice that sits on Puncak Jaya above 4,650 m altitude today is a remnant of once extensive glaciers that covered ca 2,200 km2 above 3,400 m on the high mountains across the island. At the maximum ice extent of ca 1,400 km2, the Merauke Range (Snow Mountains) were an almost continuous wall of ice with breaks east of Mt Mandala, and at the Baliem Valley and in a few passes south of Ilaga. Ice was also present on the summits of the Weyland and Nggumbulu ranges near Nabire (Hope et al. 1976; Peterson et al. 2001) but not in the Arfak Mountains. This change in the altitude of ice formation is the clearest expression that global climate change influenced Papua and we can assume that the conditions that lowered the snow line also caused changes at lower altitude. The question is whether tropical cooling coincided with the build up of large ice caps in the Northern Hemisphere. To study this, glacial history can be dated using the cosmogenic age of glacial features that have either resulted from stripping of rocks in mountain cirques or their deposition in features such as moraines or outwash deposits. Another method is to use radiocarbon dating of organic matter caught up in lake basins that were formed by the glacier. These ages will give the minimum age when ice had retreated from the basin. Cosmogenic dating is still a new technique that has only been applied to glacial features on Mt Trikora, in the northeastern Lorentz National Park. Here the southern wall of the mountains is 4,400–4,750 m in height and a small ice cap was present on the summit until the 1960s (Hope et al. 1973). The mountains lie south of a broad plateau onto which the ice flowed, leaving steep sided moraines for several kilometers. This excellent preservation has provided the best evidence for at least two glacial advances, as some moraine valleys are overrun by others that must be younger (Figure 2.7.3). Samples of limestone boulders were collected from older and younger moraine crests and the isotope Chlorine 36 measured by accelerator mass spectrometry at Lucas Heights, Australia (Prentice et al. 2005).

................. 16157$

CH11

02-08-07 10:27:52

PS

PAGE 257

258 / geoffrey s. h o p e

Figure 2.7.3. Moraine series north of Mt Trikora. The results show that the earlier retreat took place around 20,000 years ago while the latest is only 12–14,000 years old. These ages are in reasonable agreement with the last major glacial advances elsewhere in the world. Earlier glacial activity is suggested by five series of gravel terraces downstream from the moraines on the east Baliem River. The terraces are probably formed from outwash but a soil and logs were found beneath the lowest and youngest one. A log gave a radiocarbon age of 34,000 bp, reflecting a time when climates were relatively warm before the glacial advances (Hope et al. 1993). Pollen studies on two cores from Lake Habbema (3,200 m altitude and outside the glacial limit) show that forests similar to present were on the plateau by about 10,000 years ago. Dating of swamps and lake basins north of Mt Jaya suggests that ice had retreated from the moraine which dammed Lake Hogayaku by 15,100 years ago (Figure 2.7.4). By 16,500 bp at Ijomba Mire ice was well up the valley but continued to produce glacial debris in streams until around 12,600 years ago (Hope et al. 1976). Lake Hogayaku has some evidence of former advances in that a terminal moraine contains more than one till, and the lake core extends back to about 34,000 bp in rock flour (Prentice et al. 2005; Figure 2.7.5). This core is most unusual in showing an open scrub that becomes more open, perhaps prior to glacial advance. The rock flour in the core at this stage shows that ice was not far off and it may have overridden the core. The retreat after 17,700 bp correlates with the invasion of the newly exposed landscapes by a high altitude scrub and Cyathea treeferns which lasted until about 12,400 years ago. After this a richer forest slowly invades, suggesting that the temperature finally had allowed subalpine forest elements such as conifers (e.g., the podocarp Dacrycarpus compactus) to invade. The

................. 16157$

CH11

02-08-07 10:28:01

PS

PAGE 258

Paleoecology and Paleoenvironments of Papua / 259

Figure 2.7.4. Moraine maps north of Mt Jaya. Source: M. Prentice.

Figure 2.7.5. Pollen diagram from Lake Hogayaku. Composite of two cores.

................. 16157$

CH11

02-08-07 10:28:13

PS

PAGE 259

260 / geoffrey s. h o p e

Holocene subalpine forest is fairly stable until human disturbance started to remove it (Chapter 6.1). Although the dating is still too poor to be sure that maximum ice advance coincides with the peak of ice advances in the Northern Hemisphere, the general timing of ice maximum and retreat accord with Marine Isotope Stage 2. It is also in good agreement with other sites in New Guinea (Peterson et al. 2001) but the snow line appears to have been higher on Mt Jaya, at an estimated 3,900–4,000 m, compared to 3,450 m in the Star Mountains and further east (Prentice et al. 2005). This apparent rise westward in the snow line may argue for a precipitation gradient that is not as pronounced today. A possible mechanism for this is the Arafura Shelf which provided a dry plain southwards which would have reduced moisture imports with the Southeast Trades compared to the present day. This may also explain why evidence for glacier retreats and advances seem to be more common on the East Baliem and the Kemabu plateaus north of the mountains compared to further east. As the shelf flooded precipitation would have risen after 17,000 bp and this may have supported two small glacial advances noted at the Ertsberg on the southern wall of Mt Jaya at 16,000 and 13,800 bp. Prentice et al. (2005) consider that at maximum the snow line was 650–850 m lower than present, suggesting mean thermal change of about 5C. Most well preserved glacial landforms relate to the last 30,000 years and there is little evidence for older glaciations apart from the possible outwash terrace sequences. However, in Papua New Guinea an older glaciation is preserved on Mt Giluwe and dated with K/Ar ages of ca 280–220,000 bp from palagonitic breccias on the northern slopes (Lo¨ffler 1976). It is thus likely that New Guinea mountain areas have experienced long periods of extensive snow cover in general accord with glacial cycles.

The Basins of the Central Ranges From a 30,000 year record from Sirunki, in central Papua New Guinea, the tree line at the height of the glaciation seems to have been at about 2,200 m altitude (Walker and Flenley 1979). Below this, Nothofagus-dominated forest seems to have been very common (Hope 1996a,b). Numerous ideal sites for research exist in the basin and range topography of the Central Highlands and Vogelkop Peninsula, for example the massive swamplands east of Danau Paniai and the Anggi-Gigi lakes south of Manokwari. However so far only a discontinuous record has been obtained from the floor of the Baliem Valley, a large alluvial basin at 1,550 m north of Wamena. At Supulah Hill, a low sandstone hill in the center of the valley, a small hollow preserves a three meter sequence of peats that spans ca 38–34,000 bp (Hope 1998). Pollen shows that a forest dominated by beech occupied the forest floor, similar to forests on the slopes 400 m above today. This suggests cloudy conditions with consistent rainfall. The site was eroded and burnt after this time and forest fires at 29,300 bp are recorded in charcoal caught in eroding sands on the hill.

................. 16157$

CH11

02-08-07 10:28:14

PS

PAGE 260

Paleoecology and Paleoenvironments of Papua / 261

The record resumes 6 km north at Kelela Swamp where an abandoned stream meander has infilled with 5 m of peat (Haberle et al. 1991). The basal samples, dated to about 7,400 years ago, suggest the swamp at the time was a swamp forest and that a mixed montane forest with Castanopsis (oak) occupied the slopes nearby. This forest type requires warmer, sunnier conditions than beech forest. While the period of transition from one forest type to another is missing in the Baliem, records from analogous basins at Tari and Sirunki in Papua New Guinea have a similar change occurring about 14,000 years ago (Haberle 1998). From a consideration of all montane sites it seems that the boundary of mixed oak forest and beech forest rose about 700 m at the start of the Holocene, reflecting a temperature rise of 5–6C.

Northern Ranges and Lowland Trough North of the Central Ranges lies the (Mamberamo) lake plain, a large basin of subsidence through which flow the Tariku and Taritatu rivers. The swamp forest record here may be some hundreds of meters deep and extend back several hundred thousand years, because this basin is dammed by the rise of mountains such as the Foja to the north. Unfortunately, the results of oil drilling west of the Mamberamo do not seem to be available. A 33.5 m core of organic-rich muds was obtained from Cenderawasih Bay, 2 km offshore of the Coffin Hills north of Nabire, but this spans only ca 16,000 years and has very sparse pollen, mainly fern spores (Prentice et al., unpublished). Water temperatures, indicated by stable isotopes from foraminifera, indicate temperatures 3–4 colder than present at 16,000 bp. There are fluctuations of equivalent magnitude in the Holocene, suggesting rapid cooling in Cenderawasih Bay on several occasions. The longest record from Papua comes from Lake Hordorli, a swamp basin formed on a bench at ca 780 m altitude on the Cyclops Mountains 8 km north of Sentani. The Cyclops Mountains are an isolated range that cut off the Sentani area from the sea, resulting in lower rainfall around Lake Sentani. A 10 m core of lake muds was obtained with a D section corer (Figure 2.7.6) that covers about 70,000 years and records changes in the lower montane forest that reflect changes in temperature (Hope and Tulip 1994). Although the dating extends only to 4 m, the timing of changes seem to accord well with the marine isotope stages that reflect global cooling and draw-down of ocean levels (Figure 2.7.7). Before ca 14,750 bp Nothofagus was more prominent and the lake may have been deep, suggesting that the present summit beech forest was lower and the mountain slopes more humid. By 11,600 years ago the level of beech had fallen to modern levels but it rose abruptly between 9–7,000 years ago, after which it declines, being replaced by Araucaria, Castanopsis, and other montane trees. This also suggests a rise in temperature of about 4C since glacial times. The data extend back to Marine Isotope Stage 4 and suggest that the Cyclops Mts have been humid throughout the last glacial period and have always been well forested. Unfortunately this site tells us

................. 16157$

CH11

02-08-07 10:28:15

PS

PAGE 261

262 / geoffrey s. h o p e

Figure 2.7.6. Coring at Lake Hordorli.

Figure 2.7.7. Pollen diagram from Lake Hordorli.

................. 16157$

CH11

02-08-07 10:28:26

PS

PAGE 262

Paleoecology and Paleoenvironments of Papua / 263

little about conditions around Lake Sentani, which may well have been drier than today during the glaciation. A major cause of more recent change around all the coast of Papua has been the rise of the sea after the glacial maximum until it reached present levels around 6,000 years ago. The modern coastline represents only 6,000 years of adjustment, with the development of beaches, erosion of cliffs, and infilling of lagoons formed by the flooding of valleys. One striking marine spit has built out to almost enclose Jayapura Bay (Figure 2.7.8). The land is steadily rising and former beaches and reefs have been lifted from the sea to form limestones around Jayapura. Lake Sentani is a former bay cut off by tectonic uplift. On other coasts the early open coastline is gradually cut off as coral growth creates reefs that slow down the wave energy, creating lagoons.

The Southern Plain The southern lowlands of Papua have a much stronger seasonal rainfall pattern and are bounded by a shallow sea that withdrew at times of lower sea level. We might then expect that this area was drier during glacial times, when an arid plain extended southwards and the monsoon was weaker. Such climates were hypothesized by Nix and Kalma (1972) and van der Kaars (1991), who map a northern extension of the savanna woodland-rainforest boundary. They explain relictual areas of eucalypt savanna in Papua New Guinea as dating from these dry times in Marine Isotope Stages 3 and 2. There are as yet no Pleistocene lowland records from southern Papua but the post-glacial migration of the rainforest boundary southwards has been identified on the Aru Islands which were then part of Papua (Hope and Aplin 2005). A faunal sequence starts about 20,000 bp and contains savanna fauna such as agile wallaby, but some rainforest elements such as possums are present in small numbers. Around 14,000 years ago the wallabies became rarer and rainforest taxa increased, suggesting that the boundary between closed forest and savanna had reached this site. This reflected a more maritime climate as the Arafura Shelf flooded.

Figure 2.7.8. A sandspit and beach ridge complex cutting across Jayapura Bay has formed in the last few thousand years.

................. 16157$

CH11

02-08-07 10:28:28

PS

PAGE 263

264 / geoffrey s. h o p e

From lowland caves near Lake Ayamaru in the southern part of the Vogelkop, Aplin et al. (1999) record late Pleistocene faunas that include animals from higher altitude, suggesting a cooler climate and more varied vegetation within hunting range of the cave. On the south coast five cores were taken from the estuarine plain east of Timika in the Ajkwa and Tipoeka estuaries (Ellison 2005). These provide a record back to 9,000 bp that records the development of mangrove communities as the sea flooded inland. The evidence for the extension of land seawards as the tremendous sediment load from the mountains builds up and pushes the coastline southward is balanced by evidence for continuing subsidence. The sequences record flooding by the sea around 6,500–5,500 years ago and rapid siltation by mangroves. However at the study sites landward communities of Ceriops/Brugiera are replaced by more marine Rhizophora, suggesting more frequent tidal flooding despite the build up of sediment. Hence sedimentation has not kept up with subsidence in this case. Mine waste discharge from the Otomona River has reversed this trend locally by burying forest over several km2 (Paull et al. 2006). In other areas the relationship of sea and land is not well known. It is possible that land has built out onto the shelf for some kilometers in the more southerly Asmat area. Given the plentiful discharge of rivers onto the southern plain, swamp and lagoonal environments have probably always been plentiful, but the rise of sea level has resulted in a great expansion of wetlands in the last 5,000 years or so.

Discussion The vegetation history of Papua suggests that the land has mostly remained under forest despite the fluctuating climates of the Quaternary. However, at the height of glacial periods the southern lowlands supported eucalypt and Nauclea savanna while the mountain crests above 2,000 m were more open with subalpine grasslands. Some rain shadow areas such as Sentani may also have been more open woodlands with grassland patches at that time. These major formation shifts have probably repeated themselves throughout the Quaternary and have preconditioned the biota to tolerate change. The subalpine forest-grassland alternation has produced islands of habitat that have probably encouraged speciation and the development of localized endemism in the upper mountain flora. While the forest stability has been important in encouraging biodiversity and specialization to a very variable local setting, the forest composition has constantly changed. Nothofagus has become less common in the Holocene in response to drier and less cloudy conditions. At Lake Hordorli there is a shift to a higher proportion of secondary species during warmer times. This may reflect shorter tree life spans typical of lower altitude forests that are subject to high levels of insect attack and disease. There is huge potential for more paleoecological studies in Papua due to its abundant resource of swamps, lakes, caves, and other sources of sediment. Such studies provide a time scale for understanding modern ecological processes. Even

................. 16157$

CH11

02-08-07 10:28:29

PS

PAGE 264

Paleoecology and Paleoenvironments of Papua / 265

the minimal work so far undertaken in New Guinea has demonstrated the individualistic nature of site responses to environmental change. Although broad patterns of climate and anthropogenic change can be proposed, each site provides insight into the resilience and adaptability of the ecology of that area, often showing different responses. This work provides an understanding of evolutionary mechanisms. It can be analyzed to show the stability of species assemblages through time and to help us understand how representative the modern ecosystems might be.

Literature Cited Aplin, K.P., J.M. Pasveer, and W.E. Boles. 1999. Late Quaternary vertebrates from the Bird’s Head Peninsula, Irian Jaya, Indonesia, including descriptions of two previously unknown marsupial species. Records of the Western Australian Museum Supplement 57: 351–387. Ellison, J. 2005. Holocene palynology and sea-level change in two estuaries in Southern Irian Jaya. Palaeogeography, Palaeoclimatology, Palaeoecology 220: 291–309. Haberle, S.G. 1998. Late Quaternary vegetation change in the Tari Basin, Papua New Guinea. Palaeogeography, Palaeoclimatology, Palaeoecology 137: 1–24. Hope, G.S. 1989. Climatic implications of timberline changes in Australasia from 30,000 bp to present. Pp. 91–99 in Donnelly, T., and R. Wasson (eds.) CLIMANZ 3. CSIRO, Div. Water Resources, Canberra. Hope, G.S. 1996a. Quaternary change and historical biogeography of Pacific Islands. Pp. 165–190 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Process. SPB Publishing, Amsterdam. Hope, G.S. 1996b. History of Nothofagus in New Guinea and New Caledonia. Pp. 257–270 in Veblen, T.T., R.S. Hill, and J. Read (eds.) The Ecology and Biogeography of Nothofagus Forests. Yale University Press, New Haven, Connecticut. Hope, G.S. 1998. Early fire and forest change in the Baliem Valley, Irian Jaya, Indonesia. Journal of Biogeography 25: 453–461. Hope, G.S. 2005. The Quaternary in Southeast Asia. Pp. 24–37 in Gupta, A. (ed.) The Physical Geography of Southeast Asia. Oxford University Press, Oxford. Hope, G.S., and K. Aplin. 2005. Environmental change in the Aru Islands. In O’Connor, S., M. Spriggs, and P. Veth (eds.) The Archaeology of the Aru Islands, Eastern Indonesia. Pandanus Press, Canberra. Hope, G.S., J.A. Peterson, and R. Mitton. 1973. Recession of the minor ice fields of Irian Jaya. Zeitschrifte fur Glestcherkunde and Glazialgeologie IX: 73–87. Hope, G.S., J.A. Peterson, U. Radok, and I. Allison. 1976. The Equatorial Glaciers of New Guinea. A.A. Balkema, Rotterdam. Hope, G.S., and J. Tulip. 1994. A long vegetation history from lowland Irian Jaya, Indonesia. Palaeogeography, Palaeoclimatology, Palaeoecology 109: 385–398. Nix, H.A., and J.D. Kalma. 1972. Climate as a dominant control in the biogeography of northern Australia and New Guinea. Pp. 61–91in Walker, D. (ed.) Bridge and Barrier: The Natural and Cultural Heritage of the Torres Strait. Department of Biogeography and Geomorphology, Research School of Pacific Studies, Australian National University, Canberra. Paull, D., G. Banks, C. Ballard, and D. Gillieson. 2006. Monitoring the environmental

................. 16157$

CH11

02-08-07 10:28:30

PS

PAGE 265

266 / geoffrey s. h o p e impact of mining in remote locations through remotely sensed data. Geocarto International 21, 33–42. Peterson, J.A., G.S. Hope, M. Prentice, and W. Hantoro. 2001. Mountain environments in New Guinea and the late Glacial Maximum ‘‘warm seas/cold mountains’’ enigma in the West Pacific Warm Pool region. Pp. 173–187 in Kershaw, P., B. David, N. Tapper, D. Penny, and J. Brown (eds.) Bridging Wallace’s Line: Advances in GeoEcology. Catena Verlag, Reiskirchen. Prentice, M.L., G.S. Hope, K. Maryunani, and J.A. Peterson. 2005. An evaluation of snowline data across New Guinea during the last major glaciation, and area-based glacier snowlines in the Mt. Jaya region of Papua, Indonesia, during the Last Glacial Maximum. Quaternary International 138–139: 93–117. van der Kaars, W.A. 1991. Palynology of eastern Indonesian marine piston-cores: a Late Quaternary vegetational and climatic record for Australasia. Palaeogeography, Palaeoclimatology and Palaeoecology 85: 239–302. Webster, P., and N. Streten. 1978. Late Quaternary ice age climates of tropical Australasia, interpretation and reconstruction. Quaternary Research 10: 279–309. Yokoyama, Y., A. Purcell, K. Lambeck, and P. Johnston. 2001. Shore-line reconstruction around Australia during the Last Glacial Maximum and Late Glacial Stage. Quaternary International 83–85: 9–18.

Glossary of Tectonic and Geological Terms Structure Craton: Old, thick continental platform Lithosphere: The outermost layer of the earth, lying above the mantle Outwash terraces: Gravel terraces built by rivers flowing from glaciers Subduction zone: Boundary between two tectonic plates where one slides beneath the other Terrane: A geological unit incorporated into a larger land mass but not originally formed as a part of that land mass Till, moraine: Rock debris deposited by glaciers Transform fault: A boundary between two geological units, such tectonic plates, where one slides past the other without subduction or orogeny Process Orogeny: Creation of mountains, usually due to the collision of tectonic plates Subduction: The process of one tectonic plate sliding beneath another Rock Types Igneous: Rocks formed by volcanic activity Sedimentary: Rocks formed from eroded material deposited by wind or water Metamorphic: Igneous or sedimentary rocks that have been altered by heat and pressure Position Subaerial: Above water, as in reference to land masses emergent above the surface of the ocean Submarine: Lying below the surface of the ocean

................. 16157$

CH11

02-08-07 10:28:30

PS

PAGE 266

section three 

The Flora

................. 16157$

PRT3

04-13-07 13:50:07

PS

PAGE 267

................. 16157$

PRT3

02-08-07 10:21:53

PS

PAGE 268

3.1. Introduction to the Flora of Papua wayne n. takeuchi e w Gu i n e a is the world’s second largest island, with the Indonesian province of Papua (formerly Netherlands New Guinea, Irian Jaya, Papua Barat) in its western half, and the independent state of Papua New Guinea (PNG) in the east. The island comprises the greater part of a region known as Papuasia, an area consisting of New Guinea, the Bismarck Archipelago, and the Solomon Islands. Although the true size of its botanical inventory is unknown and subject to considerable speculation, there is little doubt that Papuasia is a major paleotropical center for floristic diversification. Modern estimates of the Papuasian flora have ranged widely between 11,000 (Collins et al. 1991) and 20,000 species (Womersley 1978), clearly reflecting the many gaps and uncertainties in existing knowledge. Using ferns and orchids as baselines for extrapolation, a new assessment suggests that 20,000 to 25,000 vascular plant species could be present in Papua (Supriatna et al. 1999). If correct, this projection will require a significant and corresponding adjustment in the overall register for Papuasia, but it will be many decades before a definitive judgment is possible. In 1950, the ‘‘collections index’’ (i.e., collections density; number of plant specimens/100 km2) was approximately 12 for the whole of New Guinea (Frodin 1990). At least in the eastern half of the island, collection activity increased substantially over the next 50 years, particularly during the interval from 1965 to 1975 when 40,000 gatherings were made by institutional collectors based at Lae. Since 1990 the accession rate for the PNG National Herbarium (LAE) has declined sharply, to a present 15-year average of about 400 numbers per year. Owing to earlier efforts under the territorial administration, the collections index for PNG is now near 50, the minimal benchmark for an ‘‘adequate’’ floristic inventory (Campbell 1989; Stevens 1989). By comparison, Papuan exploration has lagged considerably behind that of PNG. With a current collections index of less than 20 (Supriatna et al. 1999), Papua has shown only marginal improvement since the 1950s. Although many PNG localities are regarded as underexplored, the overall sampling intensity for the eastern side is at least 300% better than for the western side. Future progress in documentation will be ultimately contingent on the creation of a skilled corps of indigenous surveyors. Botanical commentators have made similar observations for many years, most recently Middleton (2003) and Roos (2003) for other parts of Malesia, but such recommendations have rarely been effected within Papuasia. Much of the existing information on the Papuan flora has been derived from a small number of areas, including primarily the Cyclops Mts, Freeport-Timika, the Idenburg River, Mt Jaya, Trikora-Habbema, Wissel Lakes, and the northeast

N

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

269

................. 16157$

CH12

02-08-07 10:27:49

PS

PAGE 269

270 / wayne n . t a k e u c h i

Vogelkop. These venues can be regarded as well collected only in a comparative sense relative to the rest of the province; there are no localities where sampling saturation has been achieved in a manner equivalent to the Buso-Kamiali, LaeMarkham, and Wau-Bulolo districts of PNG. Vast tracts of Papuan natural growth forest are still botanically unknown, particularly in the central parts of the province. The majority of Papuan localities are so poorly cataloged that knowledge of plant distributions is exceedingly fragmentary. Numerous species undoubtedly remain undiscovered. Although the principal gaps in floristic exploration have been identified by Campbell (1989), Prance (1977), Stevens (1989), and Supriatna et al. (1999), the examination of virtually any area would be immensely rewarding. To take just one example: a two week inspection of coastline communities near Kokas (Bomberai Peninsula) recently uncovered about eight new taxa (e.g., Takeuchi 2003a) and Papuan distributional records for Hypserpa laurina, Conandrium rhynchocarpum, Barringtonia josephstaalensis, and Zanthoxylum nitidum (Figures 3.1.1–6). Distributional maps from modern revisions (e.g., Middleton 1997; Pannell 1992) often provide implicit indications of the sampling inequalities responsible for such records. Due to the dismal state of botanical exploration in Papua, even surveys conducted using rapid-assessment methods can give returns exceeding their time investment. The scientific contrasts between the geographic halves of New Guinea are clearly reflected in the differing status of local infrastructure. Prior to the granting of PNG independence in 1975, the former Australian administration established effective facilities at a number of centers (notably Port Moresby, Bulolo, and Lae), that are today the institutional foundations for forestry science in PNG. Comparable developments have been conspicuously lacking on the Papuan side. In particular, the construction of an international grade herbarium at Lae, and the activities associated with its emergence, were never replicated in the west. The only herbarium in Papua is a small facility (Figures 3.1.7, 8) located on the campus of the Papua State University at Manokwari (MAN). Its modest total of 30,000 mounted sheets consists primarily of Boswezen Neeuwgini collections, many of which are represented by sterile duplicates. Even including the estimated 20,000 unmounted specimens in backlog (R. Maturbongs, pers. comm.), the MAN holdings are incongruously meager by comparison to the 400,000 mounted sheets at LAE. Manokwari’s limited floor space (80 m2) will constrain future expansion. The poor state of Papuan infrastructure and service capacities are clearly relevant to an understanding of the past and present inactivity in botany, though there are certainly many other contributing factors. In any evaluation of the logistical and financial obstacles to operations in developing societies, in situ programs have obvious advantages over foreign-based initiatives. Without substantive improvements to local infrastructure, it is unlikely that new collecting programs can be executed on the scales required to achieve a satisfactory inventory. Future initiatives will also be disadvantaged by the lack of a detailed forest classification and mapping system for Papua. The Regional Physical Planning Program for Transmigration (RePPProt; 1990) has developed a series of biophysical maps at 1:250,000

................. 16157$

CH12

02-08-07 10:27:50

PS

PAGE 270

Introduction to the Flora of Papua / 271

Figure 3.1.1. Glochidion daviesii (Euphorbiaceae sens. lat.; Phyllanthaceae), discovered in 2002 from populations near the Saengga River in Papua. The cauline fruits are presumably an adaptation to cassowary-aided dispersal.

................. 16157$

CH12

02-08-07 10:28:21

PS

PAGE 271

272 / wayne n . t a k e u c h i

Figure 3.1.2. Closer view of the fruit clusters in Glochidion daviesii. scale, but there is nothing equivalent to the vegetation maps for Sumatra (Laumonier et al. 1983, 1986, 1987; Laumonier 1997a,b) or to the Forest Inventory Mapping System for PNG (Hammermaster and Saunders 1995a,b). As a partial response to the planning difficulties imposed by this situation, The Nature Conservancy is currently developing a vegetation map for the lowland ecosystems of northern Papua (D. Neville and M. Summers, pers. comm.). Botanical documentation in the form of plant specimens is the primary evidence on which floristic knowledge is based, so it is no surprise that the publications portfolio for Papua is as thin as the collections index. There is little in the way of synthetic information. Much of the existing literature consists of technical articles scattered through a wide range of journals and books (written in various languages), or as unpublished material of restricted distribution. There are no general treatments like those available on the PNG side (e.g., Borrell 1989; Conn 1995; Henty 1981; Johns 1987, 1988, 1989; Peekel 1984; Percival and Womersley 1975; Womersley 1978) although the floristic overlap between east and west allows for a certain degree of cross-application. Even general accounts are relatively limited in number, consisting for the most part of expedition summaries (Archbold et al. 1942; Brass 1941; Donnelly et al. 2004; Mack and Alonso 2000; Rand and Brass 1940; Ridley 1916; van Royen 1960; Takeuchi 2003b; Vink 1965; Wollaston 1914), botanical commentaries (Conn 1994; Frodin 1988, 1990; Frodin and Gressitt 1982; Stevens 1989), checklists (Coode et al. 1997; Sidiyasa et al. 1997; Streimann 1986), conservation assessments (Craven and de Fretes 1987; Erftemeijer et

................. 16157$

CH12

02-08-07 10:28:38

PS

PAGE 272

Introduction to the Flora of Papua / 273

Figure 3.1.3. Barringtonia josephstaalensis (Lecythidaceae). A recently described species known only from the type locality (Josephstaal, PNG) and from the lowland forests along Bintuni Bay (Papua). The apparent 1,350 km disjunction between localities may be an artifact of low collecting densities. Scale bar: 2 cm.

................. 16157$

CH12

02-08-07 10:29:06

PS PAGE 273

274 / wayne n . t a k e u c h i

Figure 3.1.4. A distinctive feature of B. josephstaalensis is the smooth blue epicarp, recalling drupes of Elaeocarpus. Scale bar: 1 cm. Photo: R. Maturbongs.

al. 1989; Ruitenbeek 1992; Supriatna et al. 1999), and vegetation descriptions (van Balgooy 2001; Gibbs 1917; Hope 1976; Kalkman 1963; Mangen 1993; Rappard and van Royen 1959; van Royen 1956, 1963, 1965, 1967). Compared to neighboring PNG, there is a pronounced paucity of useful publications. Any improvement in Papuan exploration schedules would have numerous collateral benefits. There is probably no other area in Malesia with Papua’s latent potential for taxonomic discovery. Although Sumatra and Sulawesi have comparably low collecting densities (Stevens 1989; Whitten et al. 2000), their floras are decidedly less impressive (Table 3.1.1). Exploratory surveys on these islands would not be as consequential as similar efforts directed at appropriate targets in New Guinea. From considerations relating to floristic endemism, environmental quality, and biogeography, Papua is the most compelling choice as a focal site for future inventories.

Floristic Endemism Even with the limited collecting that has already occurred, it is obvious that numerous endemics are present in Papuan environments (Figures 3.1.9–14). The island of New Guinea as a whole has long been regarded as a hotspot for biotic diversification, and ongoing work from Flora Malesiana (FM) provides increasing support for this view. Compilations from completed FM revisions show more endemic species from New Guinea than from any other Malesian area, with a

................. 16157$

CH12

02-08-07 10:29:21

PS

PAGE 274

Introduction to the Flora of Papua / 275

Figure 3.1.5. An undescribed Aglaia (Meliaceae) from south Bintuni, vegetatively distinguished by elliptic-oblong leaflets (to 16-jugate) copiously marked by glandular spots. The novelty is unfortunately known only from sterile collections.

................. 16157$

CH12

02-08-07 10:29:56

PS

PAGE 275

276 / wayne n . t a k e u c h i

Figure 3.1.6. Bomberai savanna dominated by Melaleuca leucadendron sens. lat. (Myrtaceae), and Dicranopteris linearis (Gleicheniaceae). The presence of narrow endemics (e.g., Scaevola burnettii, Goodeniaceae) suggests that these communities have natural, rather than anthropogenic origins.

Figure 3.1.7. Manokwari Herbarium. Exterior view of the facilities at Papua State University (Universitas Negeri Papua). pronounced concentration of such taxa in the northern half of the island (van Balgooy et al. 1996). Supriatna et al. (1999) estimated species-level endemism in Papua at 60–90%, and Johns (1995) gives 60–70% as the overall figure for New Guinea. The lichen flora appears to have the highest endemism of any tropical area in the world (Chapter 3.2). Though the figures are highly provisional, Flora Malesiana revisions for angiosperms have yielded the following species-endemism

................. 16157$

CH12

02-08-07 10:30:21

PS

PAGE 276

Introduction to the Flora of Papua / 277

Figure 3.1.8. Interior view of the herbarium, showing the fixed shelving and aluminum containers for specimen sheets. The facility is presently without airconditioning.

Table 3.1.1. Floristic summary of major Malesian areas

Region

Collections index

Number of endemic genera

Percent species endemism

Number of phanerogam species

Borneo

35 (Kalimantan: 12)

59

37%

10,000–15,000

Java

199

10 (West Java)

5%

4,500

Malay Peninsula

⬎175

20 (41 incl. S. Thailand)

14%

7,500

New Guinea

30 (for Papuasia)

ca. 80

54%

ca. 20,000–25,000 (vascular plants)

Philippines

85

26

27–28%

8,000 (12,000 incl. all plants)

Sumatra

21–22

17

11%

8,000–10,000

Sulawesi

24

7

13–14%

5,000

Note: Collections index  number of plant specimens/100 km2; phanerogams  seed plants. Source: Ashton 1989; Biodiversity Information Center 2003 (www.pnh.com.ph); de Vogel 1989; de Wilde 1989; Frodin 1990; Johns 1995; Madulid 1982; Tan and Rojo 1989; P. van Welzen pers. comm. (cumulative statistics for percent endemism taken from Flora Malesiana revisions); Whitmore 1973; Whitten et al. 2000; Whitten et al. 2002.

................. 16157$

CH12

02-08-07 10:30:38

PS PAGE 277

278 / wayne n . t a k e u c h i

Figure 3.1.9. Anakasia simplicifolia (Araliaceae). The sole representative of a genus endemic to the westernmost parts of Papua Province (fruiting racemes were detached from another plant and placed against the pictured specimen). Faika (Monimiaceae) is an occasional associate.

................. 16157$

CH12

02-08-07 10:31:03

PS

PAGE 278

Introduction to the Flora of Papua / 279

Figure 3.1.10. Bomberai Peninsula, Papua. Typical understory habitat of Anakasia simplicifolia and Faika villosa.

................. 16157$

CH12

02-08-07 10:31:36

PS

PAGE 279

280 / wayne n . t a k e u c h i

Figure 3.1.11. Papuechites (Apocynaceae) is a vining genus endemic to New Guinea and nearby islands. It is represented by a single species, P. aambe (shown in fruit and flower), widely distributed in lowland environments. Scale bar: 1 cm. percentages for the major Malesian centers: New Guinea 54%, Borneo 37%, Philippines 27–28%, Malay Peninsula 14%, Sulawesi 13–14%, Sumatra 11%, and Java 5% (P. van Welzen, pers. comm., from revisions completed as of July 2004). Malesian endemism is concentrated primarily in the largest (unrevised) families, so the FM-based estimates are probably too conservative and will be driven upwards as statistics from the most speciose groups become available (Johns 1995). This is demonstrated by comparison with the then-current percentages from 1995: New Guinea 45%, Borneo 30%, Philippines 20%, Malay Peninsula 11%, Sulawesi 9%, Sumatra 8%, and Java 3%. The relative standing of the areas in the preceding data sets is probably more reliable than the numbers themselves. Due to the many deficiencies in current knowledge, island summaries (Table 3.1.1) are subject to considerable uncertainty and include a fair amount of conjecture. The Flora Malesiana will eventually provide a more objective basis for comparative assessment, but at present rates of revision a full accounting of the regional flora will not be completed until 2135–2150 (Geesink 1990; Roos 2003). Inter-island patterns of generic endemism in Malesia are generally consistent with currently perceived trends in species diversity. Lists of endemic genera are available for each of the major areas (Johns 1995), though certain allowances for taxonomic uncertainty are necessary when viewing such compilations. The most

................. 16157$

CH12

02-08-07 10:31:54

PS

PAGE 280

Introduction to the Flora of Papua / 281

Figure 3.1.12. Versteegia (Rubiaceae) is a genus of five species endemic to Papuasia. On the mainland of New Guinea only V. cauliflora (shown) can be considered a common species. The rare V. grandifolia, previously known only from Papua Province, was recently found on the PNG side. Scale bar: 2 cm. recent figures for New Guinea (Johns 1993, 1995) have already been significantly altered by contemporary revision, mainly as a result of the growing popularity of DNA sequencing in phylogenetic reconstruction. Among other developments, Delphyodon (Figure 3.1.15), Papuastelma, and Spathidolepis have been transferred to nonendemic genera (i.e., Parsonsia, Sarcolobus, and Dischidia, respectively; see Middleton 1997; Forster 1990, 1991). On the basis of newly available phytomolecular evidence, the monotypic Madangia will require reduction to Hoya (P. Forster, pers. comm.). Petalolophus, Faika (Figure 3.1.16), and Kairoa (Figures 3.1.17, 18) are also susceptible to future reassignments which will remove their endemic status (R. Saunders, pers. comm.; S. Renner, pers. comm.). Irrespective of nomenclatural adjustments however, New Guinea clearly leads Malesia on species and generic measures of floristic uniqueness. To the extent that documentation gains are most likely in poorly surveyed localities with high endemism, Papua will remain the great unknown and high priority target into the foreseeable future.

Environmental Quality Papua is home to some of the most picturesque landscapes on the planet. The province’s biodiversity status is attributable in large part to the presence of an

................. 16157$

CH12

02-08-07 10:32:14

PS

PAGE 281

282 / wayne n . t a k e u c h i

Figure 3.1.13. Versteegia cauliflora. Closer view of the cauline fruits and the fleshy, persistent stipules. Scale bar: 1 cm. impressive range of environments, recalling those normally seen on continental spatial scales and including some of the most biotically significant habitats in the Old World. Arrayed along the Central Divide are the highest mountains in the Asia-Pacific region, their summits crowned by the only equatorial glaciers in Malesia (Whitmore 1975; Mangen 1993). The province is endowed with unique assets of potential World Heritage quality (e.g., the ultrabasic islands and stacks of the Raja Ampat, the Vogelkop’s karst ranges, and tropical alpine lakes; Figures 3.1.19–21). In marked contrast to Western Malesia, this natural history legacy is intact and comparatively free of the human influences affecting other island ecosystems. Papua’s population density of 3.9 persons/km2 is the lowest in Malesia (Table 3.1.2). The anthropogenic pressures on environmental quality are thus far less severe than in other areas (Figure 3.1.22). On the basis of their collecting densities, the documentation urgency for Papua, Sulawesi, and Sumatra should be similar, but most of the forest ecosystems on the latter two islands have already been considerably impacted by economic development. In Sumatra, 65–80% of the lowland forest has disappeared (Whitten et al. 2000). At projected rates of forest depletion, the Sumatran lowland forest could be eradicated by 2010 (Conservation International 2003). Based on 1982 figures,

................. 16157$

CH12

02-08-07 10:32:38

PS

PAGE 282

Introduction to the Flora of Papua / 283

Figure 3.1.14. The endemic genus Chlaenandra (Menispermaceae) consists of a single species, C. ovata, found in lowland forests throughout New Guinea. The woody vine produces large clusters of glaucous fruits along the main stem (arrow). Cassowaries are avid consumers of the red fruits despite the presence of an endocarp covered with long spines. The individual drupes are approximately 60 mm long by 55 mm wide.

................. 16157$

CH12

02-08-07 10:33:08

PS

PAGE 283

284 / wayne n . t a k e u c h i

Figure 3.1.15. The distinctive fruits of Delphyodon oliganthus (Apocynaceae), a species recently transferred to Parsonsia. Scale bar: 1 cm.

................. 16157$

CH12

02-08-07 10:33:55

PS

PAGE 284

Introduction to the Flora of Papua / 285

Figure 3.1.16. Faika villosa was assigned by Philipson (1985) to a monotypic and presumably endemic genus. The corky, deeply furrowed stem (arrow) is a distinctive feature also present in Kairoa. Originally known only from northern Papua, the species/genus was recently recorded from the April River in PNG. 54% of the lowland forests in Sulawesi have similarly been removed (Whitten et al. 2002). The present values for forest destruction, though unknown, are certainly higher. In contrast to the western islands, Papuan habitats are in a markedly superior state of preservation. The remaining forest cover was estimated at over 90% by the Biodiversity Action Plan for Indonesia (Ministry of National Development Planning 1993), and at 75–80% by Supriatna et al. (1999). Although the lack of effective planning tools (e.g., forest classification and mapping systems) hampers precise assessment, Papuan ecosystems have obviously experienced less alteration than other parts of Malesia. The relationship between environmental quality and population density is apparent in per capita estimates of remaining forest: Sumatra 0.7 ha/person, Sulawesi 1.0 ha/person, and Papua 35.1 ha/person (Whitten et al. 2002). The Papuan environment is the last great wilderness in Indonesia, and one of the world’s largest remaining repositories for tropical plant life.

Biogeography New Guinea is positioned at the critical junction between the Asian and Australasian bioregions, a circumstance which prompted its description as a ‘‘keystone in Pacific botany’’ (Steenis 1950). The island’s strategic location has encouraged the

................. 16157$

CH12

02-08-07 10:34:11

PS

PAGE 285

286 / wayne n . t a k e u c h i

Figure 3.1.17. The genus Kairoa (Monimiaceae) consists of the single species K. suberosa, an endemic of NE New Guinea (Philipson 1980). The stem (at left) is often unbranched. Scale bar: 2 cm. development of a heterogeneous vegetation composed of Laurasian, Gondwanan, and neo-endemic elements. The complex interplay between biogeographically distinct components underpins much of today’s botanical diversity. Beginning with the work of Lam (1934), numerous investigators have attempted to explain the manifold and confusing features of the New Guinea flora. In retrospect, many of the earlier efforts were hindered by a faulty understanding of the island’s exceedingly complex geological history. The biotic connections to this geological dynamic are now becoming much clearer. A seminal event in the development of current perspectives was the reinterpretation of New Guinea’s tectonic structure by Pigram and Davies (1987). Although the southern half of the island is a stable extension of the Australian craton, the northern districts were interpreted as a dynamic collage of 32 discrete terranes (⬃microplates), assembled during four distinct phases of accretion. One of the most widely cited contributions in Papuasian science, the terrane-collage paradigm has inspired a host of applications by biogeographers and systematists, as exemplified in recent syntheses by van Welzen (1997), van Welzen and Turner (2001), and Heads (2001, 2003). Clear relationships have been established between patterns of floristic speciation and the phases of terrane accretion (e.g., Heads

................. 16157$

CH12

02-08-07 10:34:36

PS

PAGE 286

Introduction to the Flora of Papua / 287

Figure 3.1.18. Kairoa suberosa. The fruiting facies is indistinguishable from Steganthera. Scale bar: 2 cm.

Figure 3.1.19. One of New Guinea’s most aesthetic and pristine environments, the Raja Ampat has thousands of limestone platforms, steep-sided stacks, and ultrabasic outcrops. Floristic diversity is suppressed by low rainfalls and seasonal drought, but endemism is high. The archipelagic complex near Misool (shown) is of probable Gondwanan origin. 2001, 2003). Through this connection to geological processes, Papua has emerged as an area of particular significance and interest. While most of the terranes are oceanic in origin, those of Papua are primarily continental and were emplaced in a separate series of accretionary events. The Vogelkop (including parts of the Misool terrane) is a Gondwanan microcontinent (Pigram and Panggabean 1984; Pigram and Davies 1987).

................. 16157$

CH12

02-08-07 10:35:05

PS

PAGE 287

288 / wayne n . t a k e u c h i

Figure 3.1.20. Kawe. Located near the westernmost margin of Papua Province, Kawe is a remote island with an ultrabasic woodland of Ploiarium sessile, Exocarpos latifolius, Gymnostoma rumphianum, Decaspermum bracteatum, Ixonanthes reticulata, and Myrsine rawacensis. Livistona sp. nov. is locally common on the valley floors and in near-shoreline areas.

Figure 3.1.21. Waigeo Island. Foreground: the ultrabasic habitat of the endemic scrubland. One of the classic ultrabasic environments in New Guinea, the coastal parts of the serpentine are primarily an early-stage fire succession. Background: the placid waters of Fofak Bay.

................. 16157$

CH12

02-08-07 10:35:33

PS

PAGE 288

Introduction to the Flora of Papua / 289

Table 3.1.2. Human population densities in Malesia Region Papua

Density (individuals/km2) 3.9

Source 1990 Indonesian census

Papua New Guinea

11

2000 Papua New Guinea census

Kalimantan

17

1990 Indonesian census

Sulawesi

66

1990 Indonesian census

Sumatra

77

1990 Indonesian census

Philippines

220

US Library of Congress (1990)

Java

799

1990 Indonesian census

Source: 1990 Indonesian census data from www.indonesianembassy.org.uk; 2000 Papua New Guinea census data from National Statistical Office of Papua New Guinea.

Figure 3.1.22. Interior perspective of a mesic forest in south Misool. The arborescent taxa are mainly Flindersia amboinensis, F. laevigata var. heterophylla, Homalium foetidum, Intsia bijuga, I. palembanica, Jagera javanica ssp. javanica, Pometia pinnata, and Vatica rassak. Maniltoa spp. (M. plurijuga and M. schefferi) are often in the understory. The Misool forests are generally in pristine condition except for areas near human settlements. In New Guinea as a whole, the western and eastern ends of the island are centers for narrow endemism, having exceptional numbers of taxa known only from their type localities. At these geographic antipodes, high levels of biotic endemism are apparently related to unusual features of the paleohistorical environment. However, the true extent of such connections is obscured by the poor documentation of surrounding districts. In any consideration of the inadequately known areas of Malesia, Papua is arguably the most scientifically cost-effective and desirable geographic target. Based on the foregoing criteria of endemism, environmental quality, and biogeography, there is no other location where future inquiry has the promise of being as rewarding.

................. 16157$

CH12

02-08-07 10:35:43

PS

PAGE 289

290 / wayne n . t a k e u c h i

General Character of the Vegetation Despite its high diversity, the Papuan flora is defined by a relatively limited range of families. In lowland environments (below 1,000 m) the most important groups in terms of frequency, richness, or visual prominence, are the Anacardiaceae, Annonaceae, Arecaceae, Burseraceae, Combretaceae, Euphorbiaceae (Phyllanthaceae), Fabaceae sensu lato, Elaeocarpaceae, Flacourtiaceae (Salicaceae), Malvaceae sensu lato, Meliaceae, Monimiaceae, Moraceae, Myristicaceae, Rubiaceae, and Sapindaceae. Unlike Western Malesia, the Dipterocarpaceae is a minor family; only Anisoptera (1 sp.), Hopea (9 spp.), and Vatica (1 sp.) are present in Papua (Ashton 1982). In montane habitats the most important groups are the cryptogams, gymnosperms, and (among angiosperms) Araliaceae, Cunoniaceae, Ericaceae, Fagaceae, Gesneriaceae, Lauraceae, Myrsinaceae (Figures 3.1.23, 24), Myrtaceae (Figures 3.1.25–28), Orchidaceae, Theaceae, Urticaceae, Winteraceae, and Zingiberaceae. Diversity falls sharply at elevations above 2,500 m even in the microtherm families (see discussion in Whitmore 1975). However endemism is highest in the montane zone as a result of environmental change induced by rapid rates of geological uplift (Heads 2001, 2003).

Figure 3.1.23. Fittingia (Myrsinaceae) is a genus of seven species restricted to New Guinea (Hu Chi Ming 2001; Sleumer 1988). The most frequently collected species is F. tubiflora, shown as fresh cuttings from the little-known Karius Range. The juicy fruits are translucent-pink or red. Scale bar: 15 mm.

................. 16157$

CH12

02-08-07 10:36:04

PS

PAGE 290

Introduction to the Flora of Papua / 291

Figure 3.1.24. Fittingia tuberculata occurs primarily in Papua Province, with only one record originating from PNG. The understory shrub is locally common in forests of the Bomberai Peninsula. Drupes are opaquely red and ca 20 mm wide. Scale bar: 1 cm. In any account of the Papuan flora, the ferns (pteridophytes) and orchids require particular comment because of their extraordinary richness. With an estimated 3,000 species (Parris, Chapter 3.4), New Guinea has nearly 30% of the world’s fern diversity and more than twice as many pteridophytes as the rest of Malesia combined (see Roos 1996). Equally remarkable are the 2,800 orchid species, of which an astonishing 95% are endemic (Chapter 3.6). There are only 13 genera in New Guinea with ca 100 or more species, and the Orchidaceae has four of them (Table 3.1.3). Ironically, although ferns and orchids collectively account for one-fourth of the island’s floristic diversity, they rarely receive attention from forestry surveyors or even from most collectors. The 35 new orchids recently discovered by E. de Vogel and A. Schuiteman (pers. comm., from their CD-project fieldwork) are an indication of how much remains to be done. Future collectors can also take note of the surprising lichen inventory for New Guinea (well above 2,000 species, of which only ca 1,200 have been reliably named; Chapter 3.2). The opportunities for contributions from the field extend across the entire taxonomic spectrum. In the fern families with completed revisions, there are no Papuasian genera

................. 16157$

CH12

02-08-07 10:36:28

PS

PAGE 291

292 / wayne n . t a k e u c h i

Figure 3.1.25. Octamyrtus (Myrtaceae) is a genus of six species confined to the New Guinea region. The common O. pleiopetala is a characteristic component of montane forests. Arrows: flower buds. with more than 100 species. The genus Cyathea, represented by 78 mainland species (86 if the offshore islands are included; Holttum 1963) is the most speciose. Grammitis (64 spp.) is a distant second (Parris 1983). Sphaerostephanos has 64 species if nearby islands are included; the mainland has 58 (Holttum 1981). Selaginella is probably the largest of the groups awaiting revision (55 spp. in New Guinea and the Bismarck Archipelago; Camus 1997). Lichens generally consist of small- to moderate-size genera, but Pertusaria (presently with 70 spp.) may reach the century mark after existing collections are given a full accounting (H. Sipman, pers. comm.). Prior to its reorganization into smaller satellites, Parmelia sensu lato would have included ca 150 New Guinea species (H. Sipman, pers. comm.). For angiosperms, Syzygium and Ficus are the only large (100) genera composed primarily of arborescent taxa (in Psychotria, only P. chrysantha, P. leiophloea, and P. micralabastra attain canopy size). Myristica has 98 species in New Guinea, of which 74 have been discovered or described within the last ten years (de Wilde 1995, 1998, 2000). Because of the restricted range endemism in this genus, it is very likely that future Papuan exploration will move the generic tally substantially upward. With ca 80 species (M. Coode, Chapter 3.6 Angiosperms: Elaeocarpaceae), Elaeocarpus is the next richest tree group.

................. 16157$

CH12

02-08-07 10:36:50

PS

PAGE 292

Introduction to the Flora of Papua / 293

Figure 3.1.26. Flowering branchlets of Octamyrtus pleiopetala. The white corolla in this specimen is atypical, the species usually having red or pink petals. Most of the larger genera consist of understory or epiphytic plants. Schefflera is the colossus among woody epiphytes with 180 species (241 taxa including those at subspecific rank), but the genus is presently scheduled for dismemberment into the following segregates (D. Frodin, pers. comm.): ‘‘Brassaia’’ (27); ‘‘Cephaloschefflera’’ (4); ‘‘Pagiophylla’’ (6, all restricted to Papua); ‘‘Parapanax’’ (5); ‘‘Papuoschefflera’’ (191); ‘‘Elmeri’’ (2); ‘‘Heptapleurum’’ (2); and one species of unknown affinity. Conservatively estimated at 120 species (B.L. Burtt, pers. comm.), Cyrtandra is unusual in being overwhelmingly composed of undescribed plants. Currently with 61 species, Alpinia does not approach the century mark (M. Newman, pers. comm.), in spite of its importance and apparent diversity in Papuasian understories. Among vining plants, Freycinetia is the principal genus of note. In 1999, New Guinean Freycinetia included at least 79 species (Huynh, pers. comm.), but the conspectus was later expanded in a series of publications (Huynh 2000, 2001, 2002a,b,c, 2003), bringing the present total to 134. Most of the newer species are known only from the type locality. The dominance of nontree taxa in the New Guinea flora is reflected in elevational patterns of beta diversity. Although tree counts generally decline with elevation (Ashton 2001; Gentry 1988), the total richness on Papuasian altitudinal gradients is highest in the submontane zone (when including all taxa, irrespective

................. 16157$

CH12

02-08-07 10:37:13

PS

PAGE 293

294 / wayne n . t a k e u c h i

Figure 3.1.27. Octamyrtus glomerata in swampy forest near the Saengga River, Papua. The species is found at lower elevations than O. pleiopetala. of growth form), mainly because of the increasing contribution from epiphytes and shrubs with elevation (Takeuchi and Golman 2001). Similar patterns have been reported for the neotropics (e.g., Henderson et al. 1991).

Taxonomic Summaries In the remainder of this section, a series of taxonomic and floristic summaries have been prepared by invited authorities. A general overview of the flora is presented in a way that directs attention to major gaps in our present knowledge. It is reasonable to conclude that no family or plant group has been adequately inventoried. Political boundaries have little correlation to biotic patterns, and this is certainly true for Eastern Malesia, where the numerous relationships among geographic centers make it impractical to consider Papua in isolation. The severe deficiencies in floristic documentation effectively discourage such consideration anyway. Most of the following sections are necessarily drawn from the island of New Guinea as a whole. As will be evident from the family summaries, the taxonomic compilations are highly uneven, the inevitable outcome of a poor information base. A number of important groups have not been given any attention at all. Most of these, however, have already been treated by the Flora Malesiana, and the Papuan taxa can be extracted from those revisions. Among such excluded, but already revised, families are the Burseraceae (Leenhouts 1956), Cyperaceae (Kern 1974;

................. 16157$

CH12

02-08-07 10:37:24

PS

PAGE 294

Introduction to the Flora of Papua / 295

Figure 3.1.28. Flowering axils of Octamyrtus glomerata. Petals are bright yellow. Kern and Nooteboom 1979); Fagaceae (Soepadmo 1972); Flacourtiaceae ( Salicaceae; Sleumer 1954); legumes (Caesalpiniaceae: Ding Hou et al. 1996; Mimosaceae: Nielsen 1992); and Meliaceae (Mabberley et al. 1995). The most conspicuous omissions are the Rubiaceae and Urticaceae, families either in taxonomic disarray or under revision. It is clear that the need and opportunities for future inquiry are profound. This perception, more than any other, is a common thread through all botanical commentaries on Papua. Even after more than a century of exploration, a satisfactory understanding of the flora is still many generations away.

Literature Cited Archbold, R., A.L. Rand, and L.J. Brass. 1942. Results of the Archbold Expeditions no. 41. Summary of the 1938–1939 New Guinea expedition. Bulletin of the American Museum of Natural History 79: 197–288. Ashton, P. 1982. Dipterocarpaceae. Flora Malesiana ser. I, 9 (2): 237–552. Ashton, P. 1989. Sundaland. Pp. 91–99 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and

................. 16157$

CH12

02-08-07 10:37:37

PS

PAGE 295

296 / wayne n . t a k e u c h i

Table 3.1.3. The largest (⭓100 spp.) plant genera in New Guinea

Genus

Estimated number of species

Source

Bulbophyllum

600

Schuiteman and de Vogel, Ch. 3.6 Orchidaceae

Dendrobium

400

Schuiteman and de Vogel, Ch. 3.6 Orchidaceae

Syzygium

200

Craven, Ch. 3.6 Myrtaceae

Schefflera s.l.

180

D. Frodin pers. comm.; see text; up to 200 in Papuasia

Ficus

151

Weiblen, Ch. 3.6 Moraceae

Rhododendron

150

L. Craven pers. comm.; 155 fide Sleumer 1966

Freycinetia

134

See text

Vaccinium

132

Sleumer 1967

Phreatia

130

Schuiteman and de Vogel, Ch. 3.6 Orchidaceae

Cyrtandra

120

B. Burtt pers. comm; at least 100 undescribed spp., Q. Cronk pers. comm.

Psychotria Glomera Myristica

ca. 120 100 ca. 100

Possibly 200–300 species fide Sohmer 1988 Schuiteman and de Vogel, Ch. 3.6 Orchidaceae The actual number of species is 98 but future additions are likely

Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. Ashton, P. 2001. Ecology and conservation of Malesian flora. Pp. 253–258 in Saw, L.G., L.S.L. Chua, and K.C. Khoo (eds.) Proceedings of the Fourth International Flora Malesiana Symposium 1998. Ampang Press Sdn, Kuala Lumpur, Malaysia. Borrell, O.W. 1989. An Annotated Checklist of the Flora of Kairiru Island, New Guinea. Privately published. Brass, L.J. 1941. The 1938–39 expedition to the Snow Mountains, Netherlands New Guinea. Jour. Arn. Arb. 22 (2): 271–295, 297–342. Campbell, D.G. 1989. The importance of floristic inventory in the tropics. Pp. 5–30 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. Camus, J.M. 1997. The genus Selaginella (Selaginellaceae) in Malesia. Pp. 59–69 in Dransfield, J., M.J.E. Coode, and D.A. Simpson (eds.) Plant Diversity in Malesia III. Proceedings of the Third International Flora Malesiana Symposium 1995. Royal Botanic Gardens, Kew. Collins, N.M., J.A. Sayer, and T.C. Whitmore (eds.). 1991. The Conservation Atlas of Tropical Forests, Asia and the Pacific. Macmillan, London. Conn, B.J. 1994. Documentation of the flora of New Guinea. Pp. 123–156 in Peng, C.-I., and C.H. Chou (eds.) Biodiversity and Terrestrial Ecosystems. Academia Sinica Monograph 14. Institute of Botany, Taipei.

................. 16157$

CH12

02-08-07 10:37:38

PS

PAGE 296

Introduction to the Flora of Papua / 297 Conn, B.J. (ed.). 1995. Handbooks of the Flora of Papua New Guinea. Vol. 3. Melbourne University Press, Carlton, Victoria. Conservation International. 2003. Ecosystem profile: Sumatra, Sundaland. URL: http:// www.cepf.net/xp/cepf/where_we_work/sundaland/full_strategy.xml. Coode, M.J.E., S.C. Hinchclifte, and C.J. Marsden. 1997. Checklist of the Flowering Plants of N.E. Kepala Burung (Vogelkop), Irian Jaya, Indonesia. Royal Botanic Gardens, Kew. Craven, I., and Y. de Fretes. 1987. The Arfak Mountains Nature Conservation Area, Irian Jaya: Management Plan, 1988–1992. Directorate General of Forest Protection and Nature Conservation (PHPA), Ministry of Forestry, Republic of Indonesia, Bogor. de Vogel, E. 1989. Sulawesi (Celebes). Pp. 108–112 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. de Wilde, W.J.J.O. 1989. Sumatra. Pp. 103–107 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. de Wilde, W.J.J.O. 1995. Census of Myristica (Myristicaceae) in New Guinea anno 1994. Blumea 40 (2): 237–344. de Wilde, W.J.J.O. 1998. The myrmecophilous species of Myristica (Myristicaceae) from New Guinea. Blumea 43: 165–182. de Wilde, W.J.J.O. 2000. Myristicaceae. Flora Malesiana ser. I, 14: 1–632. Ding, H., K. Larsen, and S.S. Larsen. 1996. Caesalpiniaceae (Leguminosae-Caesalpinioideae). Flora Malesiana ser. I, 12 (2): 409–730. Donnelly, R., D. Neville, and P.J. Mous. 2004. Report on a Rapid Ecological Assessment of the Raja Ampat Islands, Papua, Eastern Indonesia, Oct. 30–Nov. 22, 2002. The Nature Conservancy Southeast Asia Center for Marine Protected Areas, Bali. Erftemeijer, P.L., G.R. Allen, and Zuwendra. 1989. Preliminary Resource Inventory of Bintuni Bay and Recommendations for Conservation and Management. Prepared for Asian Wetlands Bureau and Indonesia Directorate General of Forest Protection and Nature Conservation. AWB-PHPA Report. Forster, P.I. 1990. Notes on Asclepiadaceae, 2. Austrobaileya 3: 273–289. Forster, P.I. 1991. A taxonomic revision of Sarcolobus R.Br. (Asclepiadaceae: Marsdenieae) in Australia and Papuasia. Austrobaileya 3: 335–360. Frodin, D.G. 1988. The natural world of New Guinea: hopes, realities, and legacies. Pp. 89–138 in MacLeod, R., and P.F. Rehbock (eds.) Nature in Its Greatest Extent: Western Science in the Pacific. University of Hawaii Press, Honolulu. Frodin, D.G. 1990. Botanical progress in Papuasia. Pp. 235–247 in Baas, P., K. Kalkman, and R. Geesink (eds.) The Plant Diversity of Malesia, Proceedings of the Flora Malesiana Symposium Commemorating Prof. Dr. C.G.G.J. van Steenis. Kluwer Academic Publishers, Norwell, Massachusetts. Frodin, D.G., and J.L. Gressitt. 1982. Biological exploration in New Guinea. Pp. 87–130 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk (Monogr. Biol. 42), The Hague. Geesink, R. 1990. The general progress of Flora Malesiana. Pp. 11–16 in Baas, P., K. Kalkman, and R. Geesink (eds.) The Plant Diversity of Malesia, Proceedings of the Flora Malesiana Symposium Commemorating Prof. Dr. C.G.G.J. van Steenis. Kluwer Academic Publishers, Norwell, Massachusetts. Gentry, A. 1988. Changes in plant community diversity and floristic composition on environmental and geographical gradients. Ann. Missouri Bot. Gard. 75: 1–34.

................. 16157$

CH12

02-08-07 10:37:38

PS

PAGE 297

298 / wayne n . t a k e u c h i Gibbs, L.S. 1917. Dutch North-West New Guinea. A Contribution to the Flora and Phytogeography of the Arfak Mountains. Taylor and Francis, London. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. Pp. 99–124 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys, Leiden. Hammermaster, E.T., and J.C. Saunders. 1995a. Forest Resources and Vegetation Mapping of Papua New Guinea. PNGRIS Publ. no. 4, CSIRO and AIDAB, Canberra. Hammermaster, E.T., and J.C. Saunders. 1995b. Forest Resources and Vegetation Mapping of Papua New Guinea. 1:250,000 vegetation map overlays separately issued as working copies to PNGRIS Publ. no. 4, CSIRO and AIDAB, Canberra. Heads, M.J. 2001. Regional patterns of biodiversity in New Guinea plants. Bot. J. Linn. Soc. 136: 67–73. Heads, M.J. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Proceedings of the 5th International Flora Malesiana Symposium 2001. Telopea 10 (1): 311–449. Henderson, A., S.P. Churchill, and J.L. Luteyn. 1991. Neotropical plant diversity: are the northern Andes richer than the Amazon Basin? Nature 351: 21–22. Henty, E.E. (ed.). 1981. Handbooks of the Flora of Papua New Guinea, Vol. 2. Melbourne University Press, Melbourne. Holttum, R.E. 1963. Cyatheaceae. Flora Malesiana ser. II, 1 (2): 65–176. Holttum, R.E. 1981. Thelypteridaceae. Flora Malesiana ser. II, 1 (5): 331–599. Hope, G.S. 1976. The vegetation of Mt Jaya. Pp. 113–172 in Hope, G.S., J.A. Peterson, I. Allison, and U. Radok (eds.) The Equatorial Glaciers of New Guinea. A.A. Balkema, Rotterdam. Hu C.M. 2001. A new species of Fittingia (Myrsinaceae) from New Guinea. Blumea 46 (1): 189–191. Huynh, K.-L. 2000. The genus Freycinetia (Pandanaceae) in New Guinea (part 3). Candollea 55: 299–322. Huynh, K.-L. 2001. The genus Freycinetia (Pandanaceae) in New Guinea (part 5). Bot. Jahrb. Syst. 123 (3): 321–340. Huynh, K.-L. 2002a. The genus Freycinetia (Pandanaceae) in New Guinea (part 4). Blumea 47 (3): 513–536. Huynh, K.-L. 2002b. The genus Freycinetia (Pandanaceae) in New Guinea (part 6). Candollea 57: 55–65. Huynh, K.-L. 2002c. The genus Freycinetia (Pandanaceae) in New Guinea (part 7). Bot. Jahrb. Syst. 124 (2): 151–161. Huynh, K.-L. 2003. The genus Freycinetia (Pandanaceae) in New Guinea (part 8). Bot. Jahrb. Syst. 125 (1): 73–83. Johns, R.J. 1987. The Flowering Plants of Papuasia. Dicotyledons. Part 1: Magnoliidae. PNG University of Technology Forestry Department, Lae. Johns, R.J. 1988. The Flowering Plants of Papuasia. Dicotyledons. Part 2: Hamamelidae. Christensen Research Institute Publication No. 1. Christensen Research Institute and the PNG University of Technology Forestry Department, Lae. Johns, R.J. 1989. The Flowering Plants of Papuasia. Part 3: the Caryophyllidae. Christensen Research Institute Publication No. 2. Christensen Research Institute and the PNG University of Technology Forestry Department, Lae. Johns, R.J. 1993. Biodiversity and conservation of the native flora of Papua New Guinea. Pp. 15–75 in Beehler, B. (ed.) Papua New Guinea Conservation Needs Assessment Report. Vol. 2. PNG Dept. of Environment and Conservation, Boroko.

................. 16157$

CH12

02-08-07 10:37:39

PS

PAGE 298

Introduction to the Flora of Papua / 299 Johns, R.J. 1995. Endemism in the Malesian flora. Curtis’s Botanical Magazine 12 (2): 95–110. Kalkman, C. 1963. Description of vegetation types in the Star Mountains region, West New Guinea. Nova Guinea, Botany 15: 247–261. Kern, J.H. 1974. Cyperaceae. Flora Malesiana ser. I, 7 (3): 435–753. Kern, J.H., and H.P. Nooteboom. 1979. Cyperaceae-II. Flora Malesiana ser. I, 9 (1): 107–187. Lam, H.J. 1934. Materials toward a study of the flora of the island of New Guinea. Blumea 1 (1): 115–159. Laumonier, Y. 1997a. The Vegetation and Physiography of Sumatra. Geobotany 22. Kluwer Academic Publishers, Dordrecht. Laumonier, Y. 1997b. International map of the vegetation and of environmental conditions (3 color maps at 1:1,000,000 scale). Institut de la Carte Internationale du Tapis Ve´ge´tal and SEAMEO-BIOTROP, Toulouse and Bogor. Laumonier, Y., A. Gadrinab, and Purnajaya. 1983. International map of vegetation and environmental conditions: Southern Sumatra, scale 1:1,000,000. Institut de la Carte Internationale du Tapis Ve´ge´tal and SEAMEO-BIOTROP, Toulouse and Bogor. Laumonier, Y., Purnajaya, and Setiabudi. 1986. International map of vegetation and environmental conditions: Central Sumatra, scale 1:1,000,000. Institut de la Carte Internationale du Tapis Ve´ge´tal and SEAMEO-BIOTROP, Toulouse and Bogor. Laumonier, Y., Purnajaya, and Setiabudi. 1987. International map of vegetation and environmental conditions: Northern Sumatra, scale 1:1,000,000. Institut de la Carte Internationale du Tapis Ve´ge´tal and SEAMEO-BIOTROP, Toulouse and Bogor. Leenhouts, P.W. 1956. Burseraceae. Flora Malesiana ser. I, 5 (2): 209–296. Mabberley, D.J., C.M. Pannell, and A.M. Sing. 1995. Meliaceae. Flora Malesiana ser. I, 12 (1): 1–407. Mack, A., and L.E. Alonso (eds.). 2000. A Biological Assessment of the Wapoga River Area of Northwestern Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14, Conservation International, Washington, D.C. Madulid, D.A. 1982. Plants in peril. Filipinas J. Sci. Culture 3: 8–16. Mangen, J.M. 1993. Ecology and Vegetation of Mt Trikora, New Guinea (Irian Jaya/ Indonesia). Travaux Scientifiques du Muse´e National d’Histoire Naturelle de Luxembourg, Luxembourg. Middleton, D.J. 1997. A revision of Parsonsia R. Br. (Apocynaceae) in Malesia. Blumea 42: 191–248. Middleton, D. 2003. Progress on the flora of Thailand. Proceedings of the 5th International Flora Malesiana Symposium 2001. Telopea 10 (1): 33–42. Ministry of National Development Planning. 1993. Biodiversity Action Plan for Indonesia. Vol. 1. National Development Planning Agency, Jakarta, and The World Bank, Washington, D.C. Nielsen, I.C. 1992. Mimosaceae (Leguminosae-Mimosoideae). Flora Malesiana ser. I, 11 (1): 1–226. Pannell, C.M. 1992. A taxonomic monograph of the genus Aglaia Lour. (Meliaceae). Kew Bull. Additional Series XVI. Parris, B.S. 1983. A taxonomic revision of the genus Grammitis Swartz (Grammitidaceae: Filicales) in New Guinea. Blumea 29 (1): 13–222. Peekel, P.G. (trans. E.E. Henty). 1984. Flora of the Bismarck Archipelago for Naturalists. Kristen Press, Madang. Percival, M., and J.S. Womersley. 1975. Floristics and Ecology of the Mangrove Vegetation

................. 16157$

CH12

02-08-07 10:37:39

PS

PAGE 299

300 / wayne n . t a k e u c h i of Papua New Guinea. Botany Bulletin 8, Department of Forests, Division of Botany, Lae. Philipson, W.R. 1980. Kairoa, a new genus of Monimiaceae from Papua New Guinea. Blumea 26 (2): 367–372. Philipson, W.R. 1985. Faika and Parakibara: two new genera of Monimiaceae from Malesia. Blumea 30 (2): 417–423. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. Bureau of Mineral Resources, J. Austr. Geol. Geoph. 10: 193–211. Pigram, C.J., and H. Panggabean. 1984. Rifting of the northern margin of the Australian continent and the origin of some microcontinents in eastern Indonesia. Tectonophysics 107: 331–353. Prance, G.T. 1977. Floristic inventory of the tropics: where do we stand? Ann. Missouri Bot. Gard. 64: 659–684. Rand, A.L., and L.J. Brass. 1940. Results of the Archbold Expeditions no. 29. Summary of the 1936–37 New Guinea expedition. Bulletin of the American Museum of Natural History 77: 341–380. Rappard, F.W., and P. van Royen. 1959. Enige notities over de vegetatie in het gebied van de Wisselmeren [Some notes on the vegetation in the Wissel Lakes area]. Nova Guinea, new ser. 10 (2): 159–176. RePPProT (Regional Physical Planning Program for Transmigration). 1990. The Land Resources of Indonesia: A National Overview. Final Report. Land Resource Department of the Overseas Development Administration, London (Government of UK), and Ministry of Transmigration (Government of Indonesia), Jakarta. Ridley, N.H. 1916. Report on the botany of the Wollaston Expedition to Dutch New Guinea, 1912–13. Transactions of the Linnean Society of London, 2nd ser. Bot. 9 (1): 1–269. Roos, M. 1996. Mapping the world’s pteridophyte diversity—systematics and floras. Pp. 29–42 in Camus, J.M., M. Gibby, and R.J. Johns (eds.) Pteridology in Perspective. Proceedings of the Holttum Memorial Pteridophyte Symposium, Kew, 1995. Royal Botanic Gardens, Kew. Roos, M.C. 2003. Flora Malesiana 1991–2001. What has been achieved: revitalisation, momentum? What next? Proceedings of the 5th International Flora Malesiana Symposium 2001. Telopea 10 (1): 1–10. Ruitenbeek, H.J. 1992. Mangrove Management: An Economic Analysis of Management Options with a Focus on Bintuni Bay, Irian Jaya. School for Resource and Environmental Studies, Dalhousie University. Sidiyasa, K., T.C. Whitmore, I.G.M. Tantra, and U. Sutisna. 1997. Tree Flora of Indonesia. Check List for Irian Jaya. Ministry of Forestry, Forest Research and Development Agency, Forest and Nature Conservation Research and Development Centre, Bogor. Sleumer, H. 1954. Flacourtiaceae. Flora Malesiana ser. I, 5 (1): 1–106. Sleumer, H. 1966. Ericaceae. Flora Malesiana ser. I, 6 (4): 469–668. Sleumer, H. 1967. Ericaceae. Flora Malesiana ser. I, 6 (5): 669–914. Sleumer, H. 1988. The genera Discocalyx Mez, Fittingia Mez, Loheria Merr., and Tapeinosperma Hook. f. (Myrsinaceae) in New Guinea. Blumea 33 (1): 81–107. Soepadmo, E. 1972. Fagaceae. Flora Malesiana ser. I, 7 (2): 265–403. Sohmer, S.H. 1988. The nonclimbing species of the genus Psychotria (Rubiaceae) in New Guinea and the Bismarck Archipelago. Bishop Museum Bull. Bot. 1, Bishop Museum Press, Honolulu.

................. 16157$

CH12

02-08-07 10:37:41

PS

PAGE 300

Introduction to the Flora of Papua / 301 Stevens, P.F. 1989. New Guinea. Pp. 120–132 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. Streimann, H. 1986. Catalogue of the Lichens of Papua New Guinea and Irian Jaya. J. Cramer, Berlin. Supriatna, J., Y. de Fretes, A. Mack, C.P. Yeager, S. Olivieri, J.B. Burnett, I. Wijayanto, S. Suryadi, and A. Suhandi (eds.). 1999. The Irian Jaya Biodiversity Conservation PrioritySetting Workshop. Final Report. Conservation International, Washington, D.C. Takeuchi, W. 2003a. Two new species from the Bomberai Peninsula of Indonesian Papua, New Guinea. Harvard Papers in Botany 7 (2): 467–471. Takeuchi, W. 2003b. A community-level floristic reconnaissance of the Raja Ampat Islands in New Guinea. Sida 20 (3): 1099–1144. Takeuchi, W., and M. Golman. 2001. Botanical documentation imperatives: some conclusions from contemporary surveys in Papuasia. Sida 19 (3): 445–468. Tan, B.C., and J.P. Rojo. 1989. The Philippines. Pp. 44–62 in Campbell, D.G., and H.D. Hammond (eds.) Floristic Inventory of Tropical Countries: The Status of Plant Systematics, Collections, and Vegetation, Plus Recommendations for the Future. New York Botanical Garden, New York. van Balgooy, M.M.J. 2001. Aru, a botanical promise. Pp. 225–232 in Saw, L.G., L.S.L. Chua, and K.C. Khoo (eds.) Proceedings of the Fourth International Flora Malesiana Symposium 1998. Ampang Press Sdn, Kuala Lumpur, Malaysia. van Balgooy, M.M.J., P. Hovenkamp, and P. van Welzen. 1996. Phytogeography of the Pacific—floristic and historical distribution patterns in plants. Pp. 191–214 in Keast, A., and S. Miller (eds.) The Origin and Evolution of Pacific Island Biotas: New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Press, Amsterdam. van Royen, P. 1956. Notes on a vegetation of clay-plains in southern New Guinea. Nova Guinea, new ser. 7 (2): 175–180. van Royen, P. 1960. Sertulum Papuanum 3. The vegetation of some parts of Waigeo Island. Nova Guinea, Botany 10 (5): 25–62. van Royen, P. 1963. Sertulum Papuanum 7. Notes on the vegetation of South New Guinea. Nova Guinea, Botany 13: 195–241. van Royen, P. 1965. Sertulum Papuanum 14. An outline of the flora and vegetation of the Cycloop Mountains. Nova Guinea, Botany 21: 451–469. van Royen, P. 1967. Some observations on the alpine vegetation of Mount Biota (Papua). Acta Bot. Neerl. 15: 530–534. van Steenis, C.G.G.J. van. 1950. Desiderata for future exploration. Malaysian plant collectors and collections. Flora Malesiana ser. I, 1: cvii–cxvi, maps 2 and 3. Noordhoff/ Kolff, Jakarta (repr. 1985, Koeltz, Koenigstein). van Welzen, P.C. 1997. Increased speciation in New Guinea: tectonic causes? Pp. 363–387 in Dransfield, J., M.J.E. Coode, and D.A. Simpson (eds.) Plant Diversity in Malesia III. Proceedings of the Third International Flora Malesiana Symposium 1995. Royal Botanical Gardens, Kew. van Welzen, P.C. and H. Turner. 2001. Vicariance and dispersal in Malesian Sapindaceae: general patterns. Pp. 233–251 in Saw, L.G., L.S.L. Chua, and K.C. Khoo (eds.) Proceedings of the Fourth International Flora Malesiana Symposium 1998. Ampang Press Sdn, Kuala Lumpur. Vink, W. 1965. Botanical exploration of the Arfak Mountains. Nova Guinea, Botany 22: 471–494.

................. 16157$

CH12

02-08-07 10:37:42

PS

PAGE 301

302 / wayne n . t a k e u c h i Whitmore, T.C. 1973. Tree Flora of Malaya, Vol. II. Longman, Malaysia. Whitmore, T.C. 1975. The Rain Forests of South-East Asia. Clarendon Press, Oxford. Whitten, T., S.J. Damanik, J. Anwar, and N. Hisyam. 2000. The Ecology of Sumatra. Ecology of Indonesia Series, Volume 1. Periplus Editions (HK), Singapore. Whitten, T., G.S. Henderson, and M. Mustafa. 2002. Ecology of Sulawesi. Ecology of Indonesia Series, Vol. 4. Periplus Editions (HK), Singapore. Wollaston, A.F.R. 1914. An expedition to Dutch New Guinea. Geogr. Journ. 43 (3): 248–273. Womersley, J.S. (ed.). 1978. Handbooks of the Flora of Papua New Guinea, Vol. 1. Melbourne University Press, Melbourne.

................. 16157$

CH12

02-08-07 10:37:42

PS

PAGE 302

3.2. Lichen Biodiversity in New Guinea harrie sipman and andre´ aptroot i c he n s , or more technically, ‘‘lichenized fungi,’’ differ from other fungi in that they are associated with living algae. They obtain photosynthetic products from these algae, resulting in an association resembling an autotrophic organism. As an adaptation to this way of life they do not produce mycelia in their substrate, but instead form compact structures called thalli, which attach to the outside of the substrate and are exposed to ambient light. Consequently their affinity with the fungi was long unclear and they were treated as an independent class of organisms, the lichens. Now it has become clear that they do not form a natural group, but comprise representatives from several groups of fungi, including basidiomycetes and ascomycetes, which have in common only their means of obtaining photosynthetic products. Three major morphological types are distinguished in lichens, based on thallus structure. The simplest, crustose, type forms a crust over its substrate. The foliose type is in less direct contact with its substrate, but is often attached via specialized filaments (rhizines). The third type, fruticose, is attached to the substrate only via a narrow base, and most of the thalli stand away from the substrate like branches of a shrub. These thallus types have developed independently in several groups of lichenized fungi. Reproductive structures more clearly reflect phylogenetic affinities than do thallus types. Lichens are most conspicuous and most well studied in cold areas. However, recent research indicates that this is probably because the relative scarcity of vascular plants and the large size of lichens renders lichens more noticeable in cold areas. Lichen diversity is probably not lower in tropical forests than in more temperate areas. New Guinea has a rich lichen flora (Figure 3.2.1), which, fortunately, has received considerable attention recently, including Lambley’s (1991) overview of the macrolichens of Papua New Guinea. Currently ca 1,150 lichen species (excluding ca 75 lichenicolous fungi) are known from the area (Aptroot et al. 2002). This figure is still very incomplete and does not include many species that are represented in existing collections but are not yet completely identified due to a lack of monographic revisions. The total species number is probably well over 2,000, on the same order of magnitude as temperate, alpine, or arctic areas such as Great Britain and Ireland (Coppins 2002), Sweden and Norway (Santesson et al. 2004), and Austria (Hafellner and Tu¨rk 2001). However, the flora seems to have a larger share of endemic species in New Guinea than in these temperate areas. The available information on the lichen flora of New Guinea comes mostly from Papua New Guinea, notably the surroundings of Port Moresby, Lae, Madang, and the PNG Highlands. The Indonesian side of the island is very little

L

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

303

................. 16157$

CH13

02-08-07 10:28:40

PS

PAGE 303

304 / harrie sipman & andre´ aptroot

Figure 3.2.1. Tree trunk in the alpine zone of Mt Wilhelm covered with lichens.

................. 16157$

CH13

02-08-07 10:28:49

PS

PAGE 304

Lichen Biodiversity in New Guinea / 305

known, but the available information suggests that Papuan lichen biodiversity is very similar to that of Papua New Guinea.

The Lichenological Exploration of New Guinea The lichens of New Guinea escaped scientific attention for a long time, no doubt as a consequence of a general lack of botanical interest in the island. Until about 1965 only a few hundred specimens were available, mostly provided by general collectors who were not knowledgeable about lichen diversity. The material came mainly from the eastern part of the north coast, where it was collected during the short period of German colonization (for details, see Mattick 1942 and Streimann 1986). The results were summarized by Szatala (1956), who reported a total of 224 species. From 1965 on, interest in New Guinea lichens increased greatly, and many hundreds of specimens were collected by people who either focused on or had a strong interest in lichens. In the late 1960s, W. A. Weber and D. McVean were the first to demonstrate the significance of the New Guinea lichen flora. Their collections, mainly from the Mt Wilhelm area, appeared to contain several remarkable endemics (e.g., Calathaspis devexa, Dibaeis weberi, Megalospora weberi). Larger collections were made in the 1965–1975 period by Kurokawa and Kashiwadani’s expeditions to various parts of the Highlands of Papua New Guinea. Streimann made extensive collections while he was teaching in Bulolo in 1981–1983, and stimulated students to make collections (for results, see Streimann 1986, 1990). Koponen collected numerous lichens during his bryological exploration of the Huon Peninsula in the early 1980s. For further details on the history of lichenological exploration during this period, see Streimann (1986) summarizing the available knowledge for New Guinea in a catalog of 495 species, including much new information, mainly from the Highlands of Papua New Guinea. After 1986, the authors, partly in cooperation with P. Diederich, P. Lambley, and E. Se´rusiaux, made three expeditions to the surroundings of Port Moresby, Lae, Madang, and Goroka/Mt Wilhelm, paying special attention to lichens. As a result, Aptroot and Sipman (1991) added over 112 species, including several new species and even new genera. Aptroot et al (1995, 1997) added another 336 species, while additional species have been described or reported in more scattered publications. Streimann made extensive collections on logging sites on New Britain (Streimann and Sipman 1994). Meanwhile Stenroos (1986–88) presented the first thorough revision of a New Guinea lichen group based on ample collections, the family Cladoniaceae. After that, several other groups were revised: the genus Anzia (Yoshimura et al. 1995); Pertusaria (Archer and Elix e.g., 1998); Stereocaulon and Lobaria (Sipman 1998, 2004) Menegazzia (James et al. 2001); and Parmotrema and Hypotrachyna (Louwhoff and Elix 1999, 2002). In these studies numerous new species (many endemic) were discovered. Further relevant references can be found at http://www.nhm.uio.no/botanisk/lav/RLL/RLL.HTM. Currently some 20,000 specimens are available, housed mainly in the herbaria

................. 16157$

CH13

02-08-07 10:28:50

PS

PAGE 305

306 / harrie sipman & andre´ aptroot

ABL, B, CANB, LAE, LG, TNS, and UPNG. Approximately half of these specimens were included in modern revisions or have been otherwise identified following modern standards, while the other half await such revisions. New Guinea will continue to be a rich source of new species.

Origins of the Lichen Flora Lichens are presumed to be an ancient group of organisms that evolved very slowly. Their current diversity on New Guinea is likely a result of continental movements in the geological past, starting as far back as the Cretaceous period. The following sketch of the development of the lichen flora should be considered speculative, because the lichen flora and its distribution are still very incompletely known, as are the exact extent and elevation of landmasses during the geological past. Moreover, the contribution of long-range dispersal to disjunct lichen ranges is unclear. However, this sketch takes into account all current information on distribution and there is no convincing evidence for other scenarios. The origins of the New Guinea lichen flora seem to go back as far as the Cretaceous, when New Zealand separated from Australia. This is indicated by the lichen flora of high elevations, in particular representatives of the genus Megalospora (Sipman 1983), which are more closely related to those occurring on New Zealand and New Caledonia than to those on the Australian mainland. There seems to be little geological evidence indicating how this element may have reached New Guinea, and it is suggested here that (part of) New Guinea was initially connected with New Caledonia and New Zealand and situated in the southern temperate climate zone. During the Tertiary it must have drifted towards the equator, separating first from New Zealand, and later from New Caledonia. Meanwhile mountains must have been present, which allowed the lichen species to avoid the increasingly warm zonal climate by migrating upwards. The phanerogam genus Nothofagus provides additional support for such a history. This interpretation seems to be in conflict with the current geological view (Hall 1998) that New Guinea originated from submersed ocean floor. That scenario would, however, provide no explanation for the presence of the Megalospora flora. A different floristic element is found at middle elevations, where genera like Anzia (Figure 3.2.2), Lobaria, and Stereocaulon are diverse and rich in endemic species. The same or related species are mainly found in east Asia, and not in Australian mountains or other cool southern regions. This suggests that New Guinea after the Cretaceous did not contact Australia again until recently, and instead contacted east Asia, possibly not directly but via landmasses which later developed into the mountains of the Philippines. The actual species numbers of Anzia and Lobaria on New Guinea appear to exceed those in east Asia and the Philippines, suggesting that these genera have evolved on New Guinea, or at least found a refuge there. New Guinea’s contact with Australia, for which geological evidence is strongest, must have occurred at a time when New Guinea was climatically very similar to

................. 16157$

CH13

02-08-07 10:28:51

PS

PAGE 306

Lichen Biodiversity in New Guinea / 307

Figure 3.2.2. The genus Anzia is particularly well represented in the mountain forests of New Guinea. It has a foliose thallus, while the branchy lichen above it has a fruticose thallus and belongs to the lichen genus Usnea. the present. The contact appears to have affected principally the lowland flora, which is very similar on both sides of the Torres Strait. Because the lowland flora is very similar throughout the Malesian region, New Guinea might have been the steppingstone by which much of this flora entered Australia. It is unclear whether biogeographic delineations such as Wallace’s Line affect the lichen flora. The lichen flora of the Malesian area is still too incompletely known to provide substantial information about whether species present on one side of the line are absent on the other side. The strong endemism is no doubt a result of the geographical isolation of the high mountains of New Guinea. The closest ranges reaching elevations above 4,000 m are on the Asiatic mainland, with the exception of the very isolated and small Mt Kinabalu. An inventory on Mt Kinabalu (Sipman 1993) showed a strong affinity with the New Guinean mountains, although the number of shared species is restricted by the limited alpine surface area on Mt Kinabalu. Areas above 3,000 m are also extremely limited in the Malesian region. In contrast, the high mountains of tropical America and the Asiatic mainland are arranged in long chains which facilitate migration over thousands of kilometers. Consequently, neotropical mountain lichen species tend to have a large range in the northern Andes, often from Bolivia to Costa Rica or Mexico. Conversely, most endemic neotropical lichens occur in lower, older mountain systems, mainly in southeast Brazil and the Guyana Highlands. The high mountains of tropical Africa are geologically young and have a limited lichen flora with few endemic species. Again, endemic species occur mainly on older, lower mountains.

................. 16157$

CH13

02-08-07 10:28:54

PS

PAGE 307

308 / harrie sipman & andre´ aptroot

Ecology As autotrophic organisms, lichens are competitors with the green plants. However, lichens grow very slowly, and where sufficient nutrients and water are available, they are ineffective competitors. Instead, lichens are often found in habitats where their capacity to survive unfavorable periods in dry state supports them: on stable, impenetrable surfaces that do not retain water, such as rock, bark, wood, and leaves. Lichens may also colonize soil when plant growth is suppressed by, for example, intense grazing, a cold climate, or nutrient-poor or acid conditions. Continuously dry conditions are unfavorable, probably because lichens do not grow in a dry state. Rapid changes, such as are caused by human interference, are also unfavorable for lichens.

lichens on rock Rock is the favored substrate of most of the known lichen species in the world. Particularly in arctic and alpine environments, rocks are common and often abundantly covered with lichens. In the tropics, however, rock is much less important as a substrate for lichens; the vigorous growth of plant life and rapid chemical erosion strongly reduce the availability of free rock surface. Only at high elevations does exposed rock becomes more abundant, but there it is often covered by cyanobacteria rather than lichens. Thus rock lichens are not abundant in New Guinea. At low elevations, lichens are found mostly on the remains of raised coral reefs. Species of Porina, Verrucaria, Lithothelium, and Opegrapha occur on shady rocks in open forests. At middle elevations, rock outcroppings often appear along streams, and where the erosive forces are not too strong, crustose and foliose lichens, including Parmeliaceae, Lecanoraceae, Physciaceae, grow, and in shady and humid locations, Porina and Verrucaria are found. Rocks at high elevations, especially above ca 3,500 m, are richer in lichens. Where the rocks are moss-covered, the moss cushions include various foliose or fruticose lichens, including Parmeliaceae, Cladoniaceae, Thamnolia, Leptogium, Lobariaceae, and Stereocaulon. Moss-free rocks, when not covered by cyanobacteria, are covered in lichen species that are also found in temperate zones of the world, including Rhizocarpon geographicum and Euopsis granatina. In general, alpine rock seems poor in lichens, and the genus Umbilicaria, common in alpine situations elsewhere, has not (yet) been found. So far, only rock habitats on Mt Wilhelm have been investigated by lichen experts, and collections from other mountains may yield different lichen floras. New types of free rock surface have developed as a result of human activities, including rock exposures in road banks and other excavations, and the concrete and bricks of buildings. At middle and high elevations, road cuts may show an abundant growth of lichens, in particular of genera like Stereocaulon, Dibaeis, Baeomyces, including endemics such as Stereocaulon pseudomassartianum and Gyalidea multispora. This makes one wonder what the natural habitat of these species

................. 16157$

CH13

02-08-07 10:28:54

PS

PAGE 308

Lichen Biodiversity in New Guinea / 309

might be. Lichens on concrete and brick are not very conspicuous. Endocarpon pusillum is among the most common and is often found on old, low walls in humid situations at low elevations.

lichens on soil More than rock, soil is covered predominantly by higher plants under tropical conditions. Consequently the occurrence of soil lichens in New Guinea is very restricted, particularly at low elevations. No sites of natural open scrub vegetation on white sand, which in the neotropics may harbor a rich assortment of Cladonia species, have been found so far. At middle elevations, where the natural forest is interrupted only by streams, no natural occurrences of soil lichens have been found. Nor even have lichens been found in the dry grasslands of the intermountain valleys, perhaps because the grasslands are frequently burned by people. The situation changes for lichens above the tree line, where the often gravelly or peaty, acid soil is suitable for genera like Cladonia, Peltigera, and Baeomyces, including endemic species such as Baeomyces marginalis. However, species of Cladonia subgenus Cladina, usually climax stage lichens on acidic soil, are rare. Where roads are cut through soft material, soil lichens may grow on the banks at all elevations, though they are most abundant in the mountains. This is where two species of the genus Catapyrenium have been found. Cladonia is otherwise the most conspicuous genus here.

lichens on bark and wood Because New Guinea is largely covered by forest or other vegetation types dominated by woody plants, it is no surprise that most lichens are found on bark. The importance of trees for lichen diversity in the tropics is easily underestimated by people with experience of earth’s temperate zones. In temperate zones, rocks have the highest lichen diversity and the choice of tree species available as habitat for lichens is limited. In contrast, in tropical areas few lichens are found on rocks but many are found on numerous woody plant species. The importance of primary forest, rich in tree species, for lichen diversity in the tropics was first demonstrated by Montfoort and Ek (1990; see also Gradstein 1992). They examined cryptogamic epiphytes in a lowland forest in French Guiana and, after an examination of 32 trees, found a surprising total of 209 lichen species. More significantly, every additional tree species yielded additional lichen species and, given that the region has hundreds of tree species, it can be assumed that many more epiphytic lichens would have been found by examinations of additional tree species. Evidently, many more than 200 lichen species occur on bark in the forest area investigated. This was the first demonstration that the tropical lowlands are not always poor in lichens, but that a properly developed forest may harbor hundreds of lichen species. This finding has since been replicated in other areas. A very significant confirmation was obtained in New Guinea, when Aptroot (1997) observed 173 lichen species on a single tree near Myola, Central Province,

................. 16157$

CH13

02-08-07 10:28:55

PS

PAGE 309

310 / harrie sipman & andre´ aptroot

Papua New Guinea (Figure 3.2.3). This is a world record, more than twice the number found on the most lichen-rich host tree previously known. It can be safely assumed that an undisturbed tropical forest may harbor a minimum of 300–400 bark-inhabiting lichen species. Observations near Madang (see below) suggest that these figures are applicable to New Guinea. This high lichen diversity is most likely due to the microclimate gradient in the forest, from shady and humid near the ground to light and dry in the canopy. The lichen flora near the tree base is usually completely different from that of the canopy. Of particular interest is the lichen flora at middle height. Here the humidity level is raised by evaporation from the adjacent crowns of small trees, and considerable quantities of light are still available, passing between the crowns of the emergent trees. This height zone of the forest provides optimal conditions for the family Thelotremataceae, which has some 100 species throughout New Guinea (Sipman, in prep.). There is also a considerable altitudinal differentiation in the lichen flora of forests. In the lowlands crustose lichens predominate, while the more elaborate foliose and fruticose growth forms are scarce. Conspicuous lowland lichen families include Pyrenulaceae, Trichotheliaceae (mainly near the tree base), Thelotremata-

Figure 3.2.3. Andre´ Aptroot examining a blown-down tree with 173 lichen species, the largest number ever encountered on a single tree.

................. 16157$

CH13

02-08-07 10:29:00

PS

PAGE 310

Lichen Biodiversity in New Guinea / 311

ceae (mainly mid-trunk), Graphidaceae, Trypetheliaceae (mainly canopy). Endemic species in this altitudinal zone are found in, for example, the genus Parmotrema (see Table 3.2.1). Above 1000–1500 m, foliose lichens are more conspicuous. The dominant families here are Collemataceae (mainly near the tree base), Lobariaceae (most in the lower canopy), Parmeliaceae and Physciaceae (in the lightest places). Endemic species are common here, such as those in the genus Anzia (see Table 3.2.1). Under natural conditions hardwood trees do not always fall when they die, but may successively lose their branches and bark, leaving the wooden core of the trunk standing upright for many years. The feature is particularly common in cooler regions and at higher elevations. On New Guinea, various lichens specialized to grow on such decorticated trunks and their branches have been found, particularly members of the family Caliciaceae. To some extent the same lichens may also grow on worked wood of buildings or fences. Trees and shrubs in cultivated areas, such as plantations and parks, may also house considerable numbers of lichen species, and it may be easier to find lichens in these areas because the trees tend to be smaller and more easily accessible than in forest. However, such cultivated habitats contain mostly canopy species and do not match the total richness of natural forest.

lichens on leaves Leaves of tropical trees and shrubs often last for several years and provide an abundant, largely competition-free substrate suitable for lichens. However, growth on leaves requires a short life cycle. A large group of lichens specialized on this substrate, called foliicolous, produce fruiting bodies on tiny thalli often measuring only a few millimeters across and developing within one to three years. Foliicolous lichens form a favorable object for study and are probably better known than those found on other substrates. Important contributors to our knowledge of foliicolous lichens are R. Santesson (e.g., 1952), A. Veˇzda and R. Lu¨cking (e.g., Lu¨cking and Veˇzda 1998), and important contributions on New Guinea representatives were made by Se´rusiaux (e.g., 1997) and Aptroot and Sipman (1993). Research mainly by Lu¨cking in the neotropics has demonstrated that foliicolous lichens grow preferentially in the undergrowth of well developed, tropical lowland rainforest, where up to 200 species may be found within a few square kilometers. Only few, specialized species are found in the canopy. With increasing elevation the number of species decreases, though several species are still common around 2,500 m. Above ca 3,000 m the specialized, obligately foliicolous species are largely replaced by facultatively foliicolous species, which are not restricted to leaves in the forest undergrowth and also occur on shrub twigs. In New Guinea the study of foliicolous lichens is still underway, but first results indicate the same patterns as in the neotropics. Over 100 species are now known, including endemics such as the conspicuous Musaespora coccinea (Aptroot and Sipman 1993) from undergrowth in lowland forest, and the curious Hippocrepidea nigra from scrub on Mt Wilhelm (Aptroot et al. 1997).

................. 16157$

CH13

02-08-07 10:29:00

PS

PAGE 311

312 / harrie sipman & andre´ aptroot

Table 3.2.1. Distribution of the New Guinea lichen genera Anzia and Parmotrema (Parmeliaceae) Altitudinal range (m)

Distribution outside New Guinea

A. afromontana

2,300–3,400

east Africa, South America

A. americana

1,800–3,200

neotropical

A. corallophora

1,300–3,600

endemic

A. endoflavida

1,800–2,300

Sumatra, Java, Mt Kinabalu

A. gregoriana

1,450–3,600

Kinabalu, Sulawesi

A. isidiolenta

2,000–2,400

Philippines, Taiwan

A. isidiosa

1,800–3,750

Sri Lanka, Malaysia, Queensland

A. ornatoides

1,900–3,600

India

Taxon Anzia

A. pseudoangustata

2,300

endemic

A. pseudopustulata

2,400–3,750

endemic

A. pustulata

1,840–3,000

endemic

A. semiteres

1,850–3,750

Java, Sumatra, Mt Kinabalu

P. acrotrychum

1,480–1,600

northern Queensland

P. chinense

2,300–2,600

cosmopolitan

P. cooperi

1,000–1,900

paleotropical, incl. Africa, Australia

P. corniculans

1,200–2,100

Philippines, Java, Laos

P. crinitum

2,500–3,600

cosmopolitan

P. cristiferum

0–1,600

cosmopolitan

P. deflectens

2,200–2,600

east Africa

P. dilatatum

580–1,500

pantropical

P. durumae

1,200–1,420

east Africa

P. elacinulatum

1,450–1,840

northern Queensland

P. fasciculatum

1,600

Parmotrema

P. flaccidifolium P. gardneri P. gloriosum

1,800 0–2,000 600–2,660

P. hypomiltoides P. insuetum P. isidioinsuetum

1,900

neotropical, Africa endemic pantropical endemic neotropical

700–800

endemic

420–1,400

endemic

................. 16157$

CH13

02-08-07 10:29:00

PS

PAGE 312

Lichen Biodiversity in New Guinea / 313 P. kainantum

1,600

endemic

P. kaisenikianum

0–2,300

endemic

P. kurokawianum

40–100

Fiji

P. lambleyi P. maclayanum

30

endemic

1,000–1,600

pantropical

P. madilynae

1,200

Brazil

P. malonprotocetraricum

1,300

endemic

P. mellissii P. menyamyaense P. merrillii P. nanfongense P. naonii P. nilgherrense

800–2,200 1,200

endemic

2,000 5–100

P. praeinsuetum

Taiwan

700 2,000–3,780

P. poolii

Indonesia, Philippines, South America

400–1,600

P. pacificum

P. pigmentosum

endemic

1,200–1,530

P. overeemii

P. permutatum

pantropical

east Africa, Asia, Juan Fernandez Islands Australia, Africa, Indonesia, Tonga Bonin Islands, Mariana Islands, Ryukyu, Solomon Islands

1,300–1,580

pantropical

1,530

Brazil

800–1,900

Australia, Africa, Thailand

1,500

endemic

P. praesorediosum

650–1,300

pantropical

P. rampoddense

600–1,700

pantropical

P. robustum

1,800–2,000

Australasia, Europe, neotropical

P. saccatilobum

0–250

P. sancti-angelii

0–1,800

pantropical

P. sipmanii

sea level

endemic

P. subarnoldii P. subcorallinum

Australia, Southeast Asia, Pacific

1,300–2,000

pantropical

1,650

Indonesia, Taiwan, Mauritius

P. subrugatum

1,300–1,900

pantropical

P. sulphuratum

780–1,300

pantropical

P. tinctorum

580–1,900

cosmopolitan

P. ultralucens

780–1,700

pantropical

P. verrucatum

400

endemic

1,200

endemic

P. watutense

Note: Includes species with presence outside New Guinea (after Yoshimura et al. 1995; Louwhoff and Elix 1999). Note that several species have wide geographical and altitudinal ranges, but are scarcely recorded.

................. 16157$

CH13

02-08-07 10:29:01

PS

PAGE 313

314 / harrie sipman & andre´ aptroot

Comparison among Three Areas in Different Altitudinal Zones During four recent expeditions with P. Diederich, R. Gradstein, P. Lambley, and E. Se´rusiaux, we collected large amounts of material (approximately 10,000 specimens) in three areas situated close together, forming a kind of altitudinal transect. Each area was visited repeatedly by at least three of these lichenologists, and attempts were made towards a thorough, exhaustive sampling of all available habitats. The total amount of time spent at each of the three altitudinal areas ranges from about 6 to 14 person-days. In all localities, several tree crowns were investigated, often immediately after they fell. Consequently the lichen flora of these areas can be considered to have been fairly well sampled, and sampled to a similar degree in all three areas. Thus comparisons of the lichen floras of these three areas can be expected to yield meaningful results. For the comparison we included all identified and unidentified but recognizable species; about 60–70% of the species could be identified with certainty. A survey of the main characteristics is presented in Table 3.2.2. The species numbers of Table 3.2.2 show that the three areas contain very similar numbers of lichen species, despite having very different climates. However, clear differences are found in the frequency of life forms, vegetative reproduction, and main photobiont groups. The first altitudinal area, lowland, was located in the primary lowland forests in Madang Province. The results from several collecting localities, partly logging areas and partly scarcely disturbed primary lowland forest, are combined. The spots investigated are situated between 1 and 230 m altitude and occupy an area of about ten hectares altogether. About 500 species were recorded from this area. Crustose species dominate, while foliicolous, pantropical, and paleotropical species, and species with Trentepohlia, are abundant. According to current knowledge, ten species are endemic to New Guinea and perhaps are restricted to this region. They include conspicuous lichens such as Musaespora coccinea and Sporopodium lucidum, which are easily spotted and identified in the field by their bright red or yellow color, but have nevertheless never been found outside small areas near Madang. Many species seem to have a restricted distribution within the investigated area and are known from only one locality. There is almost no overlap in species of this area with the other two investigated areas. The second altitudinal area, Mt Gahavisuka, is a provincial park in one of the last easily accessible and intact mountain forests with Castanopsis (Fagaceae) in the PNG Highlands central valley system. The investigated area, which is part of a large area of primary, selectively logged forest, lies between 2,300 and 2,750 m and occupies about ten hectares. About 400 species were found, of which nearly 50 species may be endemic to New Guinea. Most other species are pantropical or paleotropical. In this and the next area foliose and fruticose lichens are more abundant than in the lowlands, and chlorococcoid green algae are the dominant photobionts (the photosynthetic partners of the lichens). The third altitudinal area was Mt Wilhelm, the highest mountain in PNG, bordering the central valley in the south. The investigated area reaches from the top

................. 16157$

CH13

02-08-07 10:29:01

PS

PAGE 314

Lichen Biodiversity in New Guinea / 315

Table 3.2.2. Altitudinal zonation of lichen properties in Papua New Guinea Lowland Madang

Mt Gahavisuka

Mt Wilhelm

1–230 m

2,300–2,750 m

3,500–4,500 m

500 60%

400 70%

420 70%

DISTRIBUTION Endemic Australasian Paleotropical Pantropical Temperate

5% 5% 40% 50% 0

10% 20% 35% 25% 10%

20% 20% 20% 20% 20%

SUBSTRATE Bark Leaves Soil Rock

70% 30% 0 0

75% 20% 5% 0

70% 5% 10% 15%

GROWTH FORM Fruticose Foliose Crustose

1% 15% 85%

10% 25% 65%

15% 35% 60%

PHOTOSYMBIONT Cyanobacteria Trentepohlia Chlorococcales

1% 50% 50%

10% 15% 75%

15% 10% 75%

VEGETATIVE REPRODUCTION Soredia Isidia None

15% 3% 82%

12% 6% 82%

20% 8% 72%

Altitude range Number of species Percentage of species identified

Note: Percentages are based on number of total species, except for the Distribution column, where percentages are based on number of identified species. Source: Modified from Aptroot (1997).

at 4,500 m down to 3,500 m in a valley and occupies about ten hectares. Between 4,000 and 4,500 m, the vegetation consists mostly of alpine tussock grassland. Below the natural tree line at 4,000 m the slopes are largely covered by low, sclerophyllous forest with the conifers Podocarpus and Papuacedrus. In some sites Cyathea tree ferns are dominant. In total about 450 species were found (Table 3.2.2), including abundant lichenicolous fungi. At least 80 species are endemic to New Guinea, according to current knowledge. Some of these species are very distinctive or locally common, and are thus unlikely to be overlooked, including an undescribed saxicolous Sticta, which occupies the niche of the (absent) Umbilicariaceae, and genera like Calathaspis and Compsocladium. Because alpine situations are rare

................. 16157$

CH13

02-08-07 10:29:02

PS

PAGE 315

#

316 /

´

near the equator, these species are likely to be restricted to small areas. Species with holarctic affinities are fairly abundant, although they are mostly restricted to areas above 4,000 m. For some of these species, this is the only known locality in the Southern Hemisphere. About 100 species are shared with Mt Gahavisuka. These represent the montane forest flora, including many species of Pseudocyphellaria. Virtually no species are shared with the lowland area. The highest zone is characterized by the presence of numerous saxicolous and terricolous species, which are, however, not very abundant. Sorediate species, fruticose lichens and such with cyanobacteria, are more common here than at lower altitudes.

Conservation While lichen species tend to have wide ranges, and are consequently of low protection priority, New Guinea has an unusually high number of endemic species that are restricted to the island and therefore deserving of more attention (cf. Table 3.2.1). This endemism seems unequalled by other tropical areas of the world, and exceeded only by some isolated southern temperate areas like New Zealand, Australia, and southernmost South America. In addition, lichens form an integral part of the cultural diversity for which New Guinea is famous (Figure 3.2.4). At the moment, over 1,200 species are reliably reported from New Guinea (see Aptroot et al. 2002). Even though probably only half of the flora on Papua New Guinea is known at present, lichenological knowledge now greatly exceeds that of all neighboring countries, except Australia. Notably the lichens of Indonesia, the Philippines, and Malaysia are much less known. Therefore the world distribution

Figure 3.2.4. The beard lichen Usnea is used for decoration. Little is known of other traditional use of lichens in New Guinea.

................. 16157$

CH13

02-08-07 10:29:05

PS

PAGE 316

Lichen Biodiversity in New Guinea / 317

of most Malesian species is unclear. Absence of records from surrounding countries may indicate that lichens are New Guinea endemics, or may be due to incomplete knowledge. However, it can be assumed that most of the species currently supposed to be endemics will be confirmed as such. Little can be said at this point about areas of endemism or hotspots. The available knowledge of the lichen flora comes largely from a few small areas: the surroundings of Port Moresby, Lae, and Madang, and the Highlands provinces of Papua New Guinea. Most species are known only from these areas, and there is no proof yet that they occur elsewhere on the island. It is unfortunate that almost no published information exists, for example, from the mountains of Papua, which are likely to have a rich lichen flora similar to that of the Central Highlands of Papua New Guinea. Nevertheless, it is improbable that the areas investigated happened by chance to be hotspots, with more lichen species than elsewhere on the island. Therefore it seems reasonable, for the time being, to call the whole island a ‘‘lichen hotspot,’’ in view of its many endemic species.

Recommended Action The lichen diversity of New Guinea is almost completely dependent upon primary forest. Some species appear to be able to maintain themselves on planted trees and in secondary vegetation, but it is unclear how many. Therefore the conservation of the lichen flora in New Guinea seems best served by the conservation of primary forests. Care should be taken that not only the tree diversity is maintained, but also that the forest structure is preserved, including the presence of large hardwood trees. Selective logging of such valuable trees greatly reduces the lichenological value of the forest, as can be observed regrettably in nature reserves worldwide. The most urgent research requirements seem to be inventories in other parts of New Guinea, particularly Papua, to find out if the endemic species found so far are widespread in the island or more localized. This would contribute to a more precise delimitation of hotspots of lichen diversity. Revisions of additional taxonomic groups will help to recognize additional endemic species deserving special attention. At the same time, lichen inventories should be made in the Malesian region outside New Guinea. The knowledge of the lichen flora of New Guinea currently greatly exceeds that of other Malesian areas, and it is unclear how many species seemingly restricted to New Guinea occur elsewhere in the region.

Literature Cited Aptroot, A. 1997. Lichen biodiversity in Papua New Guinea, with the report of 173 species on one tree. Biblioth. Lichenol. 68: 203–213. Aptroot, A., P. Diederich, E. Se´rusiaux, and H.J.M. Sipman. 1995. Lichens and lichenicolous fungi of Laing Island (Papua New Guinea). Biblioth. Lichenol. 57: 19–48. Aptroot, A., P. Diederich, E. Se´rusiaux, and H.J.M. Sipman. 1997. Lichens and lichenicolous fungi from New Guinea. Biblioth. Lichenol. 64: 1–256.

................. 16157$

CH13

02-08-07 10:29:06

PS

PAGE 317

#

318 /

´

Aptroot, A., P. Diederich, E. Se´rusiaux, and H.J.M. Sipman. 2002. Checklist of the lichens and lichenicolous fungi from New Guinea. Available at http: // www.biologie.uni-ham burg.de/checklists/papuanewguinea_l.htm. Aptroot, A., and H. Sipman. 1991. New lichens and lichen records from New Guinea. Willdenowia 20: 221–256. Aptroot, A., and H. Sipman. 1993. Musaespora, a genus of pyrenocarpous lichens with campylidia, and other additions to the foliicolous lichen flora of New Guinea. Lichenologist 25: 121–135. Archer, A.W., and J.A. Elix. 1998. Additional new species and two new reports in the lichen genus Pertusaria (Lichenised Ascomycotina) from Papua New Guinea. Mycotaxon 67: 155–179. Coppins, B.J. 2002. Checklist of Lichens of Great Britain and Ireland. British Lichen Society, London. Gradstein, S.R. 1992. The vanishing tropical rain forest as an environment for bryophytes and lichens. Pp. 234–258 in Bates, J.W., and A.M. Farmer (eds.) Bryophytes and Lichens in a Changing Environment. Clarendon Press, Oxford. ¨ sterreichs—eine Checkliste der Hafellner, J., and R. Tu¨rk. 2001. Die lichenisierten Pilze O bisher nachgewiesenen Arten mit Verbreitungsangaben. Stapfia (Linz) 76: 1–167. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. Pp. 69–84 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys, Leiden. James, P.W., A. Aptroot, P. Diederich, H.J.M. Sipman, and E. Se´rusiaux. 2001. New species in the lichen genus Menegazzia in New Guinea. Biblioth. Lichenol. 78: 91–108. Lambley, P.W. 1991. Lichens of Papua New Guinea. Pp. 69–84 in Galloway, D.J. (ed.) Tropical Lichens: Their Systematics, Conservation, and Ecology. Systematics Association Special Volume 43. Oxford Science Publications, Oxford. Louwhoff, S.H.J.J., and J.A. Elix. 1999. Parmotrema and allied lichen genera in Papua New Guinea. Biblioth. Lichenol. 73: 1–152. Louwhoff, S.H.J.J., and J.A. Elix. 2002. Hypotrachyna (Parmeliaceae) and allied genera in Papua New Guinea. Biblioth. Lichenol. 81: 1–149. Lu¨cking, R., and A. Veˇzda. 1998. Taxonomic studies in foliicolous species of the genus Porina (lichenized Ascomycotina: Trichotheliaceae)—II. The Porina epiphylla group. Willdenowia 28: 181–225. Mattick, F. 1942. Beitra¨ge zur Flora von Papuasien XXVI. 147. Die Flechten von NeuGuinea. 1. Allgemeines. Die Gattung Cladonia. Bot. Jahrb. 72: 151–158. Montfoort, D., and R.C. Ek. 1990. Vertical distribution and ecology of epiphytic bryophytes and lichens in a lowland rain forest in French Guiana. Unpubl. report, Utrecht. Santesson, R. 1952. Foliicolous lichens I. A revision of the taxonomy of the obligately foliicolous, lichenized fungi. Symb. Bot. Upsal. 12 (1): 1–590, pl. 1, fig. 1–92. Santesson, R., R. Moberg, A. Nordin, T. Tønsberg, and O. Vitikainen. 2004. Lichenforming and Lichenicolous Fungi of Fennoscandia. Museum of Evolution, Uppsala. Se´rusiaux, E. 1997. Sporopodiopsis, a new genus of lichens (Ectolechiaceae) from S-E Asia. Abstr. Bot. 21: 145–152. Sipman, H.J.M. 1983. A monograph of the lichen family Megalosporaceae. Biblioth. Lichenol. 18: 1–241. Sipman, H.J.M. 1993. Lichens from Mount Kinabalu. Trop. Bryol. 8: 281–314. Sipman, H.J.M. 1998. Notes on the lichen genus Stereocaulon in New Guinea. Cryptog., Bryol.-Liche´nol. 19: 229–245.

................. 16157$

CH13

02-08-07 10:29:07

PS

PAGE 318

Lichen Biodiversity in New Guinea / 319 Sipman, H. 2004. The species of Lobaria (lichenized Ascomycetes) in New Guinea. Pp. 573–606 in Do¨bbeler, P., and G. Rambold (eds.) Contributions to Lichenology, Festschrift in Honour of Hannes Hertel. Biblioth. Lichenol. 88: 1–739. Stenroos, S. 1986–1988. The family Cladoniaceae in Melanesia. 1. Cladonia sect. Unciales. Ann. Bot. Fennici 23: 161–164 (1986); 2. Cladonia section Cocciferae. Ann. Bot. Fennici 23: 239–250 (1986); 3. Cladonia sections Helopodium, Perviae and Cladonia. Ann. Bot. Fennici 25: 117–148 (1988); 4. The genera Cladia, Cladina, Calathaspis and Thysanothecium. Ann. Bot. Fennici 25: 207–217 (1988). Streimann, H. 1986. Catalogue of the Lichens of Papua New Guinea and Irian Jaya. Biblioth. Lichenol. 22: 1–145. Streimann, H. 1990. New lichen records from New Guinea. J. Hattori Bot. Lab. 68: 255–267. Streimann, H., and H.J.M. Sipman. 1994. New lichen records from the island of New Britain in Papua New Guinea. Fragm. Flor. Geobot. 39: 369–382. ¨ . 1956. Prodrome de la flore liche´nologique de la Nouvelle Guine´e. Ann. Hist. Szatala, O Mus. Nat. Hungar. n.s. 7: 15–50. Yoshimura, I., H.J.M. Sipman, and A. Aptroot. 1995. The lichen genus Anzia in New Guinea. Biblioth. Lichenol. 58: 439–469.

................. 16157$

CH13

02-08-07 10:29:07

PS

PAGE 319

3.3. Bryophytes of Papua New Guinea: Their Diversity, Ecology, and Conservation .

,

, .

, a group of non-flowering plants including mosses, liverworts, and hornworts, are evolutionary descendants of early terrestrial plants. They are distinguished by the common feature of possessing a sporophytic (diploidal, or bearing the full number of chromosomes) body that is attached to the gametophytic (haploidal, or bearing one half the normal number of chromosomes) body. The stem and leafy plant parts of bryophytes represent the sexually generated portions of the plant. As a group of terrestrial plants, bryophytes prefer humid and cool habitats, such as mountain rainforest and shaded stream banks. Po´cs (1982) and Gradstein and Po´cs (1989) presented excellent reviews on the diversity, biology, life history, ecology, and distribution of bryophytes in the tropics, especially in tropical rainforests. Because of their rather inconspicuous size, bryophytes have little economic importance to humans, but their role in the forest ecosystem is critical. Tan (2003) discussed the limited uses of mosses by people in tropical Southeast Asia, including New Guinea, where a large forest moss, Spiridens reinwardtii Nees (Figure 3.3.1), was used for body decoration by the local tribesmen.

B

History of Bryological Exploration in Papua New Guinea New Guinea has a rather unique tectonic history. The island is a landmass resulting from the juxtaposition of a southern plate with a Gondwanan flora and a northern plate that has a Laurasian flora (Piippo 1992; Hall 1998). Its complex topography has supported many types of vegetation since the Miocene (Paijmans 1976; Gressitt and Nadkarni 1978; Koponen and Norris 1983), making the island well known today for its great biotic diversity and endemism (Myers et al. 2000; Tan and Po´cs 2000). The history of taxonomic study of bryophytes of New Guinea in general, and Papua New Guinea in particular, was outlined by Schultze-Motel (1963) and Koponen and Norris (1983). The former published the first comprehensive checklist of mosses for the island (Schultze-Motel 1963), while Grolle and Piippo (1984) published the first annotated catalogue of the Hepaticae and Anthocerotae of New Guinea and Western Melanesia. A perusal of the references listed in these three publications shows that many bryologists in Europe and North America of the 19th and early 20th centuries, including Dozy and Molkenboer, Hookers, Geheeb, C. Mueller, Hampe, Stephani, Sande Lacoste, Schiffner, Mitten, Fleischer, BrothMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

320

................. 16157$

CH14

02-08-07 10:28:46

PS

PAGE 320

Bryophytes of Papua New Guinea / 321

Figure 3.3.1. Spiridens reinwardtii Nees. erus, Bescherelle, Herzog, Dixon, Verdoorn, Reimers, and Bartram, have had a hand in the study of New Guinean bryophytes.

Diversity and Richness of New Guinean Bryophytes Based on the above-mentioned checklists of mosses and hepatics and on recent monographs, the number of New Guinean bryophytes is reported to consist of 767 species, 129 genera, and 42 families of liverworts and hornworts, and some 890 species, 248 genera, and 61 families of mosses (Grolle and Piippo 1984; Tan

................. 16157$

CH14

02-08-07 10:28:49

PS

PAGE 321

322 /

.

,

,

.

and Iwatsuki 1999; see literature cited in Koponen 2000; Gradstein et al. 2002). More than 70% of the bryophytes known from Malesia and Australasia are represented in the flora of New Guinea. The wide variation in morphological and anatomical complexities seen among local bryophytes is equally impressive, ranging among mosses from the giant Dawsonia (Figure 3.3.2) with complex stem anatomy, to tiny (1–2 mm), ephemeral Trachycarpidium with spiculose capsules, and the nearly microscopic and filamentous Ephemeropsis, and, among liverworts and hornworts from large Megaceros and Conocephalum, to minute, epiphyllous Cololejeunea, Aphanolejeunea, Papillolejeunea, and Metzgeriopsis. The island is also the primary or secondary center of speciation in Malesia for several large genera, such as Frullania (ca 90 spp.), Plagiochila (54 spp.), Bazzania (ca 50 spp.), Radula (42 spp.), Macromitrium (29 spp.), Chaetomitrium (25 spp.), and Acroporium (17 spp.). For these and other genera, New Guinea harbors a disproportionately large number, if not the highest number, of endemic species by island in the region. Examples are Dawsonia (5 endemics out of 7 species), Dicranoloma (5 endemics out of 14 species), Radulina (3 endemics out of 4 species), Clastobryum (2 endemics out of 4 species), Frullania (57 endemics out of 90 species), Radula (15 endemics out of 42 species), Plagiochila (30 endemics out of 54 species), and Schistochila (14 endemics out of 19 species). The past two decades have seen a large number of modern revisions of families of New Guinean bryophytes initiated by Koponen and Norris (1983), who in 1981 made a large collection of more than 17,000 specimens of bryophytes from the Huon Peninsula and the Frieda River mining area in Papua New Guinea (Koponen 1990). The latest account of the taxonomic progress of this project documented 1,009 species (531 mosses and 478 hepatics) and 284 genera of bryophytes (Koponen 2000). Seventy-five species and five genera were found to be new to science. In addition, 223 new species records and 44 new generic records have been added to the bryofloras of New Guinea and the Solomon Islands (Koponen 2000) and more than 440 names have been reduced to new synonyms. For a breakdown of the number of species and genera of the New Guinean bryoflora studied by the Huon Peninsula project, see Koponen (1990, 1998, 2000). By and large, the regional publications on the Malesian Hepaticae have been fewer than those on Musci. As a result, reports of new taxa of hepatics predominate in recent publications of New Guinean bryoflora, while reductions of early names to synonymy predominate in recent works on the island’s mosses (Koponen 1990). To date, there are three endemic moss genera known in New Guinea (Leskeodontopsis Zant., Orthothuidium D.H. Norris & T. Kop., and Unclejackia M. Ignatov, T. Koponen & D.H. Norris), but none for the Hepaticae. The percentage of species endemism is calculated to be ca 18% (Hyvo¨nen 1989; Tan and Iwatsuki 1999) for mosses, and a much higher 38.2% for liverworts and hornworts (Piippo et al. 1987; Piippo 1994; Koponen 1990). Most of the endemic species occur at elevations above 1,500 m (Koponen 1990), are most frequent between 2,300 and 2,600 m (Piippo 1994), and are epiphytic on tree trunks and branches. According to Piippo (1994), most of the New Guinean endemics are of Gondwanan origin.

................. 16157$

CH14

02-08-07 10:28:50

PS

PAGE 322

Bryophytes of Papua New Guinea / 323

Figure 3.3.2. Dawsonia, a large moss commonly found in forests in New Guinea.

................. 16157$

CH14

02-08-07 10:29:15

PS

PAGE 323

.

324 /

,

,

.

In recent years, the project on the Huon Peninsula aiming to document the bryodiversity of Papua New Guinea (Koponen and Norris 1983) has included in its study other collections, such as those of H. Streimann, L. J. Brass, W. A. Weber and D. McVean, and R. D. Hoogland, in order to present a complete bryoflora of New Guinea and western Melanesia. These later collections were kindly made available for the revision by various herbaria (L, CANB, FH, BM). Consequently, the results also allow a more realistic prediction of about a total of 1,500 species of bryophytes in the final analysis. Over 100 of these species will be new to science, and about 300 taxa will be confirmed to be endemic to New Guinea and the Solomon Islands. Tables 3.3.1 and 3.3.2 compare the bryophyte diversity of New Guinea with that of some neighboring islands and countries of comparable size. It is apparent that the bryophyte diversity of New Guinea has the highest number of taxa at all taxonomic ranks. The differences in the number of taxa between the bryofloras of PNG and Papua (Tables 3.3.1 and 3.3.2) reflect the undercollection of these groups of plants in Papua. When intensive collections are made in Papua in the future, the bryophyte floras of the two halves of this large island will most probably be found to be similar. As it stands today, the many new and endemic species discovered provide important information relevant to conservation efforts in New Guinea. The Huon Peninsula collections are so numerous that the rarely collected species can be interpreted, with some confidence, as locally rare. It should be noted, however, that the collections from the Huon Peninsular project were made mostly from rather high elevations, and consequently, many of the lowland species have been poorly represented. Unfortunately, the frequency of collection also partially reflects the predilections of the collectors. As an example, Koponen and Norris (1983) often collected together around each of the campsites, but there are many

Table 3.3.1. Moss diversity in New Guinea and similar-size neighboring islands and countries. Location

Number of species

Number of genera

Number of families

New Guinea Papua New Guinea Papua

890 624 426

248 210 156

61 58 51

Borneoa

724

192

50

702

230

59

628

189

50

Philippines

b

Java Peninsular Malaysia

c

Indochinad

476

147

42

1,001

238

55

Source: for all locations: Tan and Iwatsuki 1999. Additional sources: a. Suleiman and Edwards 2002; Suleiman 2004; b. Tan and Mandia 2001; c. Ho and Tan 2002; Mohamed et al. 2004; d. Tan and Tran Ninh 2003; Tan et al. 2003.

................. 16157$

CH14

02-08-07 10:29:15

PS

PAGE 324

Bryophytes of Papua New Guinea / 325

Table 3.3.2. Hepatic and hornwort diversity in New Guinea and similar-size neighboring islands and countries Location

Number of species

Number of genera

Number of families

New Guinea Papua New Guineaa Papuaa

767 694 343

129 126 86

42 41 35

Borneob

623

119

35

Philippines

518

100

34

Java

a

c

315

75

31

Peninsular Malaysia

NA

NA

NA

Indochina

NA

NA

NA

d

Note: NA indicates no information available. Source: a. Grolle and Piippo 1984; Huon series (for references, see Koponen 2000; Gradstein et al. 2002); b. Menzel 1988; c. Tan and Engel 1986; d. Schiffner 1898.

species frequently collected by one but rarely collected by the other. Finally, some bryophytes are so small as to be picked up only as accidental impurities in a packet representing another species. In retrospect, a further two decades will be required to complete the inventory of the bryodiversity of New Guinea. At present, most of our information about the New Guinean bryophytes come from relatively accessible localities in PNG, such as Mt Sarawaket, Mt Kaindi, the Rawlinson Range, Cromwell Mts, Bulolo, Sattelberg, and Lae, mainly in Morobe and Madang provinces (Brotherus 1892, 1901; Bartram 1945; Noguchi 1953; Schiffner and Stephani 1901; Verdoorn 1934; Koponen 2000). New information on bryophytes of Mt Wilhelm and Mt Gahavisuka has been reported recently (Tan 2000). In Papua, the most comprehensive study of mosses is the one published for the Star Mts (Mt Antares) (van Zanten 1964). At the provincial level, Morobe Province, with more than 600 species, has the highest reported number of liverworts and mosses. This is followed by Central, Simbu, Western Highlands and Eastern Highlands in PNG, and by the regions around Jayawijaya and Manokwari in Papua (Grolle and Piippo 1984). Some areas, like West Sepik and Gulf and Western of PNG, and Fakfak and Cenderawasih of Papua, have less than 35 known species of bryophytes. These areas should be the focus of future collections.

Phytogeographic Relationships of New Guinean Bryoflora Effective conservation requires a basic knowledge of phytogeographic patterns. This is because a species that is rare on one mountain, or found once in a forest reserve, may be commonly represented elsewhere, often under another name. Assertion of species rarity depends much upon our knowledge of the flora from phytogeographically related areas. To understand the phytogeography of the New

................. 16157$

CH14

02-08-07 10:29:16

PS

PAGE 325

326 /

.

,

,

.

Guinean bryoflora, we have amalgamated a long list of bryophytes from various publications (see Shultze-Motel 1963; Grolle and Piippo 1984; Ramsay et al. 1986). Interestingly, many of the previous publications suggest a close affinity of the New Guinean flora with that of Australia. Enroth (1991) conducted a literature review of the phytogeographical affinity of the New Guinea and western Melanesian hepatic flora on the basis of 37 out of 40 families and 399 species, and concluded that the New Guinean Hepaticae has strong affinities with the floras of Southeast Asia and Oceania, and weak affinity with the Australian flora. Piippo (1992) compared the affinities of the New Guinea hepatic flora to neighboring areas, analyzing 2,906 species, 250 genera, and 62 families. The results of her study supported the analysis of Enroth (1991). Earlier Hyvo¨nen (1989; see also Piippo and Koponen 1997), who did a similarly extensive phytogeographical analysis based on 27 of the 57 families and on some 298 species and subspecific taxa of mosses, also came to the same conclusion. His findings demonstrated that the New Guinean moss flora is more closely related to the flora of Asia, for example, the Indonesian archipelago (68%) and the Philippines (52%), than to those of Australia (42%) and New Zealand (12%). Piippo and Koponen (2003) conducted an additional review of the phytogeographical connections of New Guinea’s bryoflora and reached the same conclusion. Our unpublished data on the species composition of the largest moss family, Sematophyllaceae, support this conclusion as well. In addition, several moss species of this family, such as Acroporium microcladon var. burleyii B.C. Tan, Church & Windadri, Pseudopiloecium scabrisetum Bartr., and Clastobryopsis perdecurrens (Dix.) B.C. Tan, are newly confirmed to have a disjunctive distribution between Borneo and New Guinea. Other shared mosses between Borneo and New Guinea (e.g., Brotherella longipes Broth.) have an extended range, reaching Mindanao Island (but not further north to Luzon Island) of the Philippine archipelago. Examples of New Guinean hepatics and hornworts with similar distribution patterns are Dendroceros cavernosus J. Haseg., Aphanolejeunea borneensis (Herzog) Po´cs, and Radula iwatsukii Yamada. To date, the Huon Peninsula project has primarily focused on a comparison of the bryophyte floras of Indonesia, Borneo, and the Philippines, resulting in the synonymization of many names. Future workers will similarly find other reductions in a number of putative New Guinea endemics by comparing them with endemic species known from other parts of Malesia. Unfortunately our current knowledge the bryoflora of the vast island chain of Indonesia is incomplete and precludes definitive statements regarding the distribution pattern of many of the New Guinean species.

Ecology and Ecological Adaptation of Bryophytes Bryophytes in tropical rainforests are appropriately treated as organisms that are ecologically separated from the vascular plants. They occupy different ecological niches as a result of differences in morphology and physiology. In the everhumid forested areas of the earth, bryophytes are a major contributor to ecosystem func-

................. 16157$

CH14

02-08-07 10:29:16

PS

PAGE 326

Bryophytes of Papua New Guinea / 327

tioning. In the tropics they are among the most essential plant groups in forest ecosystems (Figure 3.3.3). Several morphological and physiological features of bryophytes contribute to their role as essential parts of the forest ecosystem. Bryophytes completely lack a root system and have evolved the ability to absorb water and essential nutrients directly into all living parts of the plant. The large bryophyte mass in rainforests allows them to absorb large amounts of precipitation (Po´cs 1976) and essential nutrients (Nadkarni 1984, 1985) that leach down from the canopy. Bryophytes can effectively intercept both the water and mineral nutrients (Po´cs 1980). In an area of high bryophyte mass, up to half the mineral nutrient capital of the forest can be stored in the bodies of the bryophytes (Nadkarni 1984). The water and minerals so absorbed are then released slowly into the soil in a manner that allows root uptake by the vascular plants. In a tropical rainforest, the canopy is very uneven in height, with some species of tree dominating in the upper canopy while neighboring trees reach a lower canopy level. We have noted that on New Guinea the taller canopy trees often have greater trunk bryophyte mass than the shorter canopy trees. Because of high precipitation in an archipelagic environment, many of the rainforests on Huon Peninsula have a much larger mass of epiphytic bryophytes than any temperate forest.

Figure 3.3.3. Mossy understory of a Nothofagus-dominant forest on limestone, 05 56 55 S, 142 44 40 E, elev. 7,550 ft aneroid (2,302 m)

................. 16157$

CH14

02-08-07 10:29:35

PS

PAGE 327

328 /

.

,

,

.

Bryophytes essentially lack a well differentiated vascular system. Instead, most water and dissolved nutrients are transported along the surfaces of stems and leaves in capillary spaces created by the close proximity of individual plants. Their close proximity is made possible by the lack of a root system, which eliminates root competition as a spacing enforcer within a population. This lack of a root system in bryophytes is also associated with a capacity for dormancy during frequent periods of drying. Drought dormancy in rainforest bryophytes may be observed daily because their plant body has a large surface-to-volume ratio with the epidermal cuticle poorly developed or absent. This allows a more or less free flow of water into the plant body and leads also to high rate of evapotranspiration, thereby risking desiccation. It should be noted that the capacity for drought dormancy is not only an effective way of avoiding damage from drying, but is also an effective way of avoiding damage from higher temperatures. While physiological studies of tropical bryophytes are unfortunately few, the works so far published (Frahm 1987; literature cited in Schofield 1985) show bryophytes, even those in tropical latitudes, to be cool-loving plants unable to have a net increase in photosynthate above 25 C. Thus the midday period of high temperature is usually a period of bryophyte dormancy due to drought. In general, high temperatures associated with drought pose no problem to dormant bryophytes, but being warm and wet is often fatal. Enroth (1990; see also Piippo 1994) analyzed the species assemblage found at five different altitudinal ranges in Huon Peninsula, namely Zone 1: 0–300 m, Zone 2: 300–1,200 m, Zone 3: 1,200–2,200(–2,300) m, Zone 4: 2,200(–2,300)–2,800 (–2,900) m, and Zone 5: above 2,800(–2,900) m. The results clearly show a vertical zonation of bryophytes that parallels the scheme of altitudinal zonations constructed based on the phanerogamic (higher plants) flora and vegetation types. The New Guinean endemics and Melanesian and Malesian taxa are found to be distributed at middle to relatively high altitudes (Zones 3 and 4), whereas the widespread Oceanian, Australasian, and Asiatic taxa are represented heavily at low to middle altitudinal zones (Zones 1 to 3). Widespread but locally rare species of high latitude genera, such as mosses Andreaea, Leptodictyum, Hygrohypnum, Warnstorfia, and Limprichtia, and liverworts Andrewsianthus, Anthelia, Gymnomitrion, Marsupella, and Sphenolobopsis, occur at high altitudes (Zones 4 and 5). The vertical zonation of distribution of bryophytes seen in Papua New Guinea is most likely similar to that in Papua. Not surprisingly, Koponen and Norris (1983) and Norris (1990) have found the epiphytic bryophytes to be useful indicators of the nature of substrates and the degree of forest disturbance in Papua New Guinea. The huge number of collections that came out of the Huon Peninsula project was accompanied by the most skeletal of remarks on substrate, moisture, and sunlight exposure. The expedition was conducted with the intention of collecting as many different species as possible from as many habitats and geographic sites as possible, for ecological comparison. Although the rare bryophytes are primarily collected from trees, branches, and leaves, these collections can hardly make a definitive statement about individual ecological preference of the species, because

................. 16157$

CH14

02-08-07 10:29:35

PS

PAGE 328

Bryophytes of Papua New Guinea / 329

of the diverse and wide range of habitats occupied by many bryophyte species across the island. Rock outcrops are not frequently encountered inside rainforests in PNG, except at high elevations or along areas of frequent riverine erosion. They are found to be the habitat with the lowest diversity of bryophytes. One reason may be that most outcrops weather too rapidly in tropical sites to allow long-term persistence of any particular species. In contrast, rock outcrops in north temperate regions regularly have rare bryophytes whose low vagility (ability to move between locations) is inconsequential to persistence on the outcrops. With a large portion of the bryophyte diversity concentrated on the bark of trees, it becomes necessary to understand how bryophytes disperse in nature (van Zanten and Po´cs 1981) and the manner in which the epiphytic substratum is partitioned to allow its great bryological diversity. There is an obvious division of tree architecture into outer canopy, inner canopy, upper trunk, tree base, and leaves, and this division represents large differences in nearly all of the important environmental parameters affecting bryophytes (Gradstein 1997; Gradstein et al. 2003). The great mass of epiphytic bryophytes in tropical rainforests also should be examined in terms of the types of animals and protists that it supports. The surface of epiphytic mats of bryophytes is more extensive than the exposed soil surface, and even so, the soil surface is often very heavily cloaked in parts by bryophytes. It is unknown how many species of animals are critically dependent upon this bryophyte mass in the forest.

Conservation: Current Status, Problems, and Challenges Conservationists around the world are concerned primarily with the preservation of species and ecosystems. There are two approaches to resolving the issue, one focusing on species diversity and the other on the ecosystem. These two approaches rely upon different types of biological knowledge. On New Guinea, as well as on other tropical islands, bryophytes grow to great mass in closed tropical rainforests by utilizing their thick masses to maintain a cool temperature and regulate air humidity. Hyvo¨nen et al. (1987) reported that limited practice of shifting agriculture in thinly populated areas in Papua New Guinea did not present a serious threat to the existing bryoflora. Nonetheless, large-scale loss of bryophyte cover, with serious ecological consequences, can be triggered by opening the forest and by increasing air pollution from agriculture and mining activities. This loss of bryophyte cover can result in a decrease in relative humidity and a reduction in soil fertility that is detrimental to tropical rainforests. Species with narrow ecological tolerances and geographical ranges are usually the first to become locally extirpated (Hyvo¨nen et al. 1987). One of us has attempted to evaluate the effects deforestation on forest bryophytes in PNG (Norris 1990), but those evaluations can hardly be used as the basis of a speciescentered conservation plan.

................. 16157$

CH14

02-08-07 10:29:35

PS

PAGE 329

330 /

.

,

,

.

The island of New Guinea has been identified recently as one of the earth’s three Major Tropical Wilderness Areas for biodiversity conservation (Myers et al. 2000) and an area of high bryophyte endemism (Tan and Po´cs 2000), and Mt Wilhelm Forest Reserve has been chosen to be one of the four major threatened hotspots of high moss diversity in Malesia (Tan and Iwatsuki 1999). However, the economic development occurring in many parts of the island does not bode well for the future of New Guinean bryoflora. Aggravating the situation is the insufficient areas of nature or forest reserves allocated for protection, especially in Papua New Guinea. Another problem is how to protect effectively the rare and endangered bryophyte species on the island. Conservation decisions focused on individual rare and endangered species require a deep knowledge of the local geography of the biota in the region. Inadequate collection in many provinces of Papua New Guinea and Papua makes it difficult to design an appropriate conservation plan for the island. While we have witnessed in this past century the description and documentation of a large percentage of the island’s bryophyte flora, we must await the geographically exhaustive collecting necessary to substantiate the statement of rarity of individual species. At present, our knowledge allows statements about the world distribution of each species, a good listing of New Guinean endemic species, and an evaluation of relative abundance based on the known collection localities and the number of repeated collections of a species. Nonetheless, our studies do not yet allow a definitive selection of endangered species for species-centered conservation efforts in the island, nor in the region. In most of the northern temperate regions of the world, species-centered conservation is very effectively implemented because the distribution and taxonomic structure of the plants are well studied. Species-by-species conservation has the advantage of being legally unambiguous under rare and endangered species legislation. Unfortunately, much of the tropics do not have an adequate knowledge base for an effective conservation effort of this type. Because we are not yet sufficiently equipped for a species-centered conservation treatment of bryophytes in New Guinea, could we instead focus on the preservation of various ecosystems present locally? In the case of New Guinea, we know a lot about the distribution of various forest types, including their latitudinal and elevational limits. We also know the distribution of specific types of forest with heavy coverage of bryophytes. Conservation of a single ecosystem from the alpine/ subalpine zone to the valley and ocean is a possibility. Such a conservation approach should take advantage of the mineral- and water-absorptive capacities of bryophytes, and, in so doing, will provide a strong and convincing justification for the preservation of the entire ecosystem and its biota. With proper documentation, advocacy, and mass campaigning about the benefits that come with protection of the entire ecosystem, preservation of a combined mountain and valley ecosystem in the relatively less populated New Guinea may be acceptable to the people and local government. For conservation to be effective, future ecological studies of epiphytic bryo-

................. 16157$

CH14

02-08-07 10:29:36

PS

PAGE 330

Bryophytes of Papua New Guinea / 331

phytes in New Guinea should be carried out to address the following two questions: First, do tree species differ in terms of total bryophyte mass and the suite of bryophyte species they host? We have noted the importance of such plants (e.g., tree ferns and Pandanus) as favorable bryophyte substrates in the Huon Peninsula, but most of the field notes of bryophyte collections we examined list the substratum only as ‘‘tree,’’ without regard to species. Often there is no effort even to list the trees as smooth- or furrowed-barked. Perhaps in the rush to accomplish rapid collection in the field, the lack of effort to note the relative height of the trees from which the individual collection was obtained, is inevitable. The second question is, what kind of wealth of natural resources and bryophyte diversity is stored high in the canopy of local rainforests? In many of our New Guinean expeditions, we customarily inspected recently fallen trees to find bryophytes. Some of the species that we associate with the canopies were found also on smaller, accessible trees perched on the edge of steep cliffs. Despite this observation, we remain convinced that sufficient surveys of the canopies of New Guinean forests will result in significant additions to the flora, as well as significant changes in the rarity status of some species. As in any other part of the world, conservation in PNG, and perhaps in Papua too, is subject to constraints. Conservation efforts are variously influenced by the economics and politics of the country and the region, by the low level of biological knowledge of the area, and by the level of habitat destruction that has already occurred. This is especially true in PNG where the land alienated from tribal ownership is almost nonexistent. The economic impact of any land set aside for conservation will therefore fall disproportionately on tribal members rather than equally on people throughout the country. Unless community awareness is aroused and galvanized with help from the government agencies and nongovernmental environmental groups, any attempt to protect the biodiversity in New Guinea, which is still plentiful and significant at present, will be a difficult goal to achieve. While we have spoken of conservation of bryophytes in New Guinea as problematic, we also realize that this mental exercise accomplishes little. What can be done at our present state of knowledge? Could land at a variety of altitudes, mineralogic regimes, and moisture zones be managed so as not to seriously damage the biota? Is such an approach effective even when we know little of the interaction between bryophyte communities and their various ecologies? Pending a more satisfactory solution, we seriously suggest the land set-aside as a stop-gap measure, while research on the distribution and ecology of the biota proceed to improve the conservation program.

Acknowledgments We thank Conservation International and Dr. W. Takeuchi for the invitation to prepare this chapter on bryophyte diversity. We are grateful to Prof. T. Koponen who initiated and organized the investigation of the bryophytes of Huon Peninsula

................. 16157$

CH14

02-08-07 10:29:37

PS

PAGE 331

332 /

.

,

,

.

in PNG. We are also grateful to Mr. J. R. Shevock for reading drafts of this chapter and providing valuable comments. The first author acknowledges the financial support of DAAD, the Finnish-Chinese Botanical Foundation, and the NUS.

Literature Cited Bartram, E.B. 1945. Mosses of Morobe District, Northeast New Guinea. Bryologist 48: 110–126. Brotherus, V.F. 1892. Musci. In O. Warburg Bergpflanzen aus Kaiser Wilhelm-Land, gesammelt auf der Zo¨ller’schen Expedition in Finisterregebirge von F. Hellwig. Bot. Jahrb. 16: 1–32. Brotherus, V.F. 1901. Musci. Pp. 79–104 in Schumann, K., and K. Lauterbach (eds.) Die Flora der Deutschen Schutzgebiete in der Su¨dsee. Gebru¨der Borntra¨ger, Leipzig. Enroth, J. 1990. Altitudinal zonation of bryophytes on the Huon Peninsula, Papua New Guinea. A floristic approach, with phytogeographical considerations. Tropical Bryol. 2: 61–90. Enroth, J. 1991. On the phytogeography of Western Melanesian Hepaticae. A literature review. J. Hattori Bot. Lab. 70: 1–42. Frahm, J.-P. 1987. Which factors control the growth of epiphytic bryophytes in tropical rain forests? Symposia Biol. Hungarica 35: 639–648. Gradstein, S.R. 1997. The taxonomic diversity of epiphyllous bryophytes. Abstracta Bot. 21: 15–19. Gradstein, S.R., X.-L. He, S. Piippo, and M. Mizutani. 2002. Bryophyte flora of the Huon Peninsula, Papua New Guinea. LXVIII. Ptychanthoideae (Hepaticae). Acta Bot. Fenn. 174: 1–88. Gradstein, S.R., N.M. Nadkarni, T. Kro¨mer, I. Holz, and N. No¨ske. 2003. A protocol for rapid and representative sampling of vascular and non-vascular epiphyte diversity of tropical rain forests. Selbyana 24: 87–93. Gradstein, S.R., and T. Po´cs. 1989. Bryophytes. Pp. 311–325 in Lieth, H., and M.J.A. Werger (eds.) Tropical Rain Forest Ecosystems. Elsevier, Amsterdam. Gressitt, J.L., and N. Nadkarni 1978. Guide to Mt. Kaindi: Background to Montane New Guinea Ecology. Wau Ecology Institute Handbook No. 5, Hong Kong. Grolle, R., and S. Piippo. 1984. Annotated catalogue of Western Melanesian bryophytes. I. Hepaticae and Anthocerotae. Acta Bot. Fenn. 125: 1–86. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. Pp. 99–131 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys, Leiden. Ho, B.-C., and B.C. Tan. 2002. Additions to the moss flora of Endau Rompin National Park, Johore State, Peninsular Malaysia. Tropical Bryol. 22: 67–76. Hyvo¨nen, J. 1989. On the bryogeography of Western Melanesia. J. Hattori Bot. Lab. 66: 231–254. Hyvo¨nen, J., T. Koponen, and D.H. Norris. 1987. Human influence on the moss flora of tropical rain forest in Papua New Guinea. Symposia Biol. Hungarica 35: 621–629. Koponen, T. 1990. Bryophyte flora of Western Melanesia. Tropical Bryol. 2: 149–160. Koponen, T. 1998. Index of the Bryophyte Flora of the Huon Peninsula, Papua New Guinea. Division of Systematic Biology, University of Helsinki, Helsinki. Koponen, T. 2000. Index of the Bryophyte Flora of Western Melanesia. Division of Systematic Biology, University of Helsinki, Helsinki.

................. 16157$

CH14

02-08-07 10:29:37

PS

PAGE 332

Bryophytes of Papua New Guinea / 333 Koponen, T., and D.H. Norris. 1983. Bryophyte flora of the Huon Peninsula, Papua New Guinea. I. Study area and its bryological exploration. Ann. Bot. Fenn. 20: 15–29. Menzel, M. 1988. Annotated catalogue of the Hepaticae and Anthocerotae of Borneo. J. Hattori Bot. Lab. 65: 145–206. Mohamed, H., K.T. Yong, and G. Gunaseelan. 2004. Additions to the moss flora of Peninsular Malaysia. J. Bryol. 26: 47–52. Myers, N., R.A. Mittermeier, C.G. Mittermeier, G.A.B. Fonseca, and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853–858. Nadkarni, N. 1984. Epiphyte biomass and nutrient capital of a neotropical elfin forest. Biotropica 16: 249–256. Nadkarni, N. 1985. Biomass and nutrient capital of epiphytes in an Acer macrophyllum community of a temperate moist coniferous forest, Olympic Peninsula, Washington State. Can. J. Bot. 62: 2223–2228. Noguchi, A. 1953. Mosses of Mt. Sarawaket, New Guinea. J. Hattori Bot. Lab.10: 1–23. Norris, D.H. 1990. Bryophytes in perennially moist forests of Papua New Guinea: ecological orientation and prediction of disturbance effects. Bot. J. Linn. Soc. 104: 281–291. Paijmans, K. (ed.). 1976. New Guinea Vegetation. Australian National University Press, Canberra. Piippo, S. 1992. On the phytogeographical affinities of temperate and tropical Asiatic and Australasiatic Hepatics. J. Hattori Bot. Lab. 71: 1–35. Piippo, S. 1994. Phytogeography and habitat ecology of Western Melanesian endemic Hepaticae. J. Hattori Bot. Lab. 75: 275–293. Piippo, S., and T. Koponen. 1997. On the phytogeographic biodiversity of Western Melanesian mosses. J. Hattori Bot. Lab. 82: 191–201. Piippo, S., and T. Koponen. 2003. Review of the bryofloristic connections of New Guinea Island. Telopea 10: 467–476. Piippo, S., T. Koponen, and D.H. Norris. 1987. Endemism of the bryophyte flora in New Guinea. Symposia Biol. Hungarica 35: 361–372. Po´cs, T. 1976. The role of the epiphytic vegetation in the water balance and humus production of the rain forests of the Uluguru Mountains, East Africa. Boissiera 24b: 499–503. Po´cs, T. 1980. The epiphytic biomass and its effect on the water balance of two rain forest types in the Uluguru Mountains (Tanzania, East Africa). Acta Bot. Acad. Sci. Hungaricae 26:143–167. Po´cs, T. 1982. Tropical forest bryophytes. Pp. 59–104 in Smith, A.J.E. (ed.) Bryophyte Ecology. Chapman & Hall, London. Ramsay, H.P., H. Streimann, A.V. Ratkowsky, R. Seppelt, and A.J. Fife. 1986. Australasian alpine bryophytes. Pp. 300–335 in Barlow, B.A. (ed.) Flora and Fauna of Alpine Australasia. CSIRO and ASBS, Melbourne. Schiffner, V. 1898. Conspectus Hepaticarum Archipelagi Indici. Staatsbruckerei, Batavia. Schiffner, V., and F. Stephani. 1901. Hepaticae. Pp. 69–79 in Schumann, K., and K. Lauterbach (eds.) Die Flora der Deutschen Schutzgebiete in der Su¨dsee. Gebru¨der Borntra¨ger, Leipzig. Schofield, W.B. 1985. Introduction to Bryology. Macmillan, New York. Schultze-Motel, W. 1963. Vorla¨ufiges Verzeichnis der Laubmoose von Neuguinea. Willdenowia 3: 399–549. Suleiman, M. 2004. Moss flora of Borneo (abstract). Asian Plant Diversity and Systematics, IAPT International Symposium 2004, July 29–Aug. 2, Sakura, Japan.

................. 16157$

CH14

02-08-07 10:29:38

PS

PAGE 333

334 /

.

,

,

.

Suleiman, M., and S.R. Edwards. 2002. Mosses of Mt. Trus Madi, Sabah, Malaysia. Tropical Bryol. 21: 57–64. Tan, B.C. 2000. Additions to the moss floras of Mt. Wilhelm Nature Reserve and Mt. Gahavisuka Provincial Park, Papua New Guinea. J. Hattori Bot. Lab. 89: 173–196. Tan, B.C. 2003. Bryophytes (mosses). Pp. 193–200 in de Winter, W.P., and V.B. Amoroso (eds.) Cryptogams: Ferns and Fern Allies. Plant Resources of South-East Asia (PROSEA) 15 (2). Backhuys, Leiden. Tan, B.C., and J.J. Engel. 1986. An annotated checklist of Philippine Hepaticae. J. Hattori Bot. Lab. 60: 283–355. Tan, B.C., V.T.T. Huong, and B.-C. Ho. 2003. Trachycarpidium echinatum and Weissia platystegia, new to Vietnam and Continental SE Asia. Cryptogamie, Bryol. 24: 43–47. Tan, B.C., and Z. Iwatsuki. 1996. Hot spots of mosses in East Asia. Anales Inst. Biol. Univ. Nac. Auto´n. Me´xico, Ser. Bot. 67: 159–167. Tan, B.C., and Z. Iwatsuki. 1999. Four hot spots of moss diversity in Malesia. Bryobrothera 5: 247–252. Tan, B.C., and E.H. Mandia. 2001. New and noteworthy records of mosses from Mindoro, the Philippines, and their biogeographical implication. Gardens Bull. Singapore 53: 315–322. Tan, B.C., and T. Ninh. 2003. Vu Quang and other Vietnam mosses collected by Tran Ninh, B.C. Tan and T. Po´cs in 2002. Acta Acad. Paed. Agriensis, Sect. Biol. 25: 85–101. Tan, B.C., and T. Po´cs. 2000. Bryogeography and conservation of bryophytes. Pp. 403–448 in Shaw, A.J., and B. Goffinet (eds.) Bryophyte Biology. Cambridge University Press, Cambridge. van Zanten, B.O. 1964. Mosses of the Star Mountains Expedition. Nova Guinea, Bot. 16: 263–368. van Zanten, B.O., and T. Po´cs. 1981. Distribution and dispersal of bryophytes. Advances in Bryology 45: 479–562. Verdoorn, F. 1934. Re´sultats de l’expe´dition scientifique ne´erlandaise a` la NouvelleGuine´e. Lejeuneaceae Holostipae. Nova Guinea 18: 1–8.

................. 16157$

CH14

02-08-07 10:29:38

PS

PAGE 334

3.4. Ferns and Lycophytes of Papua . have long been grouped together because they share the characteristics of dispersal by spores and possession of internal vascular tissue, distinguishing them from bryophytes (mosses, liverworts, and hornworts) that lack internal vascular tissue, and from seed plants (gymnosperms and flowering plants) that lack spores (Moran 2004). Recent studies have shown that fern allies (lycophytes, horsetails, and whisk ferns) are not a separate group of plants allied to ferns. They are now either considered ferns (horsetails: Equisetum and whisk ferns: Psilotum and Tmesipteris) or are recognized as less closely related to ferns than are the seed plants, as are the lycophytes, such as Isoetes, Selaginella, and Lycopodiaceae (Pryer et al. 2001). However, it is convenient to treat lycophytes together with ferns for this account. Ferns and lycophytes have a long geological history. Primitive extant families (Azollaceae, Cyatheaceae, Dicksoniaceae, Dipteridaceae, Equisetaceae, Gleicheniaceae, Isoetaceae, Lycopodiaceae, Marattiaceae, Matoniaceae, Osmundaceae, Salviniaceae, Schizaeaceae, Selaginellaceae) have a fossil record dating back to the Cretaceous period or earlier (Collinson 1996) and were contemporaneous with the dinosaurs. All of these families are found in Papua. Matoniaceae is of particular interest, because fossils of 11 or more genera are known from the Upper Triassic and Upper Cretaceous, and the family was widely distributed in Eurasia, Greenland, Australia, Africa, Madagascar, and North and South America (Kato 1998). Only two genera, Matonia and Phanerosorus, each with two species, are known today and the family is now restricted to Malesia. Matonia ranges from Peninsular Malaysia to Papua, and Phanerosorus occurs only in Borneo and Papua. Ferns are commonly considered to be rather delicate inhabitants of moist shaded sites, but in fact they are found in a great variety of habitats. Some are free-floating in still or slow-moving water (e.g., Azolla); others such as Acrostichum are tolerant of salt water. Many can tolerate full sun; these may be colonizers of roadside banks (e.g., Dicranopteris and Sphenomeris), fire survivors in grassland (e.g., Pteridium), frost survivors in grassland (e.g., Cyathea), or high epiphytes in tree crowns (e.g., Pyrrosia). Ferns also vary tremendously in size, from delicate filmy ferns with fronds one cell thick and up to 5 mm long (e.g., Microgonium motleyi), to tree ferns with trunks more than 15 m tall (e.g., Cyathea contaminans) and huge palm-like ferns with fronds more than 7 m long (e.g., Angiopteris evecta). One genus, Lecanopteris, has a hollow rhizome that is inhabited by ants; the plant provides shelter for the ants and the ants provide nutrients for the plants (Gay et al. 1993). All of these are found in Papua. Our current knowledge of the ferns and lycophytes of Papua is incomplete because accounts of only 14 families have been published for Flora Malesiana and

F

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

335

................. 16157$

CH15

02-08-07 10:29:18

PS

PAGE 335

.

336 /

few other recent taxonomic treatments are available. Families treated in Flora Malesiana to date are Azollaceae (Saunders 1998), Cheiropleuriaceae (Laferrie`re 1998a), Cyatheaceae (including Dicksoniaceae; Holttum 1963), Davalliaceae (Nooteboom 1998), Equisetaceae (Laferrie`re 1998b), Gleicheniaceae (Holttum 1959a), Isoetaceae (Alston 1959; Croft 1980), Lindsaeaceae (Kramer 1971), Lomariopsidaceae (Hennipman 1978; Holttum 1978), Matoniaceae (Kato 1998), Plagiogyriaceae (Zhang and Nooteboom 1998), Polypodiaceae (Hovenkamp et al. 1998), Schizaeaceae (Holttum 1959b), and Thelypteridaceae (Holttum 1981). In addition, the account of Grammitidaceae (Parris, in prep.) is also sufficiently advanced to give an adequate assessment of species present in Papua. Relevant recent revisions of genera include Arthropteris (Holttum 1966), Asplenium section Thamnopteris (Holttum 1974), Austrogramme (Hennipman 1975), Blechnum (Chambers and Farrant 2001), Deparia (Kato 1984), Grammitis (Parris 1983, 2004), Hypolepis (Brownsey 1987), Lycopodium section Complanata (Wilce 1965), Microgonium (Croxall 1986), Ophioglossum (Wieffering 1964), Polystichum (Nakaike 1975, 1976) Syngramma (Holttum 1975), Taenitis (Holttum 1968), and the Tectaria group (Holttum 1991). Earlier, others (Cesati 1877; Christ 1911; Ridley 1916; Gepp 1917; Alderwerelt 1924; Copeland 1940a,b, 1941a,b, 1947, 1949a,b,c, 1953) have also provided records of Papuan ferns and lycophytes, some of which have been superseded by more modern accounts. Up to date treatments are badly needed for the large families Lycopodiaceae, Selaginellaceae, Aspleniaceae, and Woodsiaceae. Much of the literature available indicates the presence of species in Papua New Guinea only, and deductions must be made about their presence in Papua; little or no habitat data is available and detailed distribution within Papua is seldom provided. Kato (1990) has estimated that the fern and lycophyte flora of the neighboring island of Seram in the Moluccas contains well over 700 species. Given the small size of Seram (ca 18,000 km2) in comparison with New Guinea (ca 800,000 km2) it seems reasonable to make a conservative estimate that New Guinea has at least 3,000 species of ferns and lycophytes, over 2,000 of which are likely to occur in Papua. The world total of fern and lycophyte species is likely to be between 15,000 and 20,000; thus Papua contains an important percentage of them.

Historical Background The first fern and lycophyte collections in Papua were made by Gaudichaud (1827), who gathered a few species from Rawak ( Lawak) Island between 1817 and 1820. Beccari collected ferns and lycophytes in the Vogelkop Peninsula, particularly in the Arfak Mts, in 1872 and 1875–1876, which were identified by Cesati (1877). Versteeg collected ferns and lycophytes on the First Lorentz Expedition in 1907 and these were written up by Christ (1911), together with those of Branderhorst. Ridley (1916) documented the ferns and lycophytes collected by Kloss on Mt Carstensz ( Mt Jaya) during 1912–1913, while Gepp (1917) identified the ferns and lycophytes collected by Gibbs in the Arfak Mts during December 1913

................. 16157$

CH15

02-08-07 10:29:19

PS

PAGE 336

Ferns and Lycophytes of Papua / 337

and elsewhere, including Manokwari and Humboldt Bay, in early 1914. Alderwerelt (1924) named the ferns and lycophytes collected by several botanists including De Kock, Feuilletau de Bruyn, Gjellerup, Janowsky, Lam, Le Cocq d’Armandville, Pulle, Thomsen, Versteeg, and Von Ro¨mer; the most important of these were Lam’s collections from Mt Doorman. Brass collected ferns and lycophytes on the Third Archbold Expedition to New Guinea, particularly in the Central Range (Snow Mts), and the ferns were identified in a series of papers by Copeland (1940a,b, 1941a,b, 1947, 1949a,b,c, 1953). Subsequent important expeditions have been made by the Royal Botanic Gardens, Kew, to the Vogelkop Peninsula during the 1990s, although the results remain unpublished, and to Mt Jaya from 1998 to 2000. The high altitude collections from 3,000 m and higher on Mt Jaya will be documented in Johns et al. (in prep.).

Ecology Ferns and lycophytes occupy all of the vegetation zones in Papua, from mangrove swamps to the alpine region, extending higher than 4,000 m, but the distribution of the families in the three major habitat divisions, lowland, montane, and alpine, is very uneven. Most pteridophyte families occur in more than one habitat division, often with altitudinal replacement of genera; for example, in Lomariopsidaceae, Elaphoglossum occurs at higher altitudes than the essentially lowland members of the family, Bolbitis, Lomagramma, Lomariopsis, Teratophyllum, and Thysanosoria. The saltwater-tolerant ferns Acrostichum aureum and A. speciosum inhabit the landward margins of mangrove swamps and areas exposed to salt spray, while epiphytic members of the Polypodiaceae may be found in mangroves. Lowland and hill rainforests from sea level up to ca 1,000 m are rich in ferns and lycophytes belonging the following families: Psilotaceae, Selaginellaceae, Adiantaceae, Marattiaceae, Nephrolepidaceae, Oleandraceae, Pteridaceae, Schizaeaceae, and Vittariaceae, which are mostly lowland in distribution; and Aspleniaceae, Blechnaceae, Cyatheaceae, Davalliaceae, Dryopteridaceae, Gleicheniaceae, Hymenophyllaceae, Lindsaeaceae, Lomariopsidaceae, Lycopodiaceae, Polypodiaceae, Thelypteridaceae, and Woodsiaceae, which are also common at higher altitudes. Dicksoniaceae and Grammitidaceae are poorly represented in lowland forests. Lygodium (Schizaeaceae) is scandent (ground-rooted, scrambling through trees and shrubs for support, but not attached to them by aerial roots) at forest margins and in clearings, with Dicranopteris and Sticherus (Gleicheniaceae) forming thickets in open areas. Blechnum (Blechnaceae), Nephrolepis (Nephrolepidaceae), and Sphenomeris (Lindsaeaceae) are also typically found at forest margins. Ground-rooted climbers (Lomagramma, Lomariopsis, Teratophyllum, Thysanosoria; all Lomariopsidaceae) grow up the trunks of trees to a considerable height and produce fertile fronds well above the ground; Stenochlaena (Blechnaceae) can likewise be a climber or scandent. Large ground ferns such as Angiopteris and Marattia (Marattiaceae), Pleocnemia and Tectaria (Dryopteridaceae), and Diplazium (Woodsiaceae) are common, as are smaller ground ferns and lycophytes such

................. 16157$

CH15

02-08-07 10:29:19

PS

PAGE 337

.

338 /

as Lindsaea (Lindsaeaceae) and Selaginella (Selaginellaceae). Large tufted epiphytes such as Aglaomorpha, Drynaria, and Platycerium (Polypodiaceae) and Asplenium sect. Thamnopteris (Aspleniaceae) are characteristic of lowland and hill forests. Small epiphytes with long-creeping rhizomes are found from ground level to the crowns of trees, such as Davallia (Davalliaceae) and Pyrrosia (Polypodiaceae) and, in humid situations, Hymenophyllaceae. Montane rainforest from ca 1,000 m up to ca 2,000 m is the most species-rich fern and lycophyte habitat in Papua. The lowland families Psilotaceae, Adiantaceae, Marattiaceae, Nephrolepidaceae, Oleandraceae, Pteridaceae, Schizaeaceae, and Vittariaceae extend to montane habitats, but are less numerous than at lower altitudes. Aspleniaceae, Blechnaceae, Cyatheaceae, Davalliaceae, Dennstaedtiaceae, Dicksoniaceae, Dryopteridaceae, Equisetaceae, Gleicheniaceae, Grammitidaceae, Hymenophyllaceae, Lindsaeaceae, Lomariopsidaceae, Lycopodiaceae, Plagiogyriaceae, Polypodiaceae, Selaginellaceae, Thelypteridaceae, and Woodsiaceae are also present. Scandent climbers, large ground ferns, and large tufted epiphytes are much less common in montane rainforest than in lowland and hill rainforests, or are absent. Small epiphytes, particularly in Grammitidaceae and Hymenophyllaceae, are characteristic and abundant. In montane rainforest above 2,000 m species numbers decline with altitude up to the tree line at approximately 3,000–3,200 m. At increasing altitudes above 3,200 m there is a rapid reduction in species numbers in alpine grassland and bare rock communities, up to the limit of ferns and lycophytes at ca 4,500 m. Adiantaceae, Lindsaeaceae, Marattiaceae, Nephrolepidaceae, Oleandraceae, Pteridaceae, Schizaeaceae, and Vittariaceae are absent or virtually so from alpine areas, but Aspleniaceae, Davalliaceae, Dryopteridaceae, Gleicheniaceae, Lomariopsidaceae, Polypodiaceae, Thelypteridaceae, and Woodsiaceae have some alpine species. Blechnaceae, Cyatheaceae, Hymenophyllaceae, and Lycopodiaceae are fairly evenly distributed from the lowland to the alpine zones, while Grammitidaceae is mainly montane to alpine, Equisetaceae and Plagiogyriaceae are montane to alpine, and Isoetaceae is entirely alpine. Montane and subalpine tree fern grasslands are widespread and characterized by thick-trunked stocky species of Cyathea with rigid small fronds. Hope (1980) documents two types of tree fern grassland alliance. The montane tree fern alliance includes communities dominated by species of Cyathea (e.g., C. aenifolia) that are also common in upper montane forest. Such communities occur below ca 3,400 m in areas that do not experience severe frosts, while the subalpine tree fern alliance is dominated by species that are not usually found in forests (e.g., C. macgregorii and C. pseudomuelleri), and occupies cold valley bottoms up to ca 3,700 m. Fern-rich open heathland, sedgeland, and low shrubland is widespread at higher altitudes (Hope 1980) and can be dominated by Gleichenia and Sticherus (both Gleicheniaceae) and Plagiogyria glauca (syn. P. papuana, Plagiogyriaceae), with Polystichum cheilanthoides (Dryopteridaceae) a minor component. Isoetes forms distinctive aquatic communities in tarn and lake floors in water

................. 16157$

CH15

02-08-07 10:29:20

PS

PAGE 338

Ferns and Lycophytes of Papua / 339

averaging 50–100 cm in depth (Hope 1980) and also occurs away from water in subalpine to alpine meadows and bogs (Johns et al., in prep.).

Biogeography The ferns and lycophytes of Papua can be assigned to several biogeographical elements. The cosmopolitan element is small, ca 2%, and contains mostly species of open habitats, often occurring in numerous vegetation types over a wide altitudinal range. The Old World tropics element, ca 6%, also has numerous species of open habitats and wide ecological tolerance. The Southeast Asia-Malesia-Pacific element, in which species extend beyond the northern and western boundaries of Malesia to Taiwan, Thailand, or Indochina, or further afield to China, India, and Sri Lanka, and beyond the southern and eastern boundaries to Australia and the Solomon Islands or further into the Pacific Ocean, contains ca 13% of the species, many of which occur in lowland to lower montane rainforest. Likewise, many of the Southeast Asia-Malesia species, ca 10%, are in lowland to lower montane forest, as are many of the Malesia-Pacific species, ca 14%. Species restricted to Malesia form ca 19% of the total and ca 6% of these occur only in the Moluccas and New Guinea; they are found in a variety of habitats ranging from lowland rainforest to alpine areas. New Guinea endemic species account for ca 16% of the total, and Papua endemic species comprise the remaining ca 20%. Like the Malesian endemic species, both New Guinea endemics and Papua endemics are found in a variety of habitats. Figure 3.4.1 shows the biogeography of Papuan ferns and lycophytes.

Endemism Papuan endemic ferns and lycophytes are most numerous in montane forest between 1,400 and 2,600 m. Less than 10% of ferns and lycophytes at altitudes from sea level up to 2,200 m are endemic to Papua, but the endemics become more important with increasing altitude, to the extent that at 4,000 m and above they are 30% or more of the total species. There are no fern and lycophyte families endemic to Papua, but a monotypic genus, Thysanosoria (Lomariopsidaceae) is endemic to the Vogelkop Peninsula (Holttum 1978). Recent expeditions to Seram, in the eastern Moluccas adjacent to Papua, have collected numerous ferns previously regarded as Papua endemics (e.g., Grammitis ahenobarba). Species endemism in Papuan ferns and lycophytes can only be assessed practically for genera that have been recently revised. The number of species in Papua and endemic to Papua is listed in Table 3.4.1, which shows that endemism is important in Cyatheaceae, Dryopteridaceae (Polystichum), Grammitidaceae, Isoetes, Lomariopsidaceae, and Thelypteridaceae. Heywood and Davis (1995) list several centers of plant diversity in Papua, of which Mt Lorentz and Mamberamo-Pegunungan Jayawijaya are likely to be of major importance for ferns and lycophytes because they range from sea level to

................. 16157$

CH15

02-08-07 10:29:20

PS

PAGE 339

.

340 /

Figure 3.4.1. Biogeography of Papuan ferns and lycophytes. over 4,600 m in altitude and thus cover all of the fern and lycophyte habitats. Numerous endemics are known from both areas. The Arfak Mts (100–3,100 m alt.) and Waigeo Island (0–999 m alt.) lack upper montane and alpine habitats, but both have endemic fern and lycophyte species.

Literature Cited Alderwerelt, C.R.W.H. 1924. Pteridophyta. Nova Guinea 14: 1–172. Alston, A.H.G. 1959. Isoetaceae. Flora Malesiana ser. II, 1 (1): 62–64. Brownsey, P.J. 1987. A review of the fern genus Hypolepis (Dennstaedtiaceae) in the Malesian and Pacific regions. Blumea 32: 227–276. Cesati, V. 1877. Prospetto delle Felci raccolte dal Signor O. Beccari nella Polinesia durante il suo secondo viaggio di esplorazione in que’mari. Rend. R. Accad. Sci. Fis. Math. Napoli 16: 23–31. Chambers, T.C., and P.A. Farrant. 2001. Revision of Blechnum (Blechnaceae) in Malesia. Blumea 46: 283–350. Christ, H. 1911. Filices. Nova Guinea 8: 149–164. Collinson, M.E. 1996. ‘‘What use are fossil ferns?’’—20 years on: with a review of the fossil history of extant pteridophyte families and genera. Pp. 349–394 in Camus, J.M., M. Gibby, and R.J. Johns (eds.) Pteridology in Perspective. Royal Botanic Gardens, Kew. Copeland, E.B. 1940a. Oleandrid ferns (Davalliaceae) in New Guinea. Philipp. J. Sci. 73 (3): 345–357. Copeland, E.B. 1940b. Notes on Hymenophyllaceae. Philipp. J. Sci. 73 (4): 457–468.

................. 16157$

CH15

02-08-07 10:29:53

PS

PAGE 340

Ferns and Lycophytes of Papua / 341

Table 3.4.1. Endemism in recently revised fern and lycophyte genera

Family Cyatheaceae Davalliaceae Dicksoniaceae Dryopteridaceae

Isoetaceae Grammitidaceae

Lindsaeaceae

Lomariopsidaceae

Polypodiaceae Thelypteridaceae

Genus

Number of species in Papua

Number of species endemic to Papua

Cyathea

33

11

Davallia

13

1

Dicksonia

4

1

1 2 2 8 9 21

1 1 1 1 4 3

4

3

Acrosorus Calymmodon Ctenopteris Grammitis Themelium Xiphopteris

3 9 17 38 9 6

1 3 2 14 2 2

Lindsaea Tapeinidium

21 6

3 3

Elaphoglossum Lomagramma Thysanosoria

21 5 1

7 2 1

Aglaomorpha

6

1

Christella Coryphopteris Plesioneuron Pneumatopteris Pronephrium Sphaerostephanos

7 13 12 8 7 30

1 6 8 5 1 9

................. 16157$

CH15

Aenigmopteris Cyclopeltis Heterogonium Pleocnemia Polystichum Tectaria Isoetes

02-08-07 10:29:54

PS

PAGE 341

.

342 /

Copeland, E.B. 1941a. Gleicheniaceae of New Guinea. Philipp. J. Sci. 75 (4): 347–359. Copeland, E.B. 1941b. Miscellaneous ferns of New Guinea. Philipp. J. Sci. 76 (1): 23–25. Copeland, E.B. 1947. Cyathea in New Guinea. Philipp. J. Sci. 77 (2): 95–125. Copeland, E.B. 1949a. Pteridaceae of New Guinea. Philipp. J. Sci. 78 (1): 5–41. Copeland, E.B. 1949b. Aspleniaceae and Blechnaceae of New Guinea. Philipp. J. Sci. 78 (2): 207–229. Copeland, E.B. 1949c. Aspidiaceae of New Guinea. Philipp. J. Sci. 78 (4): 389–475. Copeland, E.B. 1953. Grammitidaceae of New Guinea. Philipp. J. Sci. 81 (2): 81–118. Croft, J.R. 1980. A taxonomic revision of Isoetes L. (Isoetaceae) in Papuasia. Blumea 26: 177–190. Croxall, J.P. 1986. Microgonium (Hymenophyllaceae) in Malesia, with special reference to Peninsular Malaysia. Kew Bull. 41 (3): 519–531. Gaudichaud, C. 1827. Botanique. Pp. 147–165 in Freycinet, L. (ed.) Voyage autour du monde sur l’Uranie et la Physicienne. Pilletaine´, Paris. Gay, H., E. Hennipman, C.R. Huxley, and F.J.E. Parrott. 1993. The taxonomy, distribution and ecology of the epiphytic Malesian Ant-fern Lecanopteris Reinw. (Polypodiaceae). Gardens’ Bull. 45: 293–335. Gepp, A. 1917. Pteridophyta. Pp. 67–78, 192–197 in Gibbs, L.S. (ed.) Dutch N.W. New Guinea. A Contribution to the Phytogeography and Flora of the Arfak Mountains Etc. Taylor and Francis, London. Hennipman, E. 1975. A re-definition of the gymnogrammoid genus Austrogramme Fournier. Fern Gaz. 11: 61–72. Hennipman, E. 1978. Bolbitis. Flora Malesiana ser. II, 1 (4): 314–330. Heywood, V.H., and S.D. Davis. 1995. Centres of Plant Diversity. A Guide and Strategy for Their Conservation. Volume 2. Asia, Australasia and the Pacific. IUCN, Cambridge. Holttum, R.E. 1959a. Gleicheniaceae. Flora Malesiana ser. II, 1 (1): 1–36. Holttum, R.E. 1959b. Schizaeaceae. Flora Malesiana ser. II, 1 (1): 37–61. Holttum, R.E. 1963. Cyatheaceae [including Dicksoniaceae]. Flora Malesiana ser. II, 1 (2): 65–158. Holttum, R.E. 1966. The genus Arthropteris in Malesia. Blumea 14: 225–229. Holttum, R.E. 1968. A re-definition of the fern genus Taenitis Willd. Blumea 16: 87–95. Holttum, R.E. 1974. Asplenium Linn., sect. Thamnopteris Presl. Gardens’ Bull. 27: 143–154. Holttum, R.E. 1975. A comparative account of the fern genera Syngramma J. Sm. and Taenitis Willd., with discussion of their relationships to each other and to other genera. Kew Bull. 30: 327–343. Holttum, R.E. 1978. Lomariopsis group. Flora Malesiana ser. II, 1 (4): 255–314. Holttum, R.E. 1981. Thelypteridaceae. Flora Malesiana ser. II, 1 (5): 331–599. Holttum, R.E. 1991. Tectaria group. Flora Malesiana ser. II, 2 (1): 1–132. Hope, G.S. 1980. New Guinea mountain vegetation communities. Pp. 153–222 in van Royen, P. (ed.) The Alpine Flora of New Guinea, Vol. 1. J. Cramer, Vaduz. Hovenkamp, P.H., M.T.M. Bosman, E. Hennipman, H.P. Nooteboom, G. Ro¨dl-Linder, and M.C. Roos. 1998. Polypodiaceae. Flora Malesiana ser. II, 3: 1–234. Johns, R.J., et al. In prep. A Guide to the Subalpine and Alpine Flora of Mount Jaya, New Guinea. Royal Botanic Gardens, Kew. Kato, M. 1984. A taxonomic study of the athyrioid fern genus Deparia with main reference to the Pacific species. J. Fac. Sci. Univ. Tokyo 3, 13: 375–429. Kato, M. 1990. The fern flora of Seram. Pp. 225–234 in Baas, P., K. Kalkman, and R. Geesing (eds.) The Plant Diversity of Malesia. Kluwer Academic, Dordrecht.

................. 16157$

CH15

02-08-07 10:29:55

PS

PAGE 342

Ferns and Lycophytes of Papua / 343 Kato, M. 1998. Matoniaceae. Flora Malesiana ser. II, 3: 289–294. Kramer, K.U. 1971. Lindsaea-group. Flora Malesiana ser. II, 1 (3): 177–254. Laferrie`re, J.E. 1998a. Cheiropleuriaceae. Flora Malesiana ser. II, 3: 285–286. Laferrie`re, J.E. 1998b. Equisetaceae. Flora Malesiana ser. II, 3: 287–288. Moran, R.C. 2004. A Natural History of Ferns. Timber Press, Portland. Nakaike, T. 1975. A contribution to the fern flora of New Guinea (I). Pp. 99–121 in Publication by The Party 1973–’74 The Botanical Expedition to Papua New Guinea. The National Science Museum, Tokyo. Nakaike, T. 1976. A contribution to the fern flora of New Guinea (II). Bull. Nat. Sci. Mus. Tokyo 2: 47–52. Nooteboom, H.P. 1998. Davalliaceae. Flora Malesiana ser. II, 3: 235–276. Parris, B.S. 1983. A taxonomic revision of the genus Grammitis Swartz (Grammitidaceae: Filicales) in New Guinea. Blumea 29: 13–222. Parris, B.S. 2004. Three new species of Grammitidaceae (Filicales) from New Guinea. Contributions to the flora of Mount Jaya XIV. Kew Bull. 59: 219–222. Parris, B.S. In prep. Grammitidaceae. Flora Malesiana ser. II. Pryer, K.M., H. Schneider, A.R. Smith, R. Cranfill, P.G. Wolf, J.S. Hunt, and S.D. Sipes (2001). Horsetails and fens are a monophyletic group and the closest living relatives to seed plants. Nature 409: 618–622. Ridley, H.N. 1916. Report on the botany of the Wollaston Expedition to Dutch New Guinea. Filices. Trans. Linn. Soc. London (Bot.) 2 (9): 251–264. Saunders, R.M.K. 1998. Azollaceae. Flora Malesiana ser. II, 3: 277–284. Wieffering, J.H. 1964. A preliminary revision of the Indo-Pacific species of Ophioglossum (Ophioglossaceae). Blumea 12: 321–337. Wilce, J.H. 1965. Section Complanata of the Genus Lycopodium. Nova Hedwigia Beihefte 19. Verlag von J. Cramer, Weinheim. Zhang, X.C., and H.P. Nooteboom. 1998. Plagiogyriaceae. Flora Malesiana ser. II, 3: 295–316.

................. 16157$

CH15

02-08-07 10:29:55

PS

PAGE 343

3.5. Gymnosperms of Papua

Number of Orders, Families, Genera, and Species are seed plants that have ‘‘naked’’ seeds, not being enclosed by carpels. They first developed in Paleozoic times and were abundant in the flora during Paleozoic and Mesozoic times (Biswas and Johri 1997). There are four orders of gymnosperms in the world, including Gnetales, Ginkgoales, Cycadales, and Coniferales. Gnetales consist of three genera: Gnetum L., Ephedra L., and Welwitschia Hook. F. Ginkgoales is a monotypic order, with one species (Ginkgo biloba L.) in central and southwestern China. Cycadales have three families, 10 genera, and about 130 species. Coniferales, the largest gymnosperm order, include eight families, 69 genera, and about 630 species. There are about 25–33 species of gymnosperms in Papua belonging to three orders, five families, and 12 genera.

G

Family Classification and Distribution Gnetales is an isolated group among seed plants; the evolutionary relationship of this order with the remaining seed plants remains elusive. They may be closely related to Pinaceae (Chaw et al. 2000), or the entire Coniferales (Chaw et al. 1997; Bowe et al. 2000), or the angiosperms (Crane 1985; Doyle and Donoghue 1986a). Gnetaceae is monotypic, and the single genus, Gnetum, has over 30 species and a wide geographic distribution in tropical and subtropical forests. Plants are dioecious (plants contain either male or female reproductive parts) climbers, or less often trees and shrubs; the leaves are opposite with netted venation; cones are arranged in whorls; seeds are enclosed in a fleshy, brightly colored envelope. Three species occur in Papua, including G. gnemon L., G. gnemonoides Brongn., and G. latifolium Bl. (Maheshwari and Vasil 1961).

This is a monophyletic order; however, its systematic position has not been satisfactorily resolved (Soltis et al. 2002). Of the three families of Cycadales (Cycadaceae, Zamiaceae, and Stangeriaceae), one (Cycadaceae) is distributed in Papua. Cycadaceae are palm-like trees that look like tree-ferns, with unbranched upright stems; the compound leaves are crowded towards the stem apex, leaflets are circinate when young, with a midvein and no lateral veins; megasporophylls are leaflike, and loosely clustered near stem apex without forming a cone. Each megasporophyll is pinnately lobed or toothed above the ovules, with two to eight ovules attached laterally at the basal portion. Four species are found in Papua: Cycas Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

344

................. 16157$

CH16

02-08-07 10:29:56

PS

PAGE 344

Gymnosperms of Papua / 345

schumanniana Lauterb. in open grasslands, C. rumphii Miq. and C. scratchleyana F. Muell. in rainforests, and C. campestris K. D. Hill in disturbed areas of lowland forests (de Laubenfels 1998).

Coniferales are distributed worldwide, with an interesting geographic structure at the family level; Araucariaceae and Podocarpaceae are mostly limited to the Southern Hemisphere, while Pinaceae, Taxaceae, and Cephalotaxaceae are distributed mainly in the Northern Hemisphere. However, Cupressaceae (including Taxodiaceae) occur in both hemispheres; nonetheless, in this family, there is a major clade containing two clusters of genera, one cluster is of the Southern Hemisphere and the other of the Northern Hemisphere (Gadek et al. 2000). Three families of the Coniferales occur in Papua.

Members of this family are trees; their leaves are simple, alternate, and spirally arranged. Plants are dioecious, or rarely monoecious (male and female cones are found on the same plant). Microsporangiate cones are cylindrical, singular or clustered, with many microsporophylls, each of which has 4–20 sporangia. Megasporangiate cones are elliptic or globular with many spirally arranged scales and bracts. Each ovuliferous scale has one ovule developing into one seed with or without wings. Bracts each have a triangular or caudate spiky tip. Each seed has two cotyledons. There are three genera in the Araucariaceae. Agathis and Araucaria have been recognized for over a century, while Wollemia was first described at the end of the twentieth century in New South Wales, Australia (Jones et al. 1995). Araucaria has a disjunct distribution between the South American continent and the South Pacific islands, whereas Agathis occurs only in South Pacific islands. In Papua, there is a single species of Araucaria, A. cunninghamii var. papuana, and a single species of Agathis, A. labillardierei (de Laubenfels 1969). Both genera grow in hill forests.

Plants of Cupressaceae are trees or shrubs. Their bark often exfoliates in long strings when mature. Leaves are persistent or rarely deciduous, simple, alternate, and distributed all around branches or twisted at base to appear 2-ranked, opposite, or whorled, scale-like. Plants are monoecious. Microsporangiate cones have spirally or oppositely arranged microsporophylls, each with two to ten microsporangia on the abaxial side. Megasporangiate cones possess peltate or basally attached and flattened scales, each with 1–20 ovules on the adaxial surface. Seeds have two or three short lateral wings and 2–15 cotyledons. Plants of this family grow in diverse habitats, from wetlands to dry soils, and from sea level to high elevations in mountains. Papuacedrus H. L. Li is the only genus in Papua and the sole species of the genus, P. papuana (F. Muell.) H. L. Li, is distributed in the montane and subalpine forests from 1,300 m to 3,600 m in elevation.

................. 16157$

CH16

02-08-07 10:29:57

PS

PAGE 345

346 /

Podocarpaceous plants are trees or shrubs. Leaves vary from broadly ovate to long linear and to scale-like. Plants are dioecious (or rarely monoecious). Microsporangiate cones cylindrical, with numerous spirally positioned microsporophylls, each with two microsporangia. Megasporangiate cones with one to many scales, each with one ovule and fused to or modified into a juicy structure, epimatium. Seeds each have two cotyledons. Podocarps are distributed in tropical and subtropical, rarely cool temperate, forests; their center of diversity is in the Southern Hemisphere of the Old World. There are 18 genera and more than 180 species in the world. In Papua there are seven genera and 14 species. Dacrycarpus (Endl.) de Lauben., a segregate genus from Podocarpus L’He´r. Ex Pers., has nine species with a geographic distribution ranging from Myanmar and southern China to Fiji and New Zealand. In Papua there are four species, including D. imbricatus (Blume) de Laub., D. steupii de Laub., D. cinctus (Pilg.) de Laub., and D. compactus (Wasscher) de Laub. (de Delaubenfels 1969). Dacrydium Sol. ex G. Forst., a genus of 21 species, grows in infertile soils and has four species in Papua (D. novo-guineense Gibbs, D. nidulum de Laub., D. beccarii Parl., and D. xanthandrum Pilg. Falcatifolium de Laub., a genus of six species, has one species in Papua (F. papuanum de Laub.). Nageia Gaertn. is widely distributed from India, southern China, Taiwan, southern Japan through Malesia to New Guinea and New Britain (Farjon, 2001). One species occurs in Papua, N. wallichianus (C. presl) Kuntze (de Laubenfels 1969; Farjon 2001). Retrophyllum C. N. Page is a segregate genus from Nageia, and has a disjunct distribution between South America and South Pacific islands. Retrophyllum vitiense (Seem.) C. N. Page occurs in Papua (de Laubenfels 1969; Farjon 2001). Podocarpus is the most diversified genus of Podocarpaceae and has over 100 species ranging from tropical and southern Africa (including Madagascar) to eastern Asia, Malesia, southwestern Pacific, eastern Australia, New Zealand, Central America, Caribbean, South America to Patagonia (Farjon 2001). Podocarpus neriifolius is common in lower elevation forest in Papua (Johns 1975). Sundacarpus (J. Buchholz & N. E. Gray) C. N. Page, a segregate monotypic genus from Podocarpus (S. amarus (Blume) C. N. Page), occurs in Papua (Johns 1975). Based on DNA sequence data, this species may be recognized as a species of Prumnopitys (Kelch 2002). It has been controversial whether to recognize Phyllocladaceae (Keng, 1973; Quinn, 1986; Chaw et al. 1997; Hill, 1998; Quinn et al. 2002). In any event, Phyllocladus Rich. ex Mirb. is closely related to Podocarpaceae (Page 1990; Quinn et al. 2002). In Papua, there is a single species of Phyllocladus, P. hypophyllus Hook.f., which occurs in the upper montane forest in association with podocarps and Nothofagus (Johns 1975).

Gaps in Floristic Documentation Gymnosperm species are currently represented by only a small number of collections (Johns 1995a). Many endemics occur in isolated mountains where future

................. 16157$

CH16

02-08-07 10:29:57

PS

PAGE 346

Gymnosperms of Papua / 347

collecting is much needed (Johns 1995b). In addition, a detailed database needs to be established for specimen-based distribution maps of each species. This is the only way through which the morphological and ecological variation of individual species of gymnosperms can be better understood and their populations more effectively conserved and managed.

Other Notes The diversity of gymnosperms is poor relative to angiosperms in Papua; however, gymnosperms are ecologically important and some of them are of economic value. For example, the wood of Phyllocladus and Papuacedrus is of good quality. Large Araucaria plantations have been established in Papua New Guinea in the Bulolo area. Seeds and leaves of Gnetum are edible and the fibrous bark is used for making baskets, ropes, and nets. Species of Podocarpaceae have linear leaves and compete successfully with flowering plants in tropical montane rainforest. Detailed ecological and evolutionary study of these gymnosperms will shed light on the forest dynamics of tropical rainforests.

Literature Cited Biswas, C., and B.M. Johri. 1997. The Gymnosperms. Narosa, New Delhi. Bowe, L.M., G. Coat, and C.W. de Pamphilis. 2000. Phylogeny of seed plants based on all three genomic compartments: extant gymnosperms are monophyletic and Gnetales’ closest relatives are conifers. Proc. Nat. Acad. Sci. 97: 4092–4097. Chaw, S., C.L. Parkinson, Y. Cheng, T.M. Vincent, and J.D. Palmer. 2000. Seed plant phylogeny inferred from all three plant genomes: monophyly of extant gymnosperms and origin of Gnetales from conifers. PNAS 97: 4086–4091. Chaw, S.M., A. Zharkikh, H.M. Sung, T.C. Lau, and W.H. Li. 1997. Molecular phylogeny of extant gymnosperms and seed plant evolution-analysis of nuclear 18S rRNA sequences. Mol. Biol. Evol. 14: 56–68. Crane, P.R. 1985. Phylogenetic analysis of seed plants and the origin of angiosperms. Ann. Missouri Bot. Gard. 72: 716–793. de Laubenfels, D.J. 1969. A revision of Malesian and Pacific rainforest conifer s. I. Podocarpaceae, in Part. J. Arnold Arbor. 50: 315–369. de Laubenfels, D.J. 1998. A taxonomic revision of the genera Cycas and Epicycas ge. nov. (Cycadaceae). Blumea. 43: 351–400. Doyle, J.A., and M.J. Donoghue. 1986a. Relationship of angiosperms and Gnetales: a numerical cladistic analysis. In Spicer, R.A., and B.A. Thomas (eds.) Systematic and Taxonomic Approaches in Palaeobotany. Oxford, Clarendon Press. The Systematics Association Special Volume No. 31: 177–198. Farjon, A. 2001. World Checklist and Bibliography of Conifers. Royal Botanical Garden, Kew. Gadek, P.A., D.L. Alpers, M.M. Heslewood, and C.J. Quinn. 2000. Relationships within Cupressaceae sensu lato: a combined morphological and molecular approach. Amer. J. Bot. 87: 1044–1057. Hill, K.D. 1998. Podocarpaceae. Pp. 547–572 in Orchard, A., and P.M. McCarthy (eds.) Flora of Australia. CSIRO 48, Melbourne.

................. 16157$

CH16

02-08-07 10:29:57

PS

PAGE 347

348 / Johns, R.J. 1975. Common forest trees of Papua New Guinea, Part I. Gymnospermae. Forestry College, Bulolo, PNG. Johns, R.J. 1995a. Introduction. Curtis’s Bot. Mag. 12: 52–62. Johns, R.J. 1995b. Endemism in the Malesian flora. Curtis’s Bot. Mag. 12: 95–110. Jones, W.G., K.D. Hill, and J.M. Allen. 1995. Wollemia nobilis, a new living Australian genus and species in the Araucariaceae. Telopea 6: 173–176. Kelch, D.G. 2002. Phylogenetic assessment of monotypic genera Sundacarpus and Manoao (Coniferales: Podocarpaceae) utilising evidence from 18S rDNA sequences. Austral. J. Bot. 15: 29–35. Keng, H. 1973. On the family Phyllocladaceae. Taiwania 18: 142–145. Maheshwari, P., and V. Vasil. 1961. Gnetum. Council Sci. Ind. Res. (CSIR), New Delhi. Page, C.N. 1990. Phyllocladaceae. Pp. 317–319 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants, vol. 1. Springer-Verlag, Berlin. Quinn, C.J. 1986. Embryogeny in Phyllocladus. New Zealand J. Bot. 24: 575–579. Quinn, C.J., R.A. Price, and P.A. Gadek. 2002. Familial concepts and relationships in the conifers based on rbcL and matK sequence comparisons. Kew Bull. 57: 513–531. Soltis, D.E., P.A. Soltis, and M.J. Zanis. 2002. Phylogeny of seed plants based on evidence from eight genes. Amer. J. Bot. 89: 1670–1681.

................. 16157$

CH16

02-08-07 10:29:58

PS

PAGE 348

3.6. Angiosperms Annonaceae of Papua . . ß

.

Number of Genera and Species is a very well defined, large family comprising ca 125 genera and ca 2,500 species worldwide. The number of genera and species is highest in Southeast Asia (ca 60 genera with ca 1,200 species) and lowest in tropical Africa. The Annonaceae in New Guinea’s flora are well represented although no revision or flora accounts are available (ca 25 genera, ca 150 species, very conservative estimation). A large number of species are endemic to New Guinea including the monotypic genus Schefferomitra subaequalis (Figure 3.6.1). Petalolophus megalopus has always been considered an endemic monotypic genus of Papua but our recent molecular and morphological studies have revealed that it should be included in Pseudouvaria (Mols et al. 2004) and it has formally been transformed (Su et al. 2005).

A

Figure 3.6.1. Schefferomitra subaequalis, a monotypic genus endemic to New Guinea. Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

349

................. 16157$

CH17

02-08-07 10:30:20

PS

PAGE 349

350 /

.

.

ß

.

Distribution and Habitat Annonaceae are characteristic understory and midstory shrubs, trees, or climbers of the tropical lowland rainforests. A very few species can be found in (upper) montane forests (e.g., Goniothalamus spp., Alphonsea spp. on Mt Kinabalu). In Papua New Guinea some species of the climbing genera Mitrella and Artabotrys occur as high as 1,800–2,100 m above sea level. The family can be found in all kinds of habitats, including peat swamp, kerangas (heath forest), and monsooninfluenced forests, but only some of the climbers prefer primary forest margins and secondary (disturbed) vegetation.

Family Classification Annonaceae has always been placed within the Magnoliales based on morphological characters, which is also supported by recent molecular studies (Angiosperm Phylogeny Group 2003; Soltis et al. 2000). The closest related family is the Eupomatiaceae, which formerly was included within the Annonaceae. Macromorphologically as well as molecularly the family is well defined and easily recognized in the field and the herbarium. Nevertheless, sterile Diospyros (Ebenaceae) specimens are regularly misidentified as species of Annonaceae. Diospyros, however, lacks the characteristic wedge-shaped inner bark of Annonaceae, and Annonaceae lack the black dots on the lower surface of leaves typical for Diospyros. The infra-familial classification has been, since the family’s naming almost 200 years ago, a major point of dispute. This is reflected in the many formal and informal accounts proposed to date. Recent molecular studies revealed the cohesiveness of the family and did not support any of the former classifications. Anaxagorea is sister to the rest of the family, which basically splits into two main clades. This large split seems to be supported by pollen characters, with one clade having inaperturate pollen and the other having aperturate pollen. Most of the Asian genera are found in the latter clade (Doyle et al. 2002; Pirie et al. 2005). As most of the African and Asian genera have not yet been taxonomically revised, this part of the family remains difficult to classify to genus. Notorious is the large (more than 120 species) genus Polyalthia which is clearly polyphyletic (Mols et al. 2004). Preliminary molecular studies combined with macromorphological characters, however, raise the possibility of subdividing the genus into smaller monophyletic groups (Mols 2004).

Features of the Family The majority of the Annonaceae are trees, and exhibit the typical monopodial growth form with more or less tiered lateral side branches. Two architectural models have been described: Roux’s model and Troll’s model. Both models are easily recognized based on the arrangement of the branches on their orthotropic axis. In species with Roux’s architectural model the branches are spirally arranged on the

................. 16157$

CH17

02-08-07 10:30:20

PS

PAGE 350

Angiosperms: Annonaceae of Papua / 351

trunk, while in species with Troll’s model the branches are distichously arranged. In the field the family can be recognized by their simple, pinnate, exstipulate leaves, mostly distichously arranged. The outer bark of young twigs is often lozenge-shaped, the inner bark wedge-shaped and has very long, strong fibers; sap in any form is absent. Habit: trees, shrubs (dwarf shrub Ellipeiopsis cherrevensis) or woody climbers (with hook-like inflorescences in Artabotrys); stilt-roots sometimes present (Xylopia); branching whorled, the crown often pyramidal. Bark without sap on cutting. Wood white or cream-colored, sometimes darkening after cutting, soft; rays wedge-shaped, fibers long and strong. Indumentum: twigs and leaves glabrous or hairy, glabrescent, hairs single or stellate (Uvaria, Rauwenhoffia both climbers). Stipules: absent. Leaves: simple, entire, alternate, usually distichous in plagiotropic shoots, indumentum on lower surface remaining or not; lateral veins distinct, interarching or not at the margin; venation reticulate or scalariform; minute oil dots; glands absent (except Desmos at the base of lamina). Inflorescences: from foliar axils or supra-axillary to ramiflorous (cauline), cauliflorous, sometimes flowers even on long subterranean suckers (Goniothalamus spp., Polyalthia spp.). Flowers actinomorphic, bisexual (unisexual in Pseudouvaria p.p.), sepals usually three, free or sometimes united, valvate or imbricate, petals six, in two whorls of three, valvate or imbricate, free, variously shaped, sometimes inner petals mitriform, forming a cap about the sexual organs (Goniothalamus, Schefferomitra, Pseuduvaria, Mitrephora, Orophea), stamens usually many, extrorse, opening by longitudinal slits, 2-loculed, connective often prolonged beyond pollensacs or covering them (uvariod type), seldom without or very short prolongation (miliusoid type), some genera tetrasporangiate (Goniothalamus, Pseuduvaria, immature pollen sacs then septate). Carpel(s) numerous to one, free (connate in introduced fruit trees of Annona), sessile or stalked, superior; style absent or short, stigma not lobed, sometimes horseshoe-shaped folded; ovule(s) numerous to one, axillary or basal if only one. Fruitlets berry-like, oblong to (sub)globose, pulp often sweet and edible when ripe, sometimes dehiscing by a longitudinal suture (Xylopia p.p.). Aril absent or present (Xylopia). Seeds with a testa of two or sometimes three layers: endocarp hard, ruminate, containing oil, sometimes starch. Embryo small, near the base of the seed.

Gaps in Floristic Documentation or Knowledge The few Asian Annonaceous genera that have recently been revised revealed a very high diversity in New Guinea, although most species are usually represented by very few specimens in the herbaria. The center of diversity of Pseuduvaria lies in New Guinea (20 of 52 species), with 95% endemism (Su 2002). It is even difficult to ascertain the number of species because many of these descriptions are based on very scanty type material. They were mostly published at the beginning of the twentieth century and many of the types were lost during World War II. In order to resolve this and other uncertainties, an increase in the

................. 16157$

CH17

02-08-07 10:30:20

PS

PAGE 351

352 /

.

.

ß

.

intensity of plant collecting (Annonaceae, but also non-Annonaceae) is considered essential. In the case of Annonaceae, collectors should try to collect more individual flowers or even flower parts, because it is difficult to study the usually leathery flowers. New and well documented material will considerably increase our general knowledge of the Annonaceae and will most likely add to the number of species recorded from New Guinea. Special attention should be given to the collection of data on flower biology because practically nothing is known about this from the eastern Malesian region.

Natural History and Floral Biology Annonaceae species may be pollinated by thrips, flies, or beetles, or sometimes even by bees (Silberbauer-Gottsberger et al. 2003). Nitidulidae (sap beetles) and Curculionidae (weevils), which live on decaying fruits, have been reported mainly for many South American species. Uvaria elmeri (Sarawak), an exception, is pollinated by cockroaches. The relatively large flowers typical of this family function in some cases as a kind of pollination chamber, in which the petals remain closed over the sexual organs. Protogyny is widespread in flowers of Annonaceae and possibly related to a preponderance of cross-pollination. Some flowers are showy, 3–4 cm in diameter (Uvaria) and bright red. Flowers of Cananga odorata with very sweet smell produce the well known ylang-ylang oil which is highly appreciated in the perfume industry. Our collections show that many trees have been planted in villages in New Guinea. Dasymaschalon and Pseuduvaria megalopus (Figure 3.6.2) might have horticultural value because of their attractive flowers, size of plant, and fruits. Flowering and fruiting generally occur throughout the year, although some species show a deciduous life cycle by flowering after leaves have been shed at the end of the dry season in Thailand, India, Indochina, and eastern Indonesia (Miliusa p.p., Mitrephora p.p.; Mols and Keßler 2003b; Weerasooriya 2001).

Fruits and Dispersal The berry-like fruitlets, often colored yellow and exuding a sweet fruity smell, suggest dispersal by birds, especially fruit pigeons and hornbills. Apes (Mezzettia and Polyalthia spp. are commonly eaten by orangutans), monkeys, fruit bats, and rodents may disperse fruits of seeds as well. Dissemination by water may occur in Miliusa macropoda, a riverine species of Borneo.

Germination and Seedlings Seeds have a short life and may remain viable for a restricted period only. Except for some climbers, they germinate only in a damp, shady environment and, therefore, natural regeneration in disturbed forest is poor. In their natural habitat re-

................. 16157$

CH17

02-08-07 10:30:21

PS

PAGE 352

Angiosperms: Annonaceae of Papua / 353

Figure 3.6.2. The attractive flowers of Pseuduvaria megalopus may have horticultural value in the future. generation is often profuse and during mast flowering years the forest floor under the mother tree is covered by seedlings. Under nursery conditions, however, the survival rate of seedlings is poor, especially when nutrients from the endosperm are exhausted. Germination is (mostly) hypogeal. The cotyledons remain within the testa; the taproot and hypocotyls emerge. The shoot is erect, initially with reduced leaves (cataphylls), borne spirally, exhibiting the Horsfieldia type and Pseuduvaria subtype (de Vogel 1979), an uncommon type in tropical woody dicotyledons.

Literature Cited Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399–436. Chatrou, L.W., H. Rainer, and P.J.M. Maas. 2004. Annonaceae. Pp. 18–20 in Smith, N., S.A. Mori, A. Henderson, D.W. Stevenson, and S.V. Heald (eds.) Flowering Plants of the Neotropics. The New York Botanical Garden and Princeton University Press, Princeton, New Jersey. de Vogel, E.F. 1979. Seedlings of Dicotyledones. Pudoc, Wageningen, the Netherlands. Doyle, J.A., P. Bygrave, and A. Le Thomas. 2002. Implications of molecular data for pollen

................. 16157$

CH17

02-08-07 10:30:41

PS

PAGE 353

354 /

.

.

ß

.

evolution in Annonaceae. Pp. 259–284 in Harley, M.M., C.M. Morton, and S. Blackmore (eds.) Pollen and Spores: Morphology and Biology. Royal Botanic Gardens, Kew. Gottsberger, G. 1999. Pollination and evolution in neotropical Annonaceae. Pl. Spec. Biol. 14: 143–152. Keßler, P.J.A. 1993. Annonaceae. Pp. 93–129 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. 2. Springer Verlag, Berlin. Mols, J.B. 2004. From Miliusa to Miliuseae to Miliusoid: identifying clades in Asian Annonaceae. Ph.D. diss., National Herbarium Nederland, Leiden. Mols, J.B., B. Gravendeel, L.W. Chatrou, M.D. Pirie, P.C. Bygrave, M.W. Chase, and P.J.A. Keßler. 2004. Identifying clades in Asian Annonaceae: monophyletic genera in the polyphyletic Miliuseae. Amer. J. Bot. 91: 590–600. Mols, J.B., and P.J.A. Keßler 2003. Studies in the Miliuseae. V. Review of the taxonomic history of a polyphyletic tribe. Telopea 10: 113–124. Mols, J.B., and P.J.A. Keßler. 2003a. The genus Miliusa (Annonaceae) in the AustroMalesian area. Blumea 48: 421–462. Pirie, M.D., L.W. Chatrou, R.H.J. Erkens, J.W. Maas, T. van der Niet, J.B. Mols, and J.E. Richardson. 2005. Phylogeny reconstruction and molecular dating in four neotropical genera of Annonaceae: the effect of taxon sampling in age estimations. Pp. 149–174 in Bakker, F.T., L.W. Chatrou, B. Gravendeel, and P.B. Pelser (eds.) Plant Species-level Systematics: New Perspectives on Pattern and Process, A.R.G. Ganter Verlag, Ruggell. Qiu, Y.L., J. Lee, F. Bernasconi-Quadroni, D.E. Soltis, P.S. Soltis, M. Zanis, E.A. Ziommer, Z. Chen, V. Savolainen, and M.W. Chase. 2000. Phylogeny of basal angiosperms: analysis of five genes from three genomes. Int. J. Pl. Sci. 161, Supplement 6: 3–27. Silberbauer-Gottsberger, I., G. Gottsberger, and A.C. Webber. 2003. Morphological and functional flower characteristics of New and Old World Annonaceae with respect to their mode of pollination. Taxon 52: 701–718. Soltis, D.E., P.S. Soltis, M.W. Chase, M.E. Mort, D.C. Albach, M. Zanis, V. Savolainen, W.H. Hahn, S.B. Hoot, M.F. Fay, M. Axtell, S.M. Swensen, L.M. Prince, W.J. Kress, K.C. Nixon, and J.S. Farris. 2000. Angiosperm phylogeny inferred from 18S rRNA, rbcL and atpB sequences. Bot. J. Linn. Soc. 133: 381–461. Su, Y.C.F. 2002. Systematics and phylogeny of Pseuduvaria (Annonaceae). Ph.D. diss., University of Hong Kong, Hong Kong. Su, Y.C.F., J.B. Mols, P.J.A. Keßler, and R.M.K. Saunders. 2005. Reassessing the generic status of Petalolophus (Annonaceae): evidence for the evolution of a distinct Sapromyophilous lineage within Pseuduvaria. Syst. Biol. 30: 494–502. Weerasooriya, A.D. 2001. Systematics, phylogeny and reproductive biology of Mitrephora (Annonaceae). Ph.D. diss., University of Hong Kong, Hong Kong.

................. 16157$

CH17

02-08-07 10:30:42

PS

PAGE 354

Apocynaceae of Papua david j. middleton Number of Genera and Species h e ap o c y na c e a e sensu lato (including Asclepiadaceae) has about 424 genera and 4,000 species, making it one of the ten largest angiosperm families. The Apocynaceae as traditionally recognized (i.e., without the inclusion of Asclepiadaceae) comprises subfamilies Rauvolfioideae with ca 915 species in 84 genera and Apocynoideae with ca 822 species in 77 genera. In Papua there are 19 genera and ca 66 species in these two subfamilies (20 genera and ca 103 species in New Guinea as a whole). The genera in Papua are Alstonia (7 species; Sidiyasa 1998), Alyxia (21 species; Middleton 2000), Carissa (1 species; Leeuwenberg and Van Dilst 2001), Cerbera (2 species; Leeuwenberg 1999), Chilocarpus (2 species; Leeuwenberg 2002), Ichnocarpus (2 species; Middleton 1994), Kopsia (1 species; Middleton 2004), Lepiniopsis (1 species), Melodinus (2 species; Leeuwenberg 2003), Ochrosia (including Neisosperma, 5 species; Hendrian 2004), Papuechites (1 species; Middleton 1995), Parsonsia (12 species; Middleton 1997a), Rauvolfia (2 species; Hendrian and Middleton 1999), Strophanthus (1 species; Beentje 1982), Tabernaemontana (2 species; Leeuwenberg 1991), Urceola (1 species; Middleton 1996b), Voacanga (1 species; Leeuwenberg 1985), and Wrightia (3 species; Middleton 2005). In addition, Anodendron (Middleton 1996a) is almost certain to be found in Papua and Carruthersia (Middleton 1997b) may be found there in the future. Lepinia is known in New Guinea only from offshore islands of eastern PNG and is unlikely to occur in Papua.

T

Distribution and Habitat Apocynaceae sensu stricto occur throughout the tropics and subtropics with a small number of species extending into the temperate zone. They occur in a wide range of habitats including wet evergreen forest, dry deciduous forest, montane forest, scrub forest, beach forest, and mangrove.

Family Classification The Apocynaceae sensu lato belongs in the Gentianales along with Gentianaceae, Loganiaceae, Gelsemiaceae, and Rubiaceae, and can be readily distinguished from those families by almost always possessing white or off-white latex. Recent phylogenetic work has suggested that the Asclepiadaceae are nested within the Apocynaceae sensu stricto (e.g., Potgieter and Albert 2001). Endress and Bruyns (2000) have proposed a combined family with five subfamilies: Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae, and Asclepiadoideae. Rauvolfioideae are characMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

355

................. 16157$

CH18

04-02-07 11:57:28

PS

PAGE 355

.

356 /

terized by stamens being free from the pistil head and mostly having corolla lobes overlapping to the left; Apocynoideae by stamens being attached to the pistil head and mostly having corolla lobes overlapping to the right; Periplocoideae by stamens with specialized structures called translators which transfer pollen, and corolla lobes valvate or overlapping to the right; Secamonoideae by the possession of non-waxy-walled pollinia and corolla lobes valvate or overlapping to the right; and Asclepiadoideae by waxy-walled pollinia and corolla lobes valvate or overlapping to the right.

Features of Subfamilies Rauvolfioideae and Apocynoideae Habit trees, shrubs or climbers, rarely herbs; buttresses and/or pneumatophores sometimes present; latex present, this usually white, less commonly creamcolored, yellowish, or bluish-white. Indumentum of simple hairs. Leaves simple, opposite or, more rarely, verticillate or spirally arranged, pinnately veined, margin entire, very rarely crenulate (Dyera in western Malesia) or toothed (Alyxia ilicifolia from Australia). Inflorescences cymose, rarely fasciculate or flowers solitary, terminal or axillary often forming panicles. Flowers bisexual, 5-merous, rarely 4-merous (in Leuconotis from western Malesia), radially symmetrical or, very rarely, slightly bilaterally symmetrical (in Allamanda from South America); calyx mostly of five more or less free lobes, sometimes fused into a tube with shortened lobes (in Voacanga, Chonemorpha), sometimes with glands at the base inside; corolla consisting of a short or long tube and erect or spreading lobes, funnelshaped, salver-shaped, platter-shaped, trumpet-shaped, urn-shaped or rotate, lobes overlapping to the left or right, more rarely valvate; stamens inserted on the inside of the corolla tube, completely included or exserted from corolla tube throat, anthers dorsifixed, sagittate or ovate, free or adnate to the pistil head, dehiscing by longitudinal slits, sometimes with the base and apex sterile; disk present or absent, if present then of two or five lobes or an annular ring, crenate or not; ovary superior or, rarely, semi-inferior, of two separate carpels united into a common style, a single bilocular ovary or a unilocular ovary; pistil head with a stigmatic base and a 2-cleft apex. Fruit a drupe, berry, capsule or follicle, solitary if from a syncarpous ovary or paired if from an apocarpous ovary, sometimes secondarily fused appearing syncarpous (in Parsonsia, Wrightia). Seeds simple, arillate, winged, with a ciliate margin or with an apical and/or basal coma.

Gaps in Floristic Documentation or Knowledge The known Apocynaceae diversity of Papua is much lower than that of Papua New Guinea but this is most likely due to the much lower density of collections in Papua compared to PNG. For example, species of the genus Anodendron are currently unknown in Papua despite the fact that there are two species, A. oblongifolium and A. whitmorei, with an apparent Moluccas/Papua New Guinea disjunct distribution. It is extremely unlikely that both species are really absent from

................. 16157$

CH18

02-08-07 10:30:09

PS

PAGE 356

Angiosperms: Apocynaceae of Papua / 357

Papua. The genus Carruthersia has not been recorded from anywhere in New Guinea despite the fact that one of the species, C. pilosa, is known from the Philippines and the Solomon Islands (Middleton 1997b). It may in the future be found in New Guinea. Also a number of taxa have extremely localized distributions, suggesting that with further collecting new localized taxa may be discovered. Others are known from very few collections, leaving gaps in our knowledge of intraspecific variation.

Additional Notes The Apocynaceae plays a particularly prominent role in our understanding of the evolution of complex floral morphologies because researchers have been able to construct a gradient of morphologically intermediate taxa between the relatively undifferentiated flowers in subfamily Rauvolfioideae and the highly modified flowers with synorganized gynostegia, pollinaria, and coronas found in subfamilies Secamonoideae and Asclepiadoideae. In four of the five subfamilies the seeds have a hair tuft at one or both ends and are dispersed by wind; in Rauvolfioideae the variety of seed dispersal types is quite large with wind dispersed seeds, edible fruits or arils, and water dispersed fruit.

Literature Cited Beentje, H.J. 1982. A monograph on Strophanthus DC. (Apocynaceae). Meded. Landb. Wag. 82–4: 1–191. Endress, M.E., and P.V. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Botanical Review 66 (1): 1–56. Hendrian. 2004. Revision of Ochrosia (Apocynaceae) in Malesia. Blumea 49: 101–128. Hendrian and D.J. Middleton. 1999. A revision of Rauvolfia L. in Malesia. Blumea 44: 449–470. Leeuwenberg, A.J.M. 1985. Voacanga Thou. Series of revisions of Apocynaceae Part XV. Wageningen Agricultural University Papers 85–3: 5–80. Leeuwenberg, A.J.M. 1991. A revision of Tabernaemontana: 1. The Old World species. Royal Botanical Gardens, Kew. Leeuwenberg, A.J.M. 1999. Series of revisions of Apocynaceae 47. The genus Cerbera L. Wageningen Agricultural University Papers 98–3: 1–64. Leeuwenberg, A.J.M. 2002. Series of revisions of Apocynaceae 52. Chilocarpus. Syst. Geogr. Pl. 72: 127–166. Leeuwenberg, A.J.M. 2003. Series of revisions of Apocynaceae 53. Melodinus. Syst. Geogr. Pl. 73: 3–62. Leeuwenberg, A.J.M., and F.J.H. Van Dilst. 2001. Series of revisions of Apocynaceae XLIX. Carissa L. Wageningen Agricultural University Papers 01.1: 3–109. Middleton, D.J. 1994. A revision of Ichnocarpus (Apocynaceae). Blumea 39: 73–94. Middleton, D.J. 1995. A revision of Papuechites (Apocynaceae). Blumea 40: 439–442. Middleton, D.J. 1996a. A revision of Anodendron A. DC. (Apocynaceae). Blumea 41: 37–68. Middleton, D.J. 1996b. A revision of Aganonerion Pierre ex Spire, Parameria Benth. and Urceola Roxb. (Apocynaceae). Blumea 41: 69–122.

................. 16157$

CH18

02-08-07 10:30:10

PS

PAGE 357

.

358 /

Middleton, D.J. 1997a. A revision of Parsonsia R. Br. (Apocynaceae) in Malesia. Blumea 42: 191–248. Middleton, D.J. 1997b. A revision of Carruthersia Seem. Blumea 42: 489–498. Middleton, D.J. 2000. Revision of Alyxia (Apocynaceae). Part 1: Asia and Malesia. Blumea 45: 1–146. Middleton, D.J. 2004. A revision of Kopsia (Apocynaceae: Rauvolfioideae). Harvard Papers in Botany 9: 89–142. Middleton, D.J. 2005. A revision of Wrightia (Apocynaceae: Apocynoideae) in Malesia. Harvard Papers in Botony 10: 161–182. Potgieter, K., and V.A. Albert. 2001. Phylogenetic relationships within Apocynaceae s.l. based on trnL intron and trnL-F spacer sequences and propagule characters. Annals of the Missouri Botanical Garden 88 (4): 523–549. Sidiyasa, K. 1998. Taxonomy, phylogeny, and wood anatomy of Alstonia (Apocynaceae). Blumea Supplement 11, Nationaal Herbarium Nederland, Leiden.

................. 16157$

CH18

02-08-07 10:30:10

PS

PAGE 358

Arecaceae of Papua . Number of Genera and Species , the palm family (Arecaceae, syn. Palmae) contains approximately 2,400 species in 189 genera (Govaerts and Dransfield 2005). Within New Guinea, more than 250 species are recognized in 31 genera: Actinorhytis (1 species), Areca (9 species), Arenga (1 species), Borassus (1 species), Brassiophoenix (2 species), Calamus (53 species), Calyptrocalyx (25 species), Caryota (2 species), Clinostigma (1 species), Cocos (1 species), Corypha (1 species), Cyrtostachys (9 species), Daemonorops (1 species), Dransfieldia (1 species), Drymophloeus (3 species), Heterospathe (19 species), Hydriastele (32 species), Korthalsia (2 species), Licuala (34 species), Linospadix (2 species), Livistona (8 species), Metroxylon (2 species), Nypa (1 species), Orania (9 species), Physokentia (1 species), Pigafetta (1 species), Pinanga (1 species), Ptychococcus (3 species), Ptychosperma (26 species), Rhopaloblaste (3 species), and Sommieria (1 species). At least 110 species occur in Papua, although this very likely represents an underestimate due to the bias towards Papua New Guinea in collecting patterns. A number of synopses of New Guinea palms are available (Essig 1977; Hay 1984; Ferrero 1997; Barfod, Banka, and Dowe 2001; Baker and Dransfield 2006a,b). A regional monograph of the palms of New Guinea is currently in preparation and will result in abundant new synonymy, many new species, and a clearer understanding of the distribution of species diversity (Baker 2002).

O

Distribution and Habitat The Arecaceae is distributed throughout the tropical and subtropical regions of the world. The highest diversity of palm species occurs in the Malesian region in which three major palm hotspots occur, namely the Sunda Shelf ( 500 species), New Guinea ( 250 species), and the Philippines (ca 140 species). New Guinea is the center of diversity for a number of genera or groups of genera, such as subtribe Linospadicinae (Calyptrocalyx, Linospadix), subtribe Ptychospermatinae (Ptychosperma, Ptychococcus, Brassiophoenix, Drymophloeus), Cyrtostachys, Heterospathe, Hydriastele (Figure 3.6.3), Orania, and Rhopaloblaste. In a number of other cases, New Guinea is a secondary center of diversity, for example, in Calamus (Figure 3.6.4) and Licuala (Figure 3.6.5), both of which are more species-rich on the Sunda Shelf, and Livistona, which is most diverse in Australia. Current phylogenetic studies of some of these groups promise to illuminate the biogeographic implications of these patterns. The majority of New Guinea palm species do not occur outside Papua or Papua New Guinea. In contrast, only three of the 29 genera are endemic to New Guinea: Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

359

................. 16157$

CH19

02-08-07 10:30:16

PS

PAGE 359

360 /

.

Figure 3.6.3. Hydriastele costata, a frequent lowland tree palm throughout New Guinea.

................. 16157$

CH19

02-08-07 10:30:45

PS

PAGE 360

Angiosperms: Arecaceae of Papua / 361

Figure 3.6.4. Calamus warburgii, one of the most common species of rattan in New Guinea.

................. 16157$

CH19

02-08-07 10:31:15

PS

PAGE 361

362 /

.

Figure 3.6.5. Licuala cf. lauterbachii, an example of this common understory palm genus.

................. 16157$

CH19

02-08-07 10:31:42

PS

PAGE 362

Angiosperms: Arecaceae of Papua / 363

Brassiophoenix, Dransfieldia, and Sommieria (Figure 3.6.6). Many of the genera are more or less widespread in New Guinea. Of the more restricted genera, Brassiophoenix, Clinostigma, and Physokentia do not occur in Papua, and Daemonorops and Dransfieldia occur only in far western Papua. In addition, Sommieria, Pigafetta (Figure 3.6.7), and Pinanga are widespread throughout Papua, but have only very limited distributions in western Papua New Guinea. Palms occur principally in primary rainforest from sea level to 2,800 m. Emergent palms can sometimes thrive outside closed forest in disturbed habitats, but smaller understory palms are usually unable to tolerate such conditions. Some species and genera are restricted to more seasonal areas in the south of the island, such as Corypha utan, Livistona benthamii, and L. muelleri. A number of taxa thrive in inundated areas, with two species, Metroxylon sagu (Figure 3.6.8) and Nypa fruticans, playing a particularly dominant role in freshwater and mangrove swamps, respectively. More unusual palm habitats include limestone cliffs and the flood zone of river banks. The relationship between palm diversity and elevation has been explored in detail (Bachman et al. 2004). Total palm species diversity decreases with increasing elevation in an almost linear fashion. However, if the decrease in available land area with elevation is taken into account, a peak in species richness is observed towards the center of the elevation range of New Guinea palms. Most genera occur throughout a large part of the total elevation range, but some have particularly broad ranges, such as Calamus, Calyptrocalyx, and Heterospathe.

Family Classification The palm family is placed in its own order, Arecales, within a large group of commelinid monocots according to the most recent phylogenetic classification of angiosperms (Angiosperm Phylogeny Group 2003). The position of the family within the commelinids is somewhat ambiguous, but it most commonly resolves as sister to all other commelinids (e.g., Chase et al. 2000). All phylogenetic studies support very strongly the monophyly of the palms. Since 1995, the palms have been subjected to a series of rigorous phylogenetic examinations (e.g., Asmussen and Chase 2001; Asmussen et al. 2006; Baker et al. 1999; Hahn 2002; Uhl et al. 1995), resulting in a new phylogenetic classification of the family (Dransfield et al. 2005). This classification recognizes five subfamilies, compared with the six included in the previous classification (Uhl and Dransfield 1987). Four of the five subfamilies, Calamoideae, Nypoideae, Coryphoideae, and Arecoideae, are represented in New Guinea.

Features of the Family Habit: trees, shrubs or climbers, often heavily armed. Stems not or only very rarely increasing in diameter with age, variously marked with nodal scars or obscured by persistent leaf bases. Leaves spirally arranged, very rarely distichous, the blade

................. 16157$

CH19

02-08-07 10:31:43

PS

PAGE 363

364 /

.

Figure 3.6.6. Sommieria leucophylla, a monotypic genus endemic to New Guinea and occurring almost exclusively in Papua.

................. 16157$

CH19

02-08-07 10:32:12

PS

PAGE 364

Angiosperms: Arecaceae of Papua / 365

Figure 3.6.7. Pigafetta filaris, a widespread calamoid palm in western Papua.

................. 16157$

CH19

02-08-07 10:32:44

PS

PAGE 365

366 /

.

strongly folded in bud, usually splitting longitudinally on emergence along either the adaxial or abaxial folds to give a compound blade, the blade palmate, pinnate or doubly-pinnate, or more rarely unsplit, in some climbing palms the leaf tip extended into a barbed whip; venation of segments parallel; leaf sheath tubular, at least at first, remaining tubular, and often producing a column at stem tip (crownshaft) or variously splitting and disintegrating into a fibrous network. Inflorescences paniculate or spicate, axillary, interfoliar, infrafoliar or sometimes aggregated in a group at the stem tip, the tip itself aborting and the stem then dying after flowering and fruiting; inflorescence bracts often conspicuous. Flowers actinomorphic, biseriate, bisexual or unisexual, the plants then either monoecious, dioecious, or polygamous; sepals almost always three, distinct or connate; petals almost always three, distinct or connate; stamens usually six but sometimes numerous or reduced to three, distinct or connate, sometimes adnate to the corolla, anthers usually dehiscing by longitudinal splits; staminodes in pistillate flower absent or variously conspicuous; gynoecium always superior, apocarpous or syncarpous, with one or three (very rarely more) fertile carpels, each with a single ovule, style absent to elongate, stigmas usually three; placentation and ovule form various; pistillode in staminate flowers absent to conspicuous, variously shaped. Fruits berries or drupes. Seeds one to three, endosperm homogeneous or ruminate. Germination adjacent or remote.

Gaps in Floristic Documentation or Knowledge A complete account of the palms of New Guinea will be available soon as a result of current taxonomic research towards a regional monograph for New Guinea palms. Currently, more than 3,300 palm specimens are available for this project, but less than a third of these originate from Papua (see Bachman et al. 2004 for distribution of collecting localities). It comes as no surprise that areas around Manokwari and Jayapura are relatively well collected. Recent collecting efforts have yielded material from new areas such as Timika, Nabire, Yapen, Biak, the Wandammen Peninsula, Etna Bay, Sorong, and the Raja Ampat Islands. Unfortunately, little is known of palm diversity in much of the remainder of Papua.

Natural History Palms are a conspicuous component of New Guinea vegetation below 2,800 m. They display a wide variety of growth forms. In addition to archetypal robust canopy tree palms (e.g., Hydriastele costata (Figure 3.6.3), Rhopaloblaste ceramica, Livistona papuana), there are moderate-sized midstory palms (e.g., Calyptrocalyx albertisianus, Hydriastele microspadix, Orania parva), diminutive understory palms (e.g., Calyptrocalyx micholitzii, Hydriastele rhopalocarpa, Linospadix albertisiana), ‘‘stemless’’ palms (e.g., Heterospathe delicatula, H. humilis), and rattans (e.g., Calamus species, Daemonorops species, Korthalsia zippelii). As mentioned above, two species dominate certain vegetation types: Nypa fruticans forms extensive mono-

................. 16157$

CH19

02-08-07 10:32:45

PS

PAGE 366

Angiosperms: Arecaceae of Papua / 367

dominant stands in brackish swamps in front of mangroves, and Metroxylon sagu (Figure 3.6.8), the sago palm, forms impenetrable, spiny forest in lowland areas inundated with fresh water. In the case of M. sagu, the extent of natural sago swamp is not known because sago stands are often semi-cultivated for the extraction of sago starch for human consumption. Nypa and sago swamps represent highly significant carbon sinks. Nypa swamps are also very important in the stabilization of coastlines and in providing breeding grounds for fish and crustaceans. Some other palm species, usually rattans, can behave as pioneers in disturbed areas (e.g., Calamus warburgii (Figure 3.6.4), Korthalsia zippelii). A small number of palms in New Guinea (e.g., Calamus reticulatus, Heterospathe macgregorii, and Hydriastele rheophytica) are adapted to grow within the flood zones of large rivers prone to seasonal flooding (Baker 1997; Dowe and Ferrero 2000). These rheophytes tend to be strongly clustered (and thus able to regenerate easily after damage) and have narrowly pinnate leaves (thus presenting little resistance when submerged). Rather little is known about the ecology of New Guinea palms. Certain morphological characteristics suggest that palms would be productive subjects for ecological studies. For example, numerous species of rattan in New Guinea possess a highly developed ocrea, a tongue-like extension of the leaf sheath beyond the insertion of the leaf petiole (Baker and Dransfield 2002). The ocrea is a papery or leathery structure ranging from a few centimeters (e.g., Calamus altiscandens) to almost one meter long (Calamus zebrinus) that usually clasps the sheathed stem and that sometimes disintegrates with age. The ocreas are often occupied by ants, creating a striking parallel with Korthalsia species in the Sunda region (Dransfield 1981). All the more remarkable is the fact that ocreas, which are rare in Calamus outside New Guinea, occur in a number of unrelated groups of New Guinean Calamus species. Ants also build nests among the leaf sheath spines of rattans and, in Calamus retroflexus, within a space created by basal leaflets reflexing across the rattan stem. A few pollination studies of New Guinea palms were carried out by Essig (1973). He made observations on Ptychosperma macarthurii, Hydriastele microspadix, and Nypa fruticans. Syrphid flies and bees belonging to the genus Nomia were the most significant transporters of pollen between the staminate and pistillate flowers in monoecious, protandrous Ptychosperma and were suggested as being the most likely pollinators. Hydriastele microspadix, while being monoecious is, in contrast, protogynous with a very rapid transition from pistillate to staminate anthesis shortly after the fall of the two large enclosing inflorescence bracts. Although several groups of insects were recorded visiting the inflorescences, Essig concluded that weevils (Curculionidae) were the most likely effective pollinators. In monoecious Nypa, Essig observed drosophilid flies transporting pollen and visiting both staminate and pistillate flowers, and concluded that they were likely to be the pollinators. Elsewhere palm pollination has proved to be a very fruitful and exciting area of research, demonstrating some remarkable examples of coevolution,

................. 16157$

CH19

02-08-07 10:32:45

PS

PAGE 367

368 /

.

Figure 3.6.8. Metroxylon sagu, very abundant in swamps throughout lowland New Guinea in wild and semi-domesticated stands.

................. 16157$

CH19

02-08-07 10:33:09

PS

PAGE 368

Angiosperms: Arecaceae of Papua / 369

and it seems very likely that detailed studies of other New Guinea palm pollination could be very rewarding. Most palm fruits are fleshy and brightly colored when ripe and are undoubtedly attractive to birds and mammals. In two unrelated groups, subtribe Ptychospermatinae (especially Ptychococcus and Brassiophoenix) and some species of Licuala (e.g., L. beccariana), seeds are defended by thick ornate, bony endocarps that bear numerous wings and grooves. It is tempting to infer that this represents a protection against damage during passage through the gut of a frugivore or against the jaws of seed predators, such as parrots and cockatoos.

Literature Cited Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: 399–436. Asmussen, C.B., and M.W. Chase. 2001. Coding and noncoding plastid DNA in palm family systematics. American Journal of Botany 88: 1103–1117. Asmussen, C.B., J. Dransfield, V. Deickmann, A.S. Barfod, J.-C. Pintaud, and W.J. Baker. 2006. A new subfamily classification of the palm family (Arecaceae): evidence from plastid DNA. Botanical Journal of the Linnean Society 151: 15–38. Bachman, S., W.J. Baker, N. Brummitt, J. Dransfield, and J. Moat. 2004. Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea. Ecography 27: 299–310. Baker, W.J. 1997. Rattans and rheophytes, palms of the Mubi river. Principes 41:148–157. Baker, W.J. 2002. The palms of New Guinea project. Flora Malesiana Bulletin 13: 35–37. Baker, W.J., C.B. Asmussen, S. Barrow, J. Dransfield, and T.A. Hedderson. 1999. A phylogenetic study of the palm family (Palmae) based on chloroplast DNA sequences from the trnL–trnF region. Plant Systematics and Evolution 219: 111–126. Baker, W.J., and J. Dransfield. 2002. Calamus longipinna (Arecaceae: Calamoideae) and its relatives in New Guinea. Kew Bulletin 57: 853–866. Baker, W.J., and J. Dransfield. 2006a. Field Guide to the Palms of New Guinea. Royal Botanic Gardens, Kew. Baker, W.J., and J. Dransfield. 2006b. Sebuah Panduan Lapangan untk Palem New Guinea. (Trans. Ary P. Keim.) Royal Botanic Gardens, Kew. Barfod, A.S., R. Banka, and J.L. Dowe. 2001. Field guide to the palms in Papua New Guinea—with a multi-access key and notes on the genera. AAU Reports 40. Department of Systematic Botany, University of Aarhus, Aarhus. Chase, M.W., D.E. Soltis, P.S. Soltis, P.J. Rudall, M.F. Fay, W.J. Hahn, S. Sullivan, J. Joseph, M. Molvray, P.J. Kores, T.J. Givnish, K.J. Sytsma, and J.C. Pires. 2000. Higherlevel systematics of the monocotyledons: an assessment of current knowledge and a new classification. Pp. 3–16 in Wilson, K.L., and D.A. Morrison (eds.) Systematics and Evolution of Monocots. CSIRO Publishing, Victoria. Dowe, J.L., and M.D. Ferrero. 2000. A new species of Rheophytic palm from New Guinea. Palms 44: 194–197. Dransfield, J. 1981. A synopsis of Korthalsia (Palmae: Lepidocaryoideae). Kew Bulletin 36: 163–194. Dransfield, J., and N.W. Uhl. 1998. Palmae. Pp. 306–389 in Kubitzki, K. (ed.) Families

................. 16157$

CH19

02-08-07 10:33:10

PS

PAGE 369

370 /

.

and Genera of Vascular Plants. Volume IV. Flowering Plants. Monocotyledons. Springer, Berlin. Dransfield, J., N.W. Uhl, C.B. Asmussen, W.J. Baker, M.M. Harley, and C.E. Lewis. 2005. A new phylogenetic classification of the palm family, Arecaceae. Kew Bulletin 60: 559–569. Essig, F.B. 1973. Pollination in some New Guinea palms. Principes 17: 75–83. Essig, F.B. 1977. A preliminary analysis of the palm flora of New Guinea. Botany Bulletin 9. Office of Forests, Division of Botany, Lae. Ferrero, M.D. 1997. A checklist of Palmae for New Guinea. Palms and Cycads 55/56: 2–39. Govaerts, R., and J. Dransfield. 2005. World Checklist of Palms. Royal Botanic Gardens, Kew. Hahn, W.J. 2002. A molecular phylogenetic study of the Palmae (Arecaceae) based on atpB, rbcL, and 18S nrDNA sequences. Systematic Biology 51: 92–112. Hay, A.J. 1984. Palmae. Pp. 195–318 in Johns, R.J., and A.J. Hay (eds.) A Guide to the Monocotyledons of Papua New Guinea, 3. Papua New Guinea University of Technology, Lae. Uhl, N.W., and J. Dransfield. 1987. Genera Palmarum: A Classification of Palms Based on the Work of H. E. Moore, Jr. International Palm Society and L. H. Bailey Hortorium, Lawrence, Kansas. Uhl, N.W., J. Dransfield, J.I. Davis, M.A. Luckow, K.S. Hansen, and J.J. Doyle. 1995. Phylogenetic relationships among palms: cladistic analyses of morphological and chloroplast DNA restriction site variation. Pp. 623–661 in Rudall, P.J, P.J. Cribb, D.F. Cutler, and C.J. Humphries (eds.) Monocotyledons: Systematics and Evolution. Royal Botanic Gardens, Kew.

................. 16157$

CH19

02-08-07 10:33:10

PS

PAGE 370

The Asclepiad Flora of New Guinea . asclepiad flora (Apocynaceae: Asclepiadoideae, Periplocoideae, Secamonoideae) of New Guinea comprises at least 193 species and four subspecies in 18 genera (Forster 1996a, 2000; Forster, Liddle, and Liddle 1997). One genus (Madangia) and 165 species (85%) are endemic. Papua (the western half of the island of New Guinea) is poorly collected, with only 47 species recorded, 12 (25%) being endemic.

T

Number of Genera and Species Worldwide the asclepiads comprise perhaps as many as 250 genera and 2,500 species. In New Guinea there are 18 genera with at least 193 species. Genera that are species rich in New Guinea are Dischidia R.Br. (with 28 species), Hoya R.Br. (with at least 74 species and 4 subspecies) and Marsdenia R.Br. (at least 50 species). There is one endemic genus Madangia P.I.Forst., D.J.Liddle & I.M.Liddle, with a single species M. inflata P.I.Forst. et al. Other nonendemic genera with small numbers of species are Brachystelma Sims (1 species), Ceropegia L. (1), Cynanchum L. (1), Finlaysonia Wallich (1), Gymnanthera R.Br. (1), Heterostemma Wight & Arn. (4), Pentatropis R.Br. (1), Phyllanthera Blume (6), Sarcolobus R.Br. (11), Secamone R.Br. (7), Tylophora R.Br. (7) and Vincetoxicum (1). A number of naturalized taxa also occur (Asclepias L. with 1 species; Calotropis with 1 species).

Distribution and Habitat Asclepiads occur in subtropical and tropical forests and semi-arid deserts throughout the world, with major concentrations of taxa in Africa, Madagascar, Malesia, Asia, South America, and New Guinea. Only a handful of genera have species that occur in temperate regions. Asclepiads are found throughout much of New Guinea, being absent only from alpine areas and anthropogenic grasslands. Some species are widespread (e.g., Dischidia littoralis Schltr., Hoya sussuela (Roxb.) Merr., Marsdenia velutina R.Br.), but many are restricted and appear to be endemic to particular valley systems. There is a wide-ranging group of species that occurs along the seashore in coastal scrub and mangroves. Many of these species may be found outside of New Guinea in adjacent regions of Malesia, the Solomon Islands, and Australia. Some overlap of species occurs between this initial group and those found in adjacent lowland coastal rainforest. A diverse and species-rich assemblage of asclepiad taxa occurs in the lower to midmontane rainforests, with considerable speciation occurring in isolated valley systems. Relatively few species are found in upper montane rainforMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

371

................. 16157$

CH20

02-08-07 10:30:55

PS

PAGE 371

.

372 /

est. It is not uncommon to find considerable species diversity within a small area, especially in rainforest types that are seasonally dry. Careful searching may reveal up to six species of Hoya, three species of Dischidia, several species of Marsdenia, a Tylophora, and occasionally a Heterostemma or Sarcolobus, all in the same patch of forest.

Family Classification The Asclepiadaceae was erected as a distinct family from the Apocynaceae by Robert Brown in 1810. It is now widely accepted, in the light of considerable morphological and molecular analysis, that the Asclepiadaceae sensu stricto is nested within the Apocynaceae sensu stricto; hence only one broadly defined family Apocynaceae is now recognized (Endress 2002; Endress and Bruyns 2002). The asclepiads comprise the three subfamilies Asclepiadoideae, Periplocoideae, and Secamonoideae, of which the Asclepiadoideae is by far the largest in terms of genera and species. The flowers of asclepiads are complex in structure, usually fleshy, and quite unlike those of any other vascular plants (Fishbein 2001). In the Asclepiadoideae the pollen of one theca (with one pollen sac) from two adjacent anthers is aggregated into two masses (pollinia) that are attached to a corpusculum (clip) by translator arms; the whole package is a pollinarium. In the Periplocoideae the pollen of one theca (with two pollen sacs) from two adjacent anthers is shed into a translator scoop. In the Secamonoideae the pollen of one theca (with two pollen sacs) from two adjacent anthers is fused into four pollinia and attached onto the end of a clip translator.

Features of the Asclepiad Subfamilies Habit: annuals or perennials. Herbs, lianas, epiphytes, rarely shrubs; mesophytic, xerophytic (often succulent); usually with latex that is white, more rarely brown, yellow or clear. Stipules: small, often greatly reduced to obsolete. Leaves: opposite (rarely whorled), simple, entire (rarely lobed); with colleters (glands or extrafloral nectaries) often present at the base of the lamina. Infloresences: extra-axillary, cymose, few to much-branched (umbelliform to racemiform), with many flower clusters or reduced to a few flowers. Flowers: five-parted, regular, bisexual. Calyx with colleters at sinus bases. Corolla sympetalous; lobes valvate or contorted in bud, free to variously fused. Annular, corolline and gynostegial coronas present or absent. Stamens five, inserted on corolla tube, almost always at the base, free above or connate into a tubular staminal column (gynostegium) that comprises the five fused stamens and the carpels. Pollen aggregated in tetrads or pollinia, dispersed in translators or pollinaria that comprise two or four pollinia per anther, a corpusculum and caudicles. Ovary superior; carpels two, free except towards base and apex; placentation axile; ovules numerous; styles connate at apex; apex enlarged, forming a secretory style-head that is depressed-globose to elongate-rostrate. Fruits: a pair of follicles (sometimes only one developing) with numerous seeds.

................. 16157$

CH20

02-08-07 10:30:56

PS

PAGE 372

Angiosperms: The Asclepiad Flora of New Guinea / 373

Seeds: Flattened, small ( 1 cm long), brown; usually comose (tuft of hairs) at one end; embryo straight; endosperm present.

Gaps in Floristic Documentation or Knowledge Pioneering collections of the New Guinean asclepiad flora were undertaken by botanist collectors such as O. Beccari, H. O. Forbes, C. L. Ledermann, R. Schlechter, O. Warburg, and C. Weinland. Foremost among these was Schlechter, who named many novelties between 1905 and 1914 (Schlechter 1905, 1913), including the supposedly endemic genus Spathidolepis Schltr., now included in Dischidia as D. torricellensis (Schltr.) P.I.Forst. Virtually no taxonomic work was then undertaken on this group until the late 1980s when revisions of the genera Marsdenia, Phyllanthera (as Cryptolepis), Sarcolobus, and Tylophora were published (Forster 1991, 1993, 1994, 1995). Despite this activity, the asclepiad flora of New Guinea as a whole must be considered poorly known. Most New Guinean asclepiad collections are from Central, Madang, and Morobe provinces in Papua New Guinea. There has been little or no collecting undertaken in many areas, especially Papua where only 47 species have been recorded (Table 3.6.1). Many taxa are known from few or single collections and it is likely that many more similarly documented species remain to be found. Species that are widespread in Papua New Guinea such as Secamone elliptica R.Br., or that are in close proximity to Papua (e.g., Marsdenia bliriensis P.I.Forst., M. mira P.I.Forst., M. primulina P.I. Forst., and Tylophora bilobata P.I.Forst.) are expected to be found there in due course.

Comparison with Adjacent Regions Asclepiads are widespread in the regions adjacent to New Guinea (Australia, Solomon Islands, Malesia, western Pacific), but there is little overlap in species (Table 3.6.2). At the species level, the strongest links are to the flora in Australia that has 23 genera with 89 native species and 11 naturalized taxa (Forster 1996b). Although the asclepiad flora of Australia is less species rich, it does have two endemic genera (Gunnessia P.I.Forst., Rhyncharrhena F.Muell.) as opposed to a single one (Madangia) in New Guinea. At the generic level, all nonendemic genera of asclepiads found in New Guinea are found in adjacent regions (Table 3.6.3).

Natural History The usual habit of an asclepiad plant is as a herbaceous twiner or woody liana, with a root system anchored in soil. In New Guinea, as in adjacent Malesia and parts of Asia, many asclepiads (particularly Marsdenieae) are epiphytic on rainforest trees, often never having contact with the soil. These epiphytes (usually species of Dischidia and Hoya) may have ascending or decumbent stems, sometimes forming large pendent masses. While some epiphytic species are robust, and differ little

................. 16157$

CH20

02-08-07 10:30:56

PS

PAGE 373

.

374 /

Table 3.6.1. Distribution of asclepiads recorded from Papua Species

Distribution outside Papua

Dischidia hirsuta

Papua New Guinea, Solomon Islands

Dischidia imbricata

Papua New Guinea, Solomon Islands

Dischidia littoralis

Australia, Papua New Guinea

Dischidia longifolia

?endemic

Dischidia major

Australia, Papua New Guinea

Dischidia soronensis

?endemic

Dischidia torricellensis

Papua New Guinea

Finlaysonia obovata

Australia, Papua New Guinea

Gymnanthera oblonga

Australia, Papua New Guinea

Heterostemma acuminatum

Australia, Papua New Guinea, Solomon Islands

Hoya anulata

Australia, Papua New Guinea

Hoya australis subsp. tenuipes

Australia, Papua New Guinea

Hoya calycina subsp. glabrifolia

Papua New Guinea

Hoya eitapensis

Papua New Guinea

Hoya globulifera

?endemic

Hoya halophila

Papua New Guinea

Hoya microphylla

Papua New Guinea

Hoya papuana

Papua New Guinea

Hoya piestolepis

Papua New Guinea

Hoya pottsii (nicholsoniae)

Australia, Papua New Guinea

Hoya pruinosa

?endemic

Hoya purpurea

?endemic

Hoya revoluta (litoralis)

Australia, Papua New Guinea, Malesia

Hoya rhodostemma

Papua New Guinea

Hoya solaniflora

Papua New Guinea

Hoya sussuela

Australia, Papua New Guinea, Malesia

Marsdenia arfakensis

endemic

Marsdenia argillicola

endemic

Marsdenia belensis

endemic

Marsdenia dischidioides

endemic

Marsdenia kebarensis

endemic

................. 16157$

CH20

02-08-07 10:30:57

PS

PAGE 374

Angiosperms: The Asclepiad Flora of New Guinea / 375 Marsdenia papillosa

Papua New Guinea

Marsdenia parva

endemic

Marsdenia praestans

Australia, Malesia

Marsdenia velutina

Australia, Papua New Guinea, Solomon Islands

Pentatropis carnosum

Australia, Papua New Guinea

Phyllanthera lancifolia

Papua New Guinea

Phyllanthera papillata

Papua New Guinea

Phyllanthera perakensis

Papua New Guinea, Malesia

Sarcolobus porcatus

Papua New Guinea

Sarcolobus retusus

Papua New Guinea

Sarcolobus secamonoides

Papua New Guinea

Sarcolobus vittatus

Australia, Papua New Guinea

Tylophora cissoides

Papua New Guinea

Tylophora flexuosa

Australia, Papua New Guinea, Malesia

Tylophora minima

endemic

Sources: Forster 1996a, 2000, unpublished.

in habit from most climbers, there is a range of (small growing species) that are restricted to thin twigs, especially in the genus Hoya (Forster, Liddle, and Liddle 1998). The asclepiads are unique among the dicotyledons in the packaging of their pollen grains into structures called pollinia (collectively a pollinarium) and translators (Kunze 1991), the only other plant family with pollinia being the Orchidaceae. These translators (subfamily Periplocoideae) and pollinaria (subfamilies Asclepiadoideae, Secamonoideae) require movement by animals for pollination. For fertilization to occur, the pollinarium or translator must be removed from a flower, and then reinserted in a particular way such that the individual pollen grains are able to germinate and effect fertilization. There is no pollen to stylehead transfer in the two subfamilies Asclepiadoideae and Secamonoideae, and the pollinia have to be inserted through slits (alar fissures), even though a style-head is still present. Asclepiads are usually pollinated by small insects (Coleoptera, Diptera, Lepidoptera), especially those species with small bowl-shaped (campanulate, rotate, salverform) flowers, although it is possible that some birds (honeyeaters) may pollinate some of the larger, flat-flowered (campanulate or rotate) species of Hoya that produce considerable nectar (Forster 1992; Ollerton and Liede 1997). Closed flowers of the South African asclepiad Microloma, that are similar in external appearance to many Dischidia flowers, are pollinated by sunbirds (Ollerton 1998), so this possibility should not be discounted. At present the pollination ecology of all New Guinean species of asclepiads is unknown. The seeds of all New Guinean asclepiads, with the exception of Finlaysonia

................. 16157$

CH20

02-08-07 10:30:57

PS

PAGE 375

.

376 /

Table 3.6.2. Distribution of non-endemic indigenous New Guinea asclepiad species Species

Distribution

Brachystelma glabriflorum

Australia

Ceropegia cumingiana

Australia, Malesia

Cynanchum ovalifolium

Australia, Malesia

Dischidia hirsuta

Malesia, Solomon Islands

Dischidia imbricata

Malesia

Dischidia littoralis

Australia

Dischidia major

Australia, Malesia

Dischidia nummularia

Australia

Dischidia ovata

Australia

Finlaysonia obovata

Australia, Malesia

Gymnanthera oblonga

Australia

Heterostemma acuminatum

Australia, Malesia

Hoya australis subsp. tenuipes

Australia

Hoya revoluta

Australia, Malesia

Hoya samoensis

Solomon Islands, SW Pacific

Marsdenia geminata

Australia

Marsdenia praestans

Malesia

Marsdenia suborbicularis

Australia

Marsdenia tricholepis

Australia

Marsdenia velutina

Australia, Malesia

Marsdenia viridiflora

Australia

Pentatropis carnosum

Australia, Malesia

Secamone elliptica

Australia, Malesia, Melanesia, Asia

Secamone timoriensis

Australia, Malesia

Tylophora cissoides

Malesia

Tylophora flexuosa

Australia

Vincetoxium brachystelmoides

Australia

Note: ‘‘Malesia’’ is exclusive of New Guinea in this table.

................. 16157$

CH20

02-08-07 10:30:57

PS

PAGE 376

Angiosperms: The Asclepiad Flora of New Guinea / 377

Table 3.6.3. Distribution of non-endemic indigenous New Guinea asclepiad genera Genus

Region

Brachystelma

Australia, Malesia

Ceropegia

Australia, Malesia

Cynanchum

Australia, Malesia

Dischidia

Australia, Malesia, SW Pacific

Finlaysonia

Australia, Malesia

Gymnanthera

Australia, Malesia

Heterostemma

Australia, Malesia, SW Pacific

Hoya

Australia, Malesia, SW Pacific

Marsdenia

Australia, Malesia, SW Pacific

Pentatropis

Australia, Malesia

Phyllanthera

Australia, Malesia

Sarcolobus

Australia, Malesia, SW Pacific

Secamone

Australia, Malesia, SW Pacific

Tylophora

Australia, Malesia, SW Pacific

Vincetoxicum

Australia, Malesia

obovata and some Sarcolobus species, possess a coma of hairs at the micropylar end. This coma assists the seed in wind dispersal. Given the epiphytic habit of many of the New Guinea species, this dispersal mechanism is fundamental to establishment of new populations of species. The dispersal ecology of all New Guinean species of asclepiads is unknown. Lepidoptera, especially Nymphalid butterflies, are dependent on the foliage of asclepiads for larval development. Little is known about the host plant preferences of these butterflies on the New Guinean asclepiad flora. A small number of asclepiads (e.g., Dischidia major Vahl) have a non-obligate relationship with ants where inflated ‘‘bladder’’ leaves may host vigorous colonies (Kleijn and van Donkelaar 2001). If the plant is knocked or predated, these ants will defend against the intruders. Ants are also known to disperse the seeds of certain asclepiads (Dischidia, Hoya).

Acknowledgment Thanks to Mary Endress for reviewing the manuscript.

Literature Cited Endress, M.E., and P.V. Bruyns. 2001. A revised classification of the Apocynaceae s.l. The Botanical Review 66: 1–56.

................. 16157$

CH20

02-08-07 10:30:58

PS

PAGE 377

.

378 /

Endress, M.E. 2002. Apocynaceae and Asclepiadaceae: united they stand. Haseltonia 8: 2–9. Fishbein, M. 2001. Evolutionary innovation and diversification in the flowers of Asclepiadaceae. Annals of the Missouri Botanical Garden 88: 603–623. Forster, P.I. 1991. A taxonomic revision of Sarcolobus R.Br. (Asclepiadaceae: Marsdenieae) in Australia and Papuasia. Austrobaileya 3: 443–466. Forster, P.I. 1992. Pollination of Hoya australis (Asclepiadaceae) by Ocybadistes walkeri sothis (Lepidoptera: Hesperiidae). Australian Entomological Magazine 19: 39–43. Forster, P.I. 1993. Conspectus of Cryptolepis R.Br. (Asclepiadaceae: Periplocoideae) in Malesia. Austrobaileya 4: 67–73. Forster, P.I. 1994. A taxonomic revision of Tylophora (Asclepiadaceae: Marsdenieae) in Papuasia. Australian Systematic Botany 7: 485–505. Forster, P.I. 1995. Circumscription of Marsdenia (Asclepiadaceae: Marsdenieae) with a revision of the genus in Australia and Papuasia. Australian Systematic Botany 8: 703–933. Forster, P.I. 1996a. A checklist of the Asclepiadaceae of Papuasia. Science in New Guinea 22: 15–22. Forster, P.I. 1996b. Asclepiadaceae. Flora of Australia 28: 197–283. Hoya, Dischidia by Forster, P.I., D.J. Liddle. CSIRO Australia, Melbourne. Forster, P.I. 2000. Cryptolepis papillata P.I.Forst. and Sarcolobus porcatus P.I.Forst. (Asclepiadaceae), newly recorded from West Papua. Austrobaileya 5: 729. Forster, P.I., D.J. Liddle, and I.M. Liddle. 1997. Madangia inflata (Asclepiadaceae: Marsdenieae), a new genus and species from Papua New Guinea. Austrobaileya 5: 53–57. Forster, P.I., D.J. Liddle, and I.M. Liddle. 1998. Diversity in the genus Hoya (Asclepiadaceae: Marsdenieae). Aloe 35: 44–48. Kleijn, D., and R. van Donkelaar. 2001. Notes on the taxonomy and ecology of the genus Hoya (Asclepiadaceae) in central Sulawesi. Blumea 46: 457–483. Kunze, H. 1991. Structure and function in asclepiad pollination. Plant Systematics and Evolution 176: 227–253. Ollerton, J. 1998. Sunbird surprise for syndromes. Nature 394: 726–727. Ollerton, J., and S. Liede. 1997. Pollination systems in the Asclepiadaceae: a survey and preliminary analysis. Bio. Jour. Linn. Soc. 62: 593–610. Schlechter, R. 1905. Periplocaceae, Asclepiadaceae. Pp. 351–369 in Schumann, K., and K. Lauterbach (eds.) Nachtra¨ge zur Flora der Deutschen Schutzgebiete in der Su¨dsee. Gebru¨der Borntra¨ger, Leipzig. Schlechter, R. 1913. Die Asclepiadaceen von Deutsch-Neu-Guinea. Botanische Jahrbu¨cher fu¨r Systematik, Pflanzengeschichte und Pflanzengeographie 50: 81–164.

................. 16157$

CH20

02-08-07 10:30:58

PS

PAGE 378

Costaceae of Papua . Number of Genera and Species

T

seven genera and at least 170 species in the family. The genera Cheilocostus and Tapeinochilos occur naturally in New Guinea.

Distribution and Habitat Costus has some 135 species in the tropics of Africa and America. Six or seven species of tropical Asia that used to be in Costus are now in Cheilocostus; two of these are found in New Guinea (Maas 1979). All eight species of Chamaecostus, both species of Dimerocostus, and the monotypic Monocostus uniflorus (Petersen) Maas occur only in tropical America, while the 15–20 species of Tapeinochilos are restricted to the Old World, being found in New Guinea and nearby parts of Indonesia and Australia. Finally, Paracostus consists of one species in Cameroon and another in Borneo.

Family Classification The most recent classification of the genera of Costaceae places all but one of the Asian species which were formerly in Costus into a new genus, Cheilocostus (Specht and Stevenson 2006). A further two genera have also been raised out of Costus, namely Chamaecostus and Paracostus. Dimerocostus, Monocostus and Tapeinochilos remain unchanged. In the older literature (e.g., Mabberley 1997) the Costaceae were placed in the Zingiberaceae subfamily Costoideae, but the evidence for recognizing them at family rank is now overwhelming (Kress et al. 2001; Larsen 1998). It is expected that the next edition of Mabberley’s Plant Book will reflect this.

Features of the Family Habit: herbs, sometimes very robust, not containing essential oils, often branching in Tapeinochilos. Leaves composed of clasping sheaths, ligule and blade, the tubular and closed sheaths forming pseudostems; ptyxis supervolute, venation pinnate. Inflorescences cone-like spikes, terminal on leafy shoots or on separate, leafless shoots, sometimes in both positions on an individual, bracts subtending one to two flowers (flowers solitary in axils of upper leaves in Monocostus). Flowers zygomorphic, bisexual, calyx fused with two to three teeth; corolla fused with three lobes; labellum lobed, margin often crisped, usually the most conspicuous organ of the flower, flanked by lateral teeth in Tapeinochilos; one fertile stamen Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

379

................. 16157$

CH21

02-08-07 10:31:07

PS

PAGE 379

.

380 /

opposite the labellum, anther with two thecae; gynoecium syncarpous, the ovary inferior with two to three locules, style passing between the anther thecae but not fused to them, stigma punctate; septal nectaries two, immersed in the apical part of the ovary; placentation axile, ovules few to many per locule, arillate, anatropous. Fruits a 3 (–2) locular capsule, dehiscing loculicidally. Seeds numerous, aril white to yellow.

Gaps in Floristic Documentation or Knowledge See the chapter on Zingiberaceae (Chapter 3.6). Dr Osia Gideon (University of Papua New Guinea) is working on a revision of Tapeinochilos (Gideon 1996). A thorough revision of Cheilocostus is required because, as noted by Maas (1979), the species limits in the C. globosus complex have yet to be resolved.

Literature Cited Gideon, O. 1996. Systematics and evolution of the genus Tapeinochilos Miq. (Costaceae, Zingiberaceae). Ph.D. thesis, James Cook University, Queensland. Kress, W.J., L.M. Prince, W.J. Hahn, and E.A. Zimmer. 2001. Unraveling the evolutionary radiation of the families of the Zingiberales using morphological and molecular evidence. Syst. Biol. 50: 926–944. Larsen, K. 1998. Costaceae. In Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. Springer-Verlag, Berlin. Maas, P.J.M. 1979. Notes on Asiatic and Australian Costoideae (Zingiberaceae). Blumea 25: 543–549. Mabberley, D.J. 1997. The Plant Book. 2nd ed. Cambridge University Press, Cambridge. Specht, C.D., and D.W. Stevenson. 2006. A new phylogeny-based generic classification of Costaceae (Zingiberales). Taxon 55 (1): 153–163.

................. 16157$

CH21

02-08-07 10:31:07

PS

PAGE 380

Elaeocarpaceae of Papua . . currently comprises 12 genera (Kubitzki 2004), including three previously placed in Tremandraceae. Four genera are endemic to Australia. The family is of southern (Gondwanan) origin and molecular evidence places it next to Cunoniaceae in Oxalidales. The family is represented in Papua by species of five genera: Aceratium, Dubouzetia, Elaeocarpus, Sericolea, and Sloanea (Coode 1978, 1981). In Papua, the family is represented by large trees, often buttressed or stiltrooted, through small trees to montane shrubs. Many species of all these genera seem to occur in disturbed or successional areas, although some are restricted to intact forest or other primary vegetation. A feature of the family is that petals are usually lobed or laciniate at the tip; this may attract insect pollinators, as may the nectariferous disk, but little is known for certain. Buzz-pollination has been suggested (Matthews et al. 2002) but this does not appear to be the whole story, because flowers of many collections have abundant nectar.

E

Aceratium There are about 17 species of Aceratium altogether (ca 12 in New Guinea of which ten are known from Papua, plus five endemic in Queensland which are distinct from the rest). Few are easy to identify to species. One, A. oppositifolium, is widespread throughout Papuasia, extending west to Buru in Maluku and east to the Solomon Islands. This species is very common and has been reported to have edible fruit; it is usually found in disturbed areas or forest margins. The other species are less common and little is known of their ecology; some are recorded as forest species. Most are small trees at low to middle altitudes, but some (e.g., A. sphaerocarpum of southern Papua) can reach 27 m. Petals in Papuan Aceratium are obviously laciniate at apex and the fruits are usually red and drupaceous with edible flesh surrounding and attached to a horny or fibrous core with one or more dry seeds. This fruit morphology indicates possible dispersal by bats. Recent as yet unpublished molecular work places Aceratium with Sericolea and both with Elaeocarpus, perhaps even embedded within it (D. Crayn, pers. comm.).

Dubouzetia Dubouzetia comprises approximately 12 species, with three or four species in New Guinea, three in northern Australia, and seven in New Caledonia. All have dehiscent, dry-walled fruits. One species, D. dentatus from Papua, may extend west Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

381

................. 16157$

CH22

02-08-07 10:31:13

PS

PAGE 381

.

382 /

.

into (low altitude) Maluku but is not well known. It may not be distinct from D. elegans, which is found throughout New Guinea in forests between 2,200 and 2,700 m and also in New Caledonia. This distribution is strange, in that it has not yet been found in the intervening Bismarck Archipelago and Solomon Islands. (The Australian and other New Caledonian species are endemic and quite distinct.) Dubouzetia galorei is perhaps the most interesting. In the south of the Vogelkop, Papua (as in the Aru Islands and along the Fly River in southern Papua New Guinea) it seems to be a species of low altitude or flood plain forest. Its pink seeds are composed of a thick spongy layer surrounding a minute ‘‘stone’’ and these seeds are said to float buoyantly; they are also (in Aru at least) eaten by cassowaries to such excess that the birds become easy to catch (field notes on van Balgooy’s collection numbered 6,596). In Papua New Guinea, however, what appears to be the same species also grows on forested slopes at altitudes up to 1,200 m. All other species of Dubouzetia have dry seeds, with or without a waxy outgrowth at one end. Petals are notched or lobed rather than laciniate. The genus seems to be most closely related to Peripentadenia (two spp. in Queensland) and Crinodendron (five spp. in South America).

Elaeocarpus Currently Elaeocarpus, the largest genus in the family, is thought to comprise about 400 species in total. Its distribution extends from Madagascar and Mauritius in the west, through Sri Lanka and southern and eastern India, Burma, Southeast Asia to Japan, throughout Malesia, Australia, New Caledonia, New Zealand, and Fiji. A few species are found in the western Pacific Islands and one is endemic to Hawaii. The patterns of variation within the genus are interesting: there are more species in Borneo (ca 70) than there are in Sulawesi (ca 30), but those of Sulawesi belong to more groups than those of Borneo (Coode 1995). New Guinea is even richer in numbers of groups (though lacking two of Borneo’s six groups, New Guinea has at least four other groups not represented in Borneo) and has more species than either (ca 80). Endemism is high throughout the range: of the 80 species found in Papuasia, only about ten extend beyond the boundaries of the region (ca 15%) and the percentage endemism in the other major islands of Malesia is only somewhat less. Elaeocarpus fruits are drupes: a single woody stone is enclosed in an often fleshy mesocarp, surrounded by epicarp, frequently blue. The blue coloration is due not to dyes but to light refraction (Lee 1991). The fruits are attractive to frugivorous birds such as cassowaries, pigeons, and hornbills, discussed for New Guinea by Frith et al. (1976) and for Queensland by Crome (1975a,b). In most species, the mesocarp eventually separates from the stone which (if large enough) can be found on the forest floor. In 1978 and 1981, 68 species were recognized for Papuasia, of which 43 were known from Papua. Since then, about nine previously unrecognized species have

................. 16157$

CH22

02-08-07 10:31:14

PS

PAGE 382

Angiosperms: Elaeocarpaceae of Papua / 383

been added, of which six are from Papua, and two added to Papua’s list that were previously known from Papua New Guinea only. In 1978, the species were placed in nine informal groups, some of which were further subdivided. These groups have been altered slightly since but still largely reflect the relationships suggested by gross morphology. Molecular studies at the species level are underway in Australia, but the results have not yet been published. Weibel (1968) reported that the seeds and their embryos can be straight (like most of the rest of the family) or curved or folded. Groups 1–6 have straight seeds in fruits from very small to very large; Groups 7 and 8 curved, in small fruits (in Borneo they may be medium-sized). Fruit was unknown in Group 9. All groups are represented in Papua. Other characters important in defining the groups are number of ovules per loculus and number of loculi per ovary; it is unfortunate that these are rather difficult to see. Many species of Elaeocarpus are found close to rivers, though in Papua none seems as closely tied to riparian habitats as are some of the Bornean species (e.g., E. macrocerus). The endemic Group 1 comprises E. bilobatus, rarely collected. It is a small to medium tree or shrub, once reported to be epiphytic. The flowers are small, ca 3 mm long; the fruits are less than 1 cm. Group 2 (Dactylosphaera) comprises species with small flowers and, uniquely, the petal laciniae are rounded to slightly swollen at tip and hooked inwards. Presumably, this has something to do with specialized pollinators. The medium-sized fruits are longer than broad and have thick-walled generally fusiform stones, suggesting also some specialization in fruit distribution. Species delimitation within this group is very difficult, suggesting recent evolution, yet the group is not endemic to New Guinea but has also been found in Halmahera and Sulawesi, but with less variation. Group 2 is not known from Australia. Group 3 (Elaeocarpus) is much more speciose in west Malesia than in New Guinea. In 1978, I unconvincingly subdivided it into small- and larger-flowered subgroups. There are three ‘‘small-flowered’’ species; their medium-sized fruits have stones that are ellipsoid but woody as in Group 2. One, E. homalioides, endemic to the New Guinea mainland, resembles Group 2 in all but swollen petal divisions and stone shape, and is also very similar to E. teysmannii from Sulawesi and Halmahera of this group. The other, E. multisectus, is widespread and variable in Papua, and extends eastwards into the Solomon Islands; it has been tried as a fast-growing plantation crop in the Markham Valley, Papua New Guinea. E. royenii has been collected only twice, both in Papua. The five species placed in the larger-flowered subgroup all have massive fruits and stones. Were this a feature unique to New Guinea, it might be thought a specialization for cassowary distribution, but very similar fruits are known in a few other species of this very large group in, for example, the Malay Peninsula and Borneo, where cassowaries are unknown. Group 3 has not been found in Australia. Group 4, with a single species, E. blepharoceras, may best be moved to Group 6. It has large fruit in which the fleshy mesocarp is replaced by dense and apparently

................. 16157$

CH22

02-08-07 10:31:14

PS

PAGE 383

.

384 /

.

persistently attached radiating fibers. I can suggest no selective advantage to this, unless it deludes cassowaries into thinking it is good to eat by also being blue, and fibers may be less costly to produce than edible flesh. Recent work in Australia suggests that this species may be close to a new species from northern New South Wales (D. Crayn, pers. comm.)—a curiously disjunct distribution. In 1978, Group 5 comprised five subgroups. Recently, I have decided to merge the first two and move the last to Group 9. The first subgroup (Ganitrus) is the most widespread of all: one variable species previously known as Elaeocarpus sphaericus (or, earlier, Elaeocarpus ganitrus) is found from Nepal to Taiwan, throughout Malesia to Australia and the Solomons and New Caledonia. Now known as E. angustifolius, this provides most of the beads known as rudraksha from which rosaries or malas are made in India, Nepal, and Malesia; the fruit stones are round and beautifully sculptured and usually have five equidistant lines denoting the number of loculi inside. There is great interest in the size of the stones and number of lines, the ‘‘best’’ commanding high prices. Apart from E. angustifolius, the most variation in the group is in New Guinea, where there are several species (including the montane E. ptilanthus with very deeply sculptured stones), and also in the Philippines where perhaps two further species can be recognized. Only E. angustifolius reaches Australia. The second subgroup links the previous with the following and comprises two species, one rare, the other common in parts of Papua New Guinea but not yet found in Papua. It has massive fruits attractive to cassowaries, and piles of stones can be found on the forest floor. The Fissipetalum subgroup currently has unsatisfactory species limits. It has a core of variants very similar in flower and nearly all with small fruits with densely woody stones. These are mostly montane trees or subalpine shrubs and it is the only group in which the variation can be seen to be greater in Papua than in Papua New Guinea, despite the relative sparsity of collections from Papua. To this core of similar taxa are added E. arnhemicus, a lowland species with somewhat different petals and small fruits with rather thin-walled stones. This species is most common in savanna or monsoon forest areas and found in northern Australia, Timor, Flores, and as far west as west Java, as well as throughout New Guinea; there are a few collections from New Britain. Finally, a herbarium specimen of the rare lowland E. sericoloides could be confused with E. arnhemicus in flower but the fruit is much larger and has the most deeply sculptured stone of all; trees can be found on the lawn of the headquarters of PT Freeport Indonesia, Kuala Kencana, Timika. The first subgroup of Group 6 (Monocera) comprises E. polyandrus; it is related to species of the western Pacific and is not yet known in Papua. The name reflects the number of stamens (up to 150). In New Guinea, the second subgroup of Group 6 is represented by E. womersleyi. This species seems to be related to E. bancroftii and E. stellaris of Queensland; they have large flowers with big 3-lobed petals and large fruits with massive woody stones. E. womersleyi resembles E. blepharoceras (above) in having persistent radi-

................. 16157$

CH22

02-08-07 10:31:14

PS

PAGE 384

Angiosperms: Elaeocarpaceae of Papua / 385

ating fibers on the stone at least at first: one can find stones on the forest floor in which the fibers have eroded away. Perhaps these too deceive the none-too-bright cassowary. The species extends westwards as least to Morotai, and is variable in Papua; it can be found from near sea level to 2,000 m and is usually found in rainforest. Further new species of this subgroup, from Sulawesi (herbarium specimens) and Queensland (D. Crayn, pers. comm.) bring the total to five. The third subgroup of Group 6 comprises seven species of which one, E. nouhuysii, is often collected in Papua. It can be a large tree and has large flowers and medium-large, flattened fruit; it often grows in seasonally flooded forest, sometimes behind mangroves or in advanced secondary regrowth. The species is also known from a single collection in Seram. The subgroup has other forest species with flattened fruit. Also, the spectacular large-flowered E. schlechterianus, with massive rounded fruit, is known in Papua from the Idenburg River at least. The subgroup extends east into the Solomons; it does not reach Australia. The fourth subgroup is probably not natural; the flowers are smaller than in the previous subgroup and not much is known about the Papuan representatives. The fifth, however, has grown recently with the addition of two interesting new species from Papua: E. crassus, known from two collections east of Sorong in the Warsamson Plain, with massively thick twigs and bunches of almost sessile leaves, and E. timikensis from near Timika, with similar thick twigs but these are full of black ants and the leaves have distinct petioles. A third distinctive species previously described from Papua, Idenburg River, is E. debruynii, with large many-nerved leaves on long stiff petioles. It is not yet clear whether this group is at all natural and related to species elsewhere in Malesia. Group 7 (Oreocarpus), with curved seeds and small fruits, comprises the widespread and variable E. culminicola, shrubs to small or medium-sized trees often in disturbed habitats from low to middle altitudes. The interest of this species is in its distribution: from Luzon, Mindoro, and Mindanao to Sulawesi, Maluku, throughout New Guinea to New Ireland and Queensland. Such a pattern is known in other groups (Baker et al. 1998). There is a rare second species with larger flowers, E. sterrophyllus, from the Cyclops Mountains near Jayapura. The group extends into Australia, where there are at least six further distinctive species. Group 8 (Coilopetalum), also with curved seeds and small fruits, has also been subdivided, albeit rather unsatisfactorily. The first two subgroups comprise two species each, all endemic to New Guinea and known from Papua, from midmontane to subalpine vegetation, often in disturbed habitats. The third subgroup, also endemic, comprises four related species from middle altitudes, only two of which are known from Papua, and the distinctive E. habbemensis, described from Papua but more often collected in Papua New Guinea. The fourth subgroup of Group 8 contains the core of the group, with thick ovate usually hairy petals and usually without distinctive apical divisions. These are mostly lowland species (E. sarcanthus is from middle altitudes) and many are successional. E. sepikanus is more variable in Papua than in Papua New Guinea, but E. elatus is confined to eastern New Guinea and New Britain. Two peripheral

................. 16157$

CH22

02-08-07 10:31:15

PS

PAGE 385

.

386 /

.

species have interesting relationships: E. branderhorstii seems closest to E. lancipetalus of Maluku. E. floridanus is a member of a group of ca six species otherwise endemic to the Lesser Sunda Islands, including Timor; E. floridanus extends far to the east through the Solomon Islands into the Pacific where it is generally known under different names (e.g., E. tonganus). In Papua, E. floridanus has so far been found in Biak only, while in Papua New Guinea it is restricted to the Bismarck Archipelago (Emira, off New Britain) and the East Papuan Islands (Rossel, Woodlark). It would be interesting if any other group of organisms displays this pattern of distribution. The subgroup as a whole extends west and north into the rest of Malesia with many species particularly in Borneo, and possibly southwards into Australia, where E. ruminatus may be related. Apart from E. floridanus, it does not extend eastwards. Group 9 was known from a single species E. schoddei from Papua New Guinea, inadequately known. Recent work suggests that the shrubby E. amabilis, described from the Arfak Mountains and collected only twice, in ‘‘fire-induced shrubbery,’’ may also belong here.

Sericolea Sericolea is a genus of small trees and shrubs of middle to high altitudes of mainland New Guinea; there is a single specimen from Fergusson Island. The fruit can be loosely termed a ‘‘berry’’; a soft watery-fleshy mesocarp (presumably of interest to birds) surrounding one or more hard ‘‘seeds.’’ Those seeds studied have slightly curved embryos. It is a very difficult genus: the revision by van Balgooy (1982) lists 15 named species. Very few of them are both distinctive and well delimited; many mountains or mountain ranges seem to have their own variants yet the same mountain may have up to four different species, sometimes occupying different niches (van Balgooy 1982). This suggests recent evolution and it appears most closely related to Elaeocarpus and Aceratium.

Sloanea Sloanea is the second-largest genus in the family, and is the most widespread, being known from Central and tropical South America as well as Madagascar, northeastern Indian subcontinent, through Burma to southern China, throughout Malesia to Australia and New Caledonia. There are about 150 species, of which ca 100 are in the New World. One of the four Australian species seems to be related to one from New Guinea, but the rest do not. The fruit is a massive woody capsule; a few non-Papuasian species have small fruits, but all dehisce. Seeds may be arillate or with a sarcotesta (Coode 1983). Some species have smooth capsules, others spiny. A few have a complex coating of detachable irritant bristle-spines; these appear to have arisen independently at least once in the New World, and perhaps three times in the Old, since their presence or absence is not correlated with any other character. What selective

................. 16157$

CH22

02-08-07 10:31:15

PS

PAGE 386

Angiosperms: Elaeocarpaceae of Papua / 387

advantage these spines may confer is conjectural; elsewhere, they may deter monkeys, but in primate-less New Guinea and Queensland forests, perhaps possums and tree kangaroos are deterred. Petals may be broad with a toothed rim, sometimes fused laterally even into a complete corona, or sepaloid (thick, ovate and acute without teeth at apex), or absent completely (Australia, New World). There are about 18 species in Papuasia in three main groups, all represented in Papua. Group 1 (Anoniodes) contains nine species that have sepaloid petals and includes those (e.g., S. sogerensis and S. pullei) which have pinnate juvenile leaves, unique in the family. Seeds are completely covered with sarcotesta and fruits nearly always have coarse spines. Only three of these species are known from Papua, but S. pullei seems restricted to it. A specimen of S. sogerensis has been collected in Maluku, and others in the Bismarck Archipelago. Otherwise the group is endemic to New Guinea and it is not clear to which other species of Sloanea it is related. Group 2 (Antholoma) contains eight species with petaloid petals, free or variously united laterally, or (S. tieghemii) fused into a toothed corona. Seeds are partly covered with arils and fruits are smooth. Two widespread species are known from Papua, S. aberrans at low to middle altitudes with free petals, and the very variable S. tieghemii mostly with coronas and mostly at middle altitudes or higher. Both may occur in primary or secondary vegetation. The group has relatives in Sulawesi and extends eastwards (via S. insularis) into the Solomons and beyond to New Caledonia (nine species). Group 3 (Cnidocarpaea) is represented by the single widespread species S. paradisiarum, which has the fruit covered with short bristle-spines and sepaloid petals. The Australian S. langii is probably related; there are bristle-spiny species on the Asiatic mainland and another in Australia, but their flowers are different. Sloanea is rather isolated in the family; it may be closest to Vallea and Aristotelia.

Present Status of Documentation It is difficult to pinpoint areas of particular richness in Papua because very few have been adequately covered by collectors and fieldworkers, and the revisionary work is not yet complete. It seems clear that the most species of Elaeocarpaceae in New Guinea occur at middle altitudes, but that some species can be at sea level and others at 3,000 or more meters. Of the better-collected areas in Papua, the Arfak Mountains would seem to be rich, with some 14 species so far recorded. About 12 were noted for the lowlands around Ayawasi (southern Vogelkop; Polak 2000), but these identifications were preliminary. Classic rich collections of Elaeocarpaceae were made by Brass from the Idenburg River and Lake Habbema; it is difficult to know whether these localities were any richer than any other as yet unvisited. There are interesting collections from the Cyclops Mountains (Jayapura) and from Yapen Island. Recent fieldwork in the Timika area revealed new species and records for Papua. Some species were found at much lower altitudes

................. 16157$

CH22

02-08-07 10:31:16

PS

PAGE 387

.

388 /

.

than expected (cold air pours down from the glaciers above) but, while many of the expected species were found, some were not, suggesting that sampling is still too patchy for anything meaningful to be said. Even so, about 23 species were found in a narrow strip from sea level to 3,000 m, but whole belts of midmontane forest were not sampled in the time available, access being difficult. Note: A recent report on the start of molecular work on the family (Maynard 2006) is most welcome.

Literature Cited Baker, W.J., M.J.E. Coode, J. Dransfield, S. Dransfield, M.M. Harley, P. Hoffmann, and R.J. Johns. 1998. Patterns of distribution of Malesian plants. Pp. 243–258 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys, Leiden. Coode, M.J.E. 1978. A conspectus of Elaeocarpaceae in Papuasia. Brunonia 1 (2): 131–302. Coode, M.J.E. 1981. Elaeocarpaceae. Pp. 38–185 in Henty, E.E. (ed.) Handbooks of the Flora of Papua New Guinea, Vol. II. Melbourne University Press, Melbourne. Coode, M.J.E. 1983. A conspectus of Sloanea (Elaeocarpaceae) in the Old World. Kew Bull. 38: 347–427. Coode, M.J.E. 1995. Elaeocarpus in the Flora Malesiana area—E. kraengensis and ten new species from Sulawesi. Kew Bull. 50: 267–294. Crome, F.H.J. 1975a. Some observations on the biology of the cassowary in northern Queensland. Emu 76: 8–14. Crome, F.H.J. 1975b. The ecology of fruit pigeons in tropical northern Queensland. Austr. Wildlife Research 2: 155–185. Frith, H.J., F.H.J. Crome, and T.O. Wolfe. 1976. Food of fruit pigeons in New Guinea. Emu 76: 49–58. Kubitzki, K. 2004. The Families and Genera of Vascular Plants VI. Springer, Berlin. Lee, D.W. 1991. Ultrastructural basis and function of iridescent blue colour of fruits in Elaeocarpus. Nature 349: 260–263. Matthews, M.L., and P.K. Endress. 2002. Comparative floral structure and systematics in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Bot. J. Linn. Soc. 140: 321–381. Maynard, D.J. 2006. A molecular phylogeny for Australian Elaeocarpus (Elaeocarpaceae) and the affinities of a putative new taxon. Australian Systematic Botany Society Newsletter 126: 17–19. Polak, M. 2000. The botanical diversity in the Ayawasi area, Irian Jaya, Indonesia. Biodiversity and Conservation 9: 1345–1375. van Balgooy, M.M.J. 1982. A revision of Sericolea Schlechter (Elaeocarpaceae). Blumea 28: 103–141.

................. 16157$

CH22

02-08-07 10:31:16

PS

PAGE 388

Ericaceae of Papua lyn a. craven Number of Genera and Species h e er i c a ce a e , as recently redefined to include Epacridaceae, comprises about 4,000 species in approximately 120 genera. The family is relatively well collected in New Guinea and taxonomically is quite well known at the species level due largely to the revisionary studies of H. Sleumer for Flora Malesiana (Sleumer 1964, 1966) that have been complemented by van Royen (1982) and van Royen and Kores (1982). The genera occurring in New Guinea are: Agapetes (ca 95 species in genus/ca 10 species in New Guinea), Decatoca (1/1), Dimorphanthera (ca 75/ca 60), Diplycosia (ca 100/20), Gaultheria (ca 130/6), Rhododendron (ca 1,000/ ca 150), Styphelia sensu lato (ca 300/4), Trochocarpa (ca 12/8), Vaccinium (ca 450/ ca 130).

T

Distribution and Habitat Ericaceae is a cosmopolitan and speciose family. The species predominantly occur in temperate to subtemperate zones and, where they do occur in the tropics, the species occur mainly in montane to subalpine regions. Within New Guinea, species may occur as epiphytic shrubs in montane rainforest, terrestrial shrubs in open ridge communities and land-slip areas, in subalpine to alpine shrubberies, and in grassland communities in montane and subalpine regions. With the exception of the endemic Decatoca, all the New Guinean genera are widespread, especially Agapetes, Gaultheria, Rhododendron and Vaccinium, the last two genera occurring widely in the Northern Hemisphere.

Family Classification The family is classified in the Ericales (Cronquist 1981). This order has been expanded considerably in recent years and now includes about 20 families (Angiosperm Phylogeny Group 2003). Following analysis of morphological and DNA sequence data, the related families Empetraceae and Epacridaceae were found to nest into Ericaceae and have been subsumed within the latter taxon (Kron et al. 2002; Stevens et al. 2004).

Features of the Family Habit shrubs, sometimes lianas, trees or herbs, sometimes echlorophyllous. Leaves alternate, decussate or whorled, often pseudowhorled, simple, exstipulate. Buds perulate or naked. Indumentum various, often a mixture of unicellular and multiMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

389

................. 16157$

CH23

04-02-07 11:57:34

PS

PAGE 389

.

390 /

cellular hairs, the multicellular hairs glandular or not, unbranched or not, sometimes peltate. Inflorescences terminal or axillary, often racemose or umbellate, sometimes one-flowered. Flowers bisexual; sepals or calyx lobes (1–)4–5(–9), usually connate, sometimes obsolete or wanting; petals(3–)4–5(–9), usually connate; stamens (2–)5–10(–16), free from the corolla or adnate, sometimes connate, included or exserted, the anthers dorsifixed or sometimes basifixed, the apex sometimes extended into two tubules and the body may have two or more appendages, dehiscing by pores or slits; ovary superior to inferior, (2–)4–5(–14)-locular, the style terminal, the stigma truncate to capitate, the placentation usually axile to intruded parietal, rarely apical or basal, the ovules one to many per locule, anatropous or hemitropous to nearly campylotropous. Fruits a capsule, berry, or drupe; seeds commonly numerous, testa usually single-layered, sometimes winged or tailed, endosperm well developed.

Gaps in Floristic Documentation or Knowledge The revisionary studies of H. Sleumer culminated in his account of Ericaceae and Epacridaceae (hereafter the name Ericaceae is used for the merged families) for Flora Malesiana. Sleumer’s work has been the basis of all subsequent studies of the Ericaceae in New Guinea. Because of their showy flowers and appeal to horticulturalists, species of Rhododendron are commonly collected in New Guinea and this genus is well known relative to the comparably-sized Vaccinium. The preparation of a review of Rhododendron in New Guinea is warranted so that the species described (and those reduced to synonymy) since the Flora Malesiana account can be put in context. Vaccinium (sensu Sleumer) in New Guinea requires a great deal more fieldwork (i.e., collecting) before knowledge of its basic taxonomy can be brought up to the same level as that which exists for Rhododendron. For Vaccinium, the development of an interactive identification system may be of great assistance to field and herbarium botanists grappling with the difficulty of naming material in fruit. Agapetes is relatively well collected but its taxonomic status is the subject of current debate with some botanists advocating the transfer of some of the New Guinean species to the Pacific-centered genus Paphia (e.g., Stevens 2004). Another and perhaps preferable possibility is that, rather than circumscribe additional genera, Agapetes, Dimorphanthera, and Vaccinium be merged into a single genus, as has been widely accepted for the various elements brought together in Rhododendron. A similar situation exists with Styphelia, which some botanists (e.g., Quinn et al. 2003) consider should be defined much more narrowly than did Sleumer (1964).

Comparison with Adjacent Regions The relationships of the New Guinean species of Ericaceae indicate that the family in New Guinea has had diverse origins. Agapetes, Dimorphanthera, Diplycosia, Rhododendron, and Vaccinium are genera with their relationships predominantly

................. 16157$

CH23

02-08-07 10:31:22

PS

PAGE 390

Angiosperms: Ericaceae of Papua / 391

to western Malesia and Asia. Although Rhododendron and Vaccinium are very widespread genera outside the New Guinea region, the relationships of their New Guinea species are with species to the west. The poor, or lack of, representation of Agapetes, Dimorphanthera, Diplycosia, Rhododendron, and Vaccinium in Australia is perhaps indicative of the relatively recent dispersal of these genera into New Guinea from the west, with limited onward dispersal into Australia. Of this group of genera, Australia has only two species of Rhododendron and one of Agapetes. The epacridaceous Decatoca, Styphelia, and Trochocarpa are part of the Australiancentered group of genera. Their presence in New Guinea presumably is a result of these species or their progenitors dispersing from Australia. (Geologically, much of New Guinea, together with Australia and Tasmania, comprises a single tectonic mass and logically can be treated as a single biogeographic unit. The name Sahul is sometimes applied to this land.) Gaultheria is an interesting genus. It occurs quite widely (south, east and Southeast Asia, Malesia, eastern Australia, Tasmania, New Zealand, and North and South America) and its New Guinean representatives are related to groups centered in western Malesia–Asia–North America on the one hand, and in Malesia–Australasia–South America on the other (Middleton 1991). Heads (2003) has discussed the biogeography of New Guinea Ericaceae within a broader study of the family in Malesia.

Natural History The New Guinean species of Ericaceae are mainly shrubs, usually of small to medium height, although there are a few species that develop into small to medium height trees. Some species of Agapetes and Dimorphanthera are climbers, occasionally reaching high into the canopy. While perhaps the majority of the species are terrestrial, a significant number are epiphytic. This is not surprising given that the family is essentially heliophytic; the adoption of an epiphytic life form in rainforest brings a plant into a much higher light zone than is found on the ground. The New Guinean Ericaceae predominantly occur in montane to subalpine regions and may comprise a major part of subalpine shrubberies. Knowledge of the pollinators of New Guinea Ericaceae, and of the interactions between flower and pollinator, is very limited, as indeed it is with most New Guinea plants, and field studies should be initiated. Species of the family with small, whitish flowers are probably insect-pollinated; this flower type is characteristic of Gaultheria, Diplycosia, Vaccinium, Styphelia, Decatoca, and Trochocarpa. The larger, usually brightly colored flowers (often reddish or pinkish) of Agapetes and Dimorphanthera apparently are bird-pollinated. The greatest floral diversity in the family in New Guinea, however, is found in Rhododendron. The Rhododendron species may be grouped into five main classes on floral features (Stevens 1976). These classes are: long, tubular, fragrant, white flowers that are probably mothpollinated; subrotate to funnel-shaped, yellow to orange flowers that are probably butterfly- and other insect-pollinated; large, campanulate, fragrant, white to pinkish flowers that are probably moth-pollinated (although blossom bats have also

................. 16157$

CH23

02-08-07 10:31:22

PS

PAGE 391

.

392 /

been suggested as possible pollinators); small- to medium-size, tubular, pink to red flowers that are probably bird-pollinated; medium to large, campanulate (often with the corolla tube curved), pink to red flowers that are quite clearly birdadapted. Although there are few field observations pertaining to the reproductive biology of New Guinea Rhododendron species, a considerable body of knowledge on cultivated species has been built upon observations and experimental research by E. G. Williams and J. L. Rouse and their collaborators (Craven 2003; Williams and Rouse 1988; Williams et al. 1990, 1991). The latter research is largely informative on aspects of the breeding system, notably on pollen-style-ovule interactions, but provides insight into the factors involved in the evolution and maintenance of the remarkable floral diversity exhibited by New Guinean rhododendrons. Rhododendron has capsular fruit and its light, tailed seeds are believed to be winddispersed. The fruits of Agapetes, Dimorphanthera, and Vaccinium are berries and their seeds are probably dispersed by mammals and birds. Gaultheria and Diplycosia have succulent to semisucculent calyces at fruit maturity and dispersal in these genera is doubtless also by mammals and birds. Styphelia, Decatoca, and Trochocarpa have berry-like drupaceous fruit and are no doubt also dispersed endozoically.

Literature Cited Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot J Linn Soc 141: 399–436. Craven, L.A. 2003. Rhododendron publications of John L. Rouse. Available at http://www .vireya.net/references-Rouse.htm (accessed 12 August 2005). Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia University Press, New York. Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Pp. 311–449 in Everett, J., and P.G. Wilson (eds.) Proc. 5th Int. Fl. Malesiana Symp. 2001. Royal Botanic Gardens, Sydney. Kron, K.A., W.S. Judd, P.F. Stevens, D.M. Crayn, A.A. Anderberg, P.A. Gadek, C.J. Quinn, and J.L. Luteyn. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Bot Rev 68: 335–423. Middleton, D.J. 1991. Infrageneric classification of the genus Gaultheria L. (Ericaceae). Bot J Linn Soc 106: 229–258. Quinn, C.J., D.M. Crayn, M.M. Heslewood, E.A. Brown, and P.A. Gadek. 2003. A molecular estimate of the phylogeny of Styphelieae (Ericaceae). Austral Syst Bot 16: 581–594. Sleumer, H. 1964. Epacridaceae. Pp. 422–444 in Van Steenis, C.G.G.J. (ed.) Flora Malesiana ser. I, 6. Wolters-Noordhoff, Groningen. Sleumer, H. 1966–1967. Ericaceae. Pp. 469–914 in Van Steenis, C.G.G.J. (ed.) Flora Malesiana ser. I, 6. Wolters-Noordhoff, Groningen. Stevens, P.F. 1976. The altitudinal and geographical distributions of flower types in Rhododendron section Vireya, especially in the Papuasian species, and their significance. Bot J Linn Soc 72: 1–33.

................. 16157$

CH23

02-08-07 10:31:23

PS

PAGE 392

Angiosperms: Ericaceae of Papua / 393 Stevens, P.F. 2004. New taxa in Paphia and Dimorphanthera (Ericaceae) in Papuasia and the problem of generic limits in Vaccinieae. Edinb J Bot 60: 267–298. Stevens, P.F., J. Luteyn, E.G.H. Oliver, T.L. Bell, E.A. Brown, R.K. Crowden, A.S. George, G.J. Jordan, P. Ladd, K. Lemson, C.B. McLean, Y. Menadue, J.S. Pate, H.M. Stace, and C.M. Weiller. 2004. Ericaceae. Pp. 145–194 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. Springer-Verlag, Berlin. van Royen, P. 1982. Epacridaceae. Pp. 1901–1926 in The Alpine Flora of New Guinea 3. Cramer, Vaduz. van Royen, P., and P. Kores. 1982. Ericaceae. Pp. 1485–1900 in The Alpine Flora of New Guinea 3. Cramer, Vaduz. Williams, E.G., and J.L. Rouse. 1988. Disparate style lengths contribute to isolation of species in Rhododendron. Austral J Bot 36: 183–191. Williams, E.G., J.L. Rouse, V. Kaul, and R.B. Knox. 1991. Reproductive timetable for the tropical Vireya Rhododendron, R. macgregoriae. Sex Pl Reprod 4: 155–165. Williams, E.G., J.L. Rouse, B.E. Palser, and R.B. Knox. 1990. Reproductive biology of Rhododendron. Hort Rev 12: 1–67.

................. 16157$

CH23

02-08-07 10:31:23

PS

PAGE 393

Euphorbiaceae of Papua . Number of Genera and Species 326 genera of Euphorbiaceae are known, with about 7,750 species. In Malesia about 115 genera and 1,600 species are present. New Guinea comprises 54 genera with ca 460 species of which 73% are endemic.

W

Distribution and Habitat The Euphorbiaceae are a cosmopolitan family, present from the tropical to subarctic regions. New Guinea is a major center of biodiversity, though most genera are less speciose than in Western Malesia (Malaysia, Sumatra, Borneo, Java). About a quarter of the Euphorbiaceae species in New Guinea have a widespread distribution, either throughout Malesia or also present in Australia or the Pacific Islands. Euphorbiaceae can be found in all habitats, ranging from lowland to upper montane areas, and on (almost) all kinds of soil. Some prostrate species of Euphorbia will be present in cracks in the pavement and herbs in the genera Acalypha and Phyllanthus are usually present on waste areas in cities. Most species of Euphorbiaceae are usually found in the more secondary surroundings, though they also occur in primary rainforest. Quite a few species are pioneers in primary forest, as soon as a gap occurs they will appear and may remain present while primary forest species establish themselves (see ‘‘Pioneers’’ section, below). Few species are adapted to a single habitat. Excoecaria agallocha is a mangrove species, though it is also found outside the mangrove on silty areas inland. Some species, like Euphorbia atoto, are typical of sandy beaches.

Family Classification Traditionally the Euphorbiaceae are subdivided into five subfamilies (RadcliffeSmith 2001). The Oldfieldioideae and Phyllanthoideae have two ovules per locule; the Acalyphoideae, Crotonoideae, and Euphorbioideae have a single ovule per locule. All five subfamilies are represented in New Guinea. On the basis of molecular phylogenetic research the family is about to be divided into five separate families (Angiosperm Phylogeny Group 2003). The uni-ovulate subfamilies will then form the Euphorbiaceae sensu stricto. The Oldfieldioideae will become the Picrodendraceae, the Phyllanthoideae the Phyllanthaceae, the genera Panda, Galearia, and Microdesmis will become the Pandaceae (not to be confused with Pandanaceae), and the tribe Drypeteae in the Phyllanthoideae will become the Putranjivaceae. Most of these new families, like the Euphorbiaceae in the broad sense, cannot be defined by specific characteristics, because there are always exceptions. This Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

394

................. 16157$

CH24

02-08-07 10:31:28

PS

PAGE 394

Angiosperms: Euphorbiaceae of Papua / 395

chapter considers the Euphorbiaceae in the broad sense, comprising five subfamilies.

Features of the Family The Euphorbiaceae are an extremely variable family; summarizing all characteristics is impossible because of the many exceptions. The only common characteristic they have is the superior ovary, so the Euphorbiaceae usually key out together with any family with a superior ovary. Most typical of the Euphorbiaceae is their fruit: a dry capsule that ‘‘explodes,’’ leaving loose seeds, loose fruit parts, and a persistent column (but again, many genera provide exceptions, having drupes or berries). The most recent revision of all New Guinean Euphorbiaceae is by Airy Shaw (1980). Habit: prostrate to erect herbs, shrubs, trees, or lianas, sometimes spiny and/or succulent, monoecious or dioecious. Indumentum absent to present all over, hairs simple, bundled, stellate, lepidote, and/or glandular; in some genera glandular scale hairs (Macaranga, Mallotus, Octospermum). Stipules usually present, usually caducous, free or sometimes connate. Leaves usually simple, some 3-foliolate (Bischofia, Hevea) or digitate (Annesijoa), alternate to opposite to whorled, usually petiolate with, in several genera, raised glands at the apex, petiole usually basally and/or apically pulvinate; blade very variable in size, shape, base, margin, and apex; venation penninerved, basally 3-nerved, or palminerved; often glandular areas (extrafloral nectaries) near base, apex or margin on upper or lower surface, marginal teeth often with apical glands. Inflorescences cauliflorous to axillary to terminal, either cymose, thyrsiform, racemose, spicate, paniculate, or flowers in fascicles, sometimes opposite to the leaf (Suregada); in case of monoecy both sexes often in separate inflorescences, or if in one inflorescence at different positions, seldom both sexes originating from the same node; bracts with or without glands usually present, with a single to several flowers. Flowers unisexual (exceptionally bisexual, such as in several species of Aporosa in New Guinea; Schot 2004), usually actinomorphic, very small, usually (shortly) pedicelled. Sepals seldom absent, free to united, valvate or imbricate, usually ca five lobes of often different sizes, in a few genera pistillate ones enlarging in fruit. Petals absent to present in a single to both sexes, usually free, sometimes brightly coloured. Disk often present, in staminate flowers often separate glands, in pistillate flowers usually an annular ring. Stamens one to 1,000, free to variously connate, sometimes branching (Ricinus, Spathiostemon), filaments short to long (stamens exserted), anthers 2–4-locular, erect or inflexed in bud (Croton), thecae in various sizes, shapes, and arrangements, opening with lengthwise slits, often a gland on the connective. Ovary superior, usually 3-locular, smooth or with spines, ovules one or two per locule, apical, hanging, anatropous or hemitropous; style absent to present, stigmas often apically split (to completely split), usually with long (branching) papillae adaxially. Fruit usually dry capsular, usually dehiscing loculicidally and septicidally, sometimes only one type partly so, leaving V-shaped segments, smooth to armed with

................. 16157$

CH24

02-08-07 10:31:28

PS

PAGE 395

.

396 /

spines, after dehiscence an angular, apically somewhat winged column remaining; indehiscent drupes and berries also present. Seeds one or two per locule or (by abortion) one per fruit (Antidesma, part of Bridelia), usually more or less obovoid with a ventral ridge, often an apical caruncle (fleshy appendage), sometimes a fleshy sarcotesta or aril; endosperm absent to copious; embryo straight, curved, or folded.

Gaps in Floristic Documentation or Knowledge Much about the ecology of Euphorbiaceae is still unknown. Probably many new species are yet uncollected, and most distributions of species are incomplete because of largely undercollected areas, especially in Papua.

Natural History Little is known about the floral biology of Euphorbiaceae. Most species have inconspicuous flowers that are not attractive to many insects. Moog et al. (2002) demonstrated that Macaranga hulletii King ex Hook.f. (Southeat Asia mainland and western Malay Archipelago) is pollinated by thrips of the genus Neoheegeria. Probably most species of Macaranga are pollinated by thrips, especially on Borneo (U. Moog, pers. comm.). Also recently, a team of Japanese botanists showed that some Southeast Asian mainland species of Glochidion are pollinated by moths of the genus Epicephala (Gracillariidae) (Kato, Takimura, and Kawakita 2003). A female actively collects pollen from staminate flowers and ‘‘injects’’ the pollen into the stigmatal cone with her proboscis, after which she oviposits into the style. The larvae of the moth eat only a part of the developing seeds. This mechanism seems likely for almost all species of Glochidion because the stigmas are united into a cone and no free pollen-receiving stigma papillae are present, and these papillae are all hidden inside the cone. Kato and colleagues stress that this places Glochidion in a position similar to that of Ficus and Yucca, genera in which the species are also obligately pollinated by a specialized group of insects. Members of the same team have now demonstrated that two related taxa, Breynia and Phyllanthus subgenus Gomphidium, are also pollinated by the same moths (Kawakita and Kato 2004).

Little is known about seed or fruit dispersal in Euphorbiaceae. Labels on specimens regularly indicate that the seeds are eaten and perhaps dispersed by birds. The capsular fruit of Homalanthus novoguineensis are consumed by an array of birds of paradise (Beehler 1983) and Macaranga giganteus is favored by species of manucode (B. Beehler, pers. comm.). Most seeds usually have no or almost no outer fleshy part that can be rewarding for birds; therefore, probably many are damaged by passing through the bird’s intestines. Ants probably transport to their

................. 16157$

CH24

02-08-07 10:31:29

PS

PAGE 396

Angiosperms: Euphorbiaceae of Papua / 397

nests seeds that have at least a caruncle. The most common dispersal mechanism may be simple barochory (i.e., the seeds drop to the ground after dehiscence of the fruit). The dehiscence of the fruits can be somewhat explosive and seeds may be strewn away from the parent tree. The more fleshy fruits or seeds with a thick fleshy sarcotesta are probably eaten by birds, animals, and humans (e.g., Antidesma, Baccaurea, Bridelia). No phylogenetic signal has so far been detected in fruit development (e.g., from capsular to drupaceous) in the most recent molecular phylogenies (e.g., Kathriarachchi et al. 2005). An overview of some of the Euphorbiaceae seeds and seedlings can found in Ng (1991). Seeds of the climax species have no or a short dormancy (up to one year). Seeds of the pioneer species probably have a long dormancy; they germinate from the seedbank as soon as a gap in the forest appears. Two groups of Macaranga (not present on New Guinea) are renowned for their symbiosis with ants. The plants provide housing, and the ants can even farm their bugs inside the plant. Macaranga and Mallotus in New Guinea often show leaves with extrafloral nectaries. In fact many species in the uni-ovulate Euphorbiaceae show all kinds of extrafloral glands, not only on the leaves, but quite often also in the inflorescences. These are usually visited by ants, which probably means that ants live on the plants in exchange for protection against insects, epiphytes, and neighboring plants. Recently it was shown that the Bornean large-leaved species of Macaranga and Mallotus are pioneer species that germinate from the seedbank as soon as disturbance occurs. Because of their large leaves, they provide shade that enables other species to germinate, which then replace most of the Macaranga and Mallotus pioneers after one or two decades. Macaranga and Mallotus also appeared to be good indicator species for the type and age of the disturbance. Different species tend to germinate after selective logging, burning or repeated burning, and clearcutting (Slik, Keßler, and van Welzen 2003). In New Guinea species with the same syndrome are present: small-leaved species live in climax vegetation while largeleaved species are pioneers. Generally, Macaranga and Mallotus are lowland species; in the mountains, species of Homalanthus (Omalanthus) are found in a similar ecological role (H.-J. Esser, pers. comm.).

Literature Cited Airy Shaw, H.K. 1980. The Euphorbiaceae of New Guinea. Kew Bull. Add. Ser. 8: 1–243. Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399–436.

................. 16157$

CH24

02-08-07 10:31:29

PS

PAGE 397

398 /

.

Beehler, B.M. 1983. Fruigivory and polygamy in birds of paradise. Auk 100: 1–12. Kathriarachchi, H., P. Hoffmann, R. Samuel, K.J. Wurdack, and M.W. Chase. 2005. Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3’ndhF, rbcL, and nuclear PHYC). Molec. Phylog. Evol. 36: 112–134. Kato, M., A. Takimura, and A. Kawakita. 2003. An obligate pollination mutualism and reciprocal diversification in the tree genus Glochidion (Euphorbiaceae). PNAS 100: 5264–5267. Kawakita, A., and M. Kato. 2004. Obligate pollination mutualism in Breynia (Phyllanthaceae): further documentation of pollination mutualism involving Epicephala moths (Gracillariidae). Am. J. Bot. 91: 1319–1325. Moog, U., B. Fiala, W. Federle, and U. Maschwitz. 2002. Thrips pollination of the dioecious ant plant Macaranga hullettii (Euphorbiaceae) in southeast Asia. Am. J. Bot. 89: 50–59. Ng, F.S.P. 1991. Manual of forest fruits, seeds and seedlings 1. Malayan For. Rec. 34: 67–83, figs. 57–69, 321–366. Radcliffe-Smith, A. 2001. Genera Euphorbiacearum. Royal Botanic Gardens, Kew. Schot, A.M. 2004. Systematics of Aporosa. Blumea Supplement 17: 1–377. Slik, J.W.F., P.J.A. Keßler, and P.C. van Welzen. 2003. Macaranga and Mallotus species (Euphorbiaceae) as indicators for disturbance in the mixed lowland dipterocarp forest of East Kalimantan (Indonesia). Ecol. Indic. 2: 311–324.

................. 16157$

CH24

02-08-07 10:31:30

PS

PAGE 398

Melastomataceae of Papua . in the narrow sense, excluding their sister family Memecylaceae (Angiosperm Phylogeny Group 2003), comprise some 3,000 species in the neotropics, 240 in Africa, 230 on Madagascar, and about 1,000 in Asia. Papua harbors 12 genera, with perhaps 160 species (Table 3.6.4). Most of the Papuan species are woody climbers (Catanthera, Dissochaeta, Poikilogyne, Medinilla section Heteroblemma), epiphytic shrubs (Medinilla), or terrestrial shrubs (Astronia, Astronidium, Pternandra). Interestingly, Papua seems to lack herbaceous Melastomataceae and has very few tree species (some Astronia and Pternandra). Most species occur in primary and secondary forests, often near water; a few of the herbaceous species are confined to rock outcrops.

M

Historical Background The earliest contribution focused on the Melastomataceae of Papua is a study by Mansfeld (1925) of collections made mainly in eastern New Guinea by Rudolf Schlechter and E. Keyser in what was then German New Guinea. The other important early collector was J. J. Smith who traveled in the western, then Dutch, half of the island. Schlechter paid two lengthy visits to New Guinea, one in 1901–1902 and the other in 1907–1908, and during these expeditions he visited New Ireland and New Britain (see Chapter 1.2). Mansfeld described two new genera, Phyllapophysis, with a single species, P. schlechteri, now considered a species of Catanthera, and Scrobicularia, which included only S. callantha, now considered a species of Poikologyne. A third genus from Papua, Bamlera (Schumann and Lauterbach 1900), is now treated as a synonym of Beccarianthus. All of the new taxa were climbers, with highly polymorphic leaves and inflorescences.

Morphological Characteristics of Melastomataceae Inflorescences of Melastomataceae are terminal or axillary, paniculate or umbellate, usually with many white or purplish flowers. The calyx is truncate with external teeth or lobed. Flowers are usually 4-merous, more rarely 5-merous (some Medinilla and Pachycentria); usually there are twice as many stamens as petals, but sometimes the inner stamens are staminodial (some species of Dissochaeta) or completely reduced (Catanthera, some Dissochaeta). The stamens open apically by one or two pores or by slits (e.g., Pternandra, Astronia, Astronidium, Beccarianthus), and the connective is ventrally or dorsally lobed or adorned with a tuft of papillae or hairs. The ovary is more or less deeply inferior and has 4 or rarely 3, 5 or 6 locules. Placentation of seeds is axillary except in Pternandra, where the plaMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

399

................. 16157$

CH25

02-08-07 10:31:36

PS

PAGE 399

................. 16157$

CH25

02-08-07 10:31:36

PS

India to Indo-China and throughout the Malesian region and Oceania

East Africa and throughout the Malesian region

Burma, Thailand, and throughout the Malesian region to New Guinea

Centered in Borneo and New Guinea

Thailand to Borneo and New Guinea

Melastoma

Ochthocharis

Pachycentria

Poikilogyne

Pternandra

Note: Estimates of species numbers in Papua are speculative.

Burma to Indo-China and throughout the Malesian region

Medinilla

Burma to Indo-China and throughout the Malesian region

New Guinea, Java, Philippines

Creochiton

Dissochaeta

New Guinea, Borneo, Sumatra

Catanthera

Astronidium

Borneo, Philippines, New Guinea

New Guinea to the Pacific (Society Islands)

Astronia

Beccarianthus

Distribution

Philippines, Borneo, New Guinea

Genus

15

20

8

7

20

150–200

40

10

13–17

22

67

59

Total number of species

4

7

2

3

6

15

5–10

4–5

9

10

50

30

Number of species in Papua

.

Maxwell 1981 (including Kibessia )

Mansfeld 1925; Nayar 1973; Maxwell 1982 (incl. Dicerospermum Bakh.f. and Scrobicularia Mansf. )

Clausing 2000 (including Pogonanthera Bl.)

Hansen and Wickens 1981

Meyer 2001 (including Otanthera )

Bodegom and Veldkamp 2001

Maxwell 1980a,b, 1983 (incl. Anplectrum, Backeria, Dalenia, Diplectria, Macrolenes, Omphalopus); see also Veldkamp et al.1978; Renner et al. 2001

Maxwell 1983 (incl. Eisocreochiton Quisumb. & Merrill and Anplectrella Furtado)

Nayar 1982 (incl. Hederella Stapf, Phyllapophysis Mansf. )

Maxwell and Veldkamp 1990b (including Bamlera Lauterbach & K. M. Schum. Everetttia Merrill )

Maxwell and Veldkamp 1990b

Maxwell and Veldkamp 1990a

Taxonomic treatment

Table 3.6.4. Regional distribution of Melastomataceae with recent taxonomic treatments

400 /

PAGE 400

Angiosperms: Melastomataceae of Papua / 401

centae are located on the outer walls of the locules, and in Astronia, Astronidium, Beccarianthus (and their relative Astrocalyx, not found in New Guinea) where placentae are located basally in each locule (Maxwell and Veldkamp 1990a,b). Fruits in Melastomataceae are fleshy or dry and ribbed. The seeds are straight, oblong, ovoid, or wedge-shaped. Papua has an unusually large number of climbing and epiphytic species. Climbing has evolved several times in Melastomataceae, and the respective species exhibit a wide range of morphological adaptations, including branch-borne roots, reduced leaves, unusual wood, and hollow roots or stems occupied by ants (Clausing and Renner 2001a). Among the species occurring on Papua, scrambling growth is found in Dissochaeta and its close relatives (or synonyms) Macrolenes and Diplectria. Root climbing, on the other hand, is found in Catanthera, Kendrickia, and Medinilla section Heteroblemma. A morphological cladistic analysis (Renner 1993) and subsequent molecular phylogenetic work (Clausing and Renner 2001b; Renner 2004) showed that Pternandra and Astronieae (Astronia, Astronidium, Astrocalyx, and Beccarianthus) are successive sister groups to the remaining Melastomataceae. The basal placement of these lineages in the family tree fits with their retaining ancestral traits, such as anther dehiscence by two longitudinal slits instead of by one or two pores as in the more derived Melastomataceae. It may be biogeographically significant that both these basal clades are well represented on New Guinea. The sister group of Melastomataceae, Memecylaceae, includes 360 species in four genera in Africa, Madagascar, and Southeast Asia, and some 90 species in two genera in tropical America. The next closely related clade (Alzateaceae, Crypteroniaceae, Oliniaceae, Penaeaceae, and Rhynchocalycaceae) occurs mainly in Africa and Southeast Asia, and is thought to date back to the mid-Cretaceous (see Renner 2004, for references). The geographic origin and age of the New Guinea Melastomataceae flora would make an interesting subject for a future molecular clock-based study.

Ecology and Biology of Papuan Melastomataceae Though there are no published studies on the pollination or seed dispersal of any melastomes in New Guinea, some aspects can be extrapolated from congeneric species with similar flowers or fruits that have been studied elsewhere. Genetic self-incompatibility and self-compatibility have both been documented in Melastomataceae, and so has asexual seed formation. One may therefore assume that these mating systems are also found among Papuan species. The incidence of polyploidy in Melastomataceae overall seems high, but only about eight percent of the species worldwide have been counted (and none from Papua). Most Melastomataceae have bisexual flowers, but unisexual flowers distributed on different (male and female) individuals are known from a few species of Astronia. The principal mode of promoting outcrossing in Melastomataceae, however, is spatial separation of pollen and stigma, achieved by the pollen being enclosed in poricidal anthers that have to be manipulated by an animal, usually a pollen-collecting bee,

................. 16157$

CH25

02-08-07 10:31:37

PS

PAGE 401

.

402 /

to release pollen grains. The reward offered in a typical melastome flower is pollen; nectar production is very rare. Bee species known to visit Melastomataceae are listed in Renner (1993) and comprise a wide spectrum of the world’s bee lineages, including carpenter bees and stingless bees (that may steal the pollen). It is not yet clear whether the conspicuous stamen appendages found in many Melastomataceae have a function in the pollination mechanism beyond that of enhancing the visual attractiveness of the flowers and making the stamens easier to grasp for the bees. Seed dispersal in those Melastomataceae that have capsular fruits is by wind. In species with berries, the fruits are usually taken by birds, but also by marsupials, primates, bats, other mammals, turtles, and other reptiles.

Literature Cited Angiosperm Phylogeny Group 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Botanical Journal of the Linnean Society 141: 399–436. Bodegom, S., and J.F. Veldkamp. 2001. A revision of the pseudo-stipular species of Medinilla Gaud. ex DC. (Melastomataceae–Melastomatoideae–Miconieae). Blumea 46: 527–567. Clausing, G. 2000. Revision of Pachycentria (Melastomataceae). Blumea 45: 341–375. Clausing, G., and S.S. Renner. 2001a. Evolution of growth form in epiphytic Dissochaeteae (Melastomataceae). Org. Divers. Evol. 1: 45–60. Clausing, G., and S.S. Renner. 2001b. Molecular phylogenetics of Melastomataceae and Memecylaceae: Implications for character evolution. Am. J. Bot. 88: 486–498. Hansen, C., and G.E. Wickens. 1981. A revision of Ochthocharis (Melastomataceae) including Phaeoneuron in Africa. Kew Bull. 36: 13–29, Pl. 2, 3. Mansfeld, R. 1925. Die Melastomataceen von Papuasien. Bot. Jahrb. Syst. 60: 115–130. Maxwell, J.F. 1980a. Taxonomic revision of Diplectrinae Maxw. and Dissochaetinae Naud. (Dissochaeteae) Melastomataceae. Ph.D. diss., Department of Botany, University of Singapore, Singapore. Maxwell, J.F. 1980b. Taxonomic notes on the tribe Dissochaeteae (Naud.) Triana (Melastomataceae). Gard. Bull. (Singapore) 33: 312–327. Maxwell, J.F. 1981. A revision of the genus Pternandra (Melastomataceae). Gard. Bull. (Singapore) 34: 1–90. Maxwell, J.F. 1982. Taxonomic and nomenclatural notes on Oxyspora DC., Anelrincleistus Korth., Poikilogyne Baker f., and Allomorphia Bl. (Melastomataceae, tribe Oxysporeae). Gard. Bull. (Singapore) 35: 209–226. Maxwell, J.F. 1983. Taxonomic studies of the Melastomataceae (Part 1). A revision of subtribes Diplectriinae Maxw. and Dissochaetinae (Naud.) Triana (Genera Diplectria (Bl.) [sic], Dissochaeta Bl., Macrolenes Naud., Creochiton Bl., and Pseudodissochaeta Nayar). Fed. Mus. Journal 29: 45–117. Maxwell, J.F., and J.F. Veldkamp. 1990a. Notes on the Astronieae (Melastomataceae) I. Astrocalyx, Astronia. Blumea 35: 71–114. Maxwell, J.F., and J.F. Veldkamp. 1990b. Notes on the Astronieae (Melastomataceae) II. Astronidium, Beccarianthus. Blumea 35: 115–165.

................. 16157$

CH25

02-08-07 10:31:37

PS

PAGE 402

Angiosperms: Melastomataceae of Papua / 403 Meyer, K. 2001. Revision of the Southeast Asian genus Melastoma (Melastomataceae). Blumea 46: 351–398. Nayar, M.P. 1973. Three new species of Poikilogyne (Melastomataceae) from New Guinea. J. Jap. Bot. 48: 238–241. Nayar, M.P. 1982. Revision of the genus Catanthera F. v. Muell. (Melastomataceae). Reinwardtia 10: 35–61. Renner, S.S. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae. Nord. J. Bot. 13: 519–540. Renner, S.S. 2004. Bayesian analysis of combined data partitions, using multiple calibrations, supports recent arrival of Melastomataceae in Africa and Madagascar. Am. J. Botany 91: 1427–1435. Renner, S.S., G. Clausing, N. Cellinese, and K. Meyer. 2001. Melastomataceae. Flora of Thailand 7: 412–497. Schumann, K., and K. Lauterbach. 1900. Die Flora der Deutschen Schutzgebiete in der Su¨dsee. Borntraeger, Leipzig. Veldkamp, J.F., N.A.P. Franken, M.C. Roos, and M.P. Nayar. 1978. A revison of Diplectria (Melastomataceae). Blumea 24: 405–430.

................. 16157$

CH25

02-08-07 10:31:38

PS

PAGE 403

Moraceae of Papua . Number of Genera and Species T 37 genera of Moraceae have a broad range of inflorescence forms, pollination syndromes, and breeding systems (Datwyler and Weiblen 2004). Most of the 1,100 species are figs (Ficus) known for a unique inflorescence and obligate pollination mutualism with fig wasps. In Papua, there are ten genera and 173 described species, dominated by Ficus, with 151 species, and followed by Artocarpus, with seven.

Distribution and Habitat Moraceae are distributed from tropical to temperate forests throughout the world but the great majority of species are restricted to tropical rainforest. In New Guinea, the family occurs from the lowlands to cloud forest up to 2,400 m above sea level. Ficus is a prominent member of forest communities throughout the island in terms of local species richness and abundance (Weiblen 1998). About 70% of these species are endemic to the island and alpha diversity is extreme. In a lowland rainforest, for example, it is not uncommon to encounter up to 50 Ficus species within a few hundred hectares. Species turnover at a regional scale, on the other hand, appears to be quite low. In a comparison of four lowland rainforests across the Ramu and Sepik river basins, for instance, at least half of the Ficus species are shared between any two sites, even those separated by 500 km (G. D. Weiblen, unpublished data). The uneven density of collections across the island make it difficult to discern patterns of local endemism but most species appear to be widespread, perhaps related to dispersal by vertebrate frugivores (Dumont et al. 2004). Merely seven Ficus species (4%) appear to be limited to Papua, and only four (2%) to New Britain and the Bismarck Archipelago. The generally widespread distribution of Ficus species runs contrary to the biogeographic notion that mountainous New Guinea is an engine of plant speciation because of geographic isolation of populations. As we shall see, the mode of speciation in Ficus is likely related to the uniquely specialized mode of pollination (Weiblen and Bush 2002). The fossil record and molecular divergence of Moraceae suggest that the family is at least 90 million years old and the distribution of tribes suggests either a Gondwanan or Laurasian origin (Datwyler and Weiblen 2004).

Family Classification Molecular data indicate that Moraceae are part of the Rosidae and are closely allied to the Urticalean rosids, including Cannabaceae, Celtidaceae, Urticaceae, Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

404

................. 16157$

CH26

02-08-07 10:31:45

PS

PAGE 404

Angiosperms: Moraceae of Papua / 405

Cecropiaceae, and Ulmaceae (Sytsma et al. 2002). The Urticalean rosids differ from most other rosids in the presence of solitary ovules, laticifers, cystoliths, paired inflorescences in the leaf axils, and unisexual flowers. Urticaceae plus Cecropiaceae are sister to the Moraceae, distinguished from the latter in having lacifers only in the bark, clear latex, and orthotropous ovules (Sytsma et al. 2002). Moraceae are divided into five tribes according to a recent phylogenetic classification of the family (Datwyler and Weiblen 2004). These are represented in New Guinea by the monotypic Ficeae, the Artocarpeae with Artocarpus, Parartocarpus (1 species) and Prainea (1), the Castilleae with Antiaris (1) and Antiaropsis (1), and the Moreae with Fatuoa (1), Maclura (3), Streblus (3), and Trophis (4). Paperbark mulberry (Broussonetia) is introduced.

Features of the Family Habit includes trees, shrubs, hemiepiphytes, climbers and herbs. Stipules are present. Leaves are alternate, simple, entire to lobed to serrate, with milky latex in all parenchymatous tissue. Inflorescences distinguish the tribes. Ficeae are characterized by a syconium, an urn-shaped receptacle lined with flowers and enclosed at the apex by a ring of bracts. Plants are monoecious with bisexual inflorescences or gynodioecious but functionally dioecious (Weiblen 2000). Artocarpeae are dioecious or monoecious with unisexual inflorescences ranging from spikes to globes to solitary flowers. Castilleae are dioecious or monoecious with discoid to urceolate receptacles ringed by bracts. Moreae are dioecious to monoecious with relatively simple racemes, spikes or globose heads. Flowers are reduced and, when present, the perianth is 4- or 5-merous, tepaloid, and membranous. Filaments are either straight or inflexed in bud. Inflexed stamens in the Moreae are associated with a pistillode against which the anthers are appressed in bud. These stamens, springing outward at anthesis to release their pollen, are associated with wind pollination. The perianths of pistillate flowers are often fused to the receptacle, a condition hypothesized to protect the flowers against phytophagous insects (Berg 1990). Fruits are aggregates of drupes or achenes, often with a fleshy receptacle.

Gaps in Floristic Documentation or Knowledge Probably the greatest turnover in floristic composition of Moraceae occurs across the altitudinal gradient from lowland forest to montane forest. However, the altitudinal limits of many Ficus species in particular are not well documented. There is a need to explore the species limits of montane versus lowland taxa, as well as patterns of intraspecific variation across the island, and species limits in the complexes of climbing figs (35 New Guinean species in subgenus Synoecia), and cauliflorous figs (50 New Guinean species in subgenus Sycomorus).

Natural History Fig pollination is one of the best known examples of obligate mutualism between plants and insects, and was reviewed by Cook and Rasplus (2003). In general, the

................. 16157$

CH26

02-08-07 10:31:45

PS

PAGE 405

.

406 /

associations between figs and their pollinators, microscopic wasps in the subfamily Agaoninae of parasitic Hymenoptera, are species-specific. It appears that each Ficus species in New Guinea is associated with a unique pollinating wasp species (Wiebes 1982). Female fig wasps actively or passively pollinate fig flowers while galling a fraction of fig ovaries. Each fig wasp larva feeds on and destroys a single developing ovule. The fate of ovaries as either seeds or galls depends on the intricate interactions between the style lengths of fig flowers and the ovipositor lengths of pollinators (Weiblen 2004). In monoecious fig species, seeds and wasps develop within each fig due to the presence of range of style lengths. In functionally dioecious species, seeds and wasps develop in different figs on separate plants (Weiblen et al. 2001). The pollination mutualism in New Guinea is also impacted by nonpollinating fig wasps, including gallers and parasitoids. In contrast to the host-specific associations between Ficus species and their pollinator wasps, seed dispersal is accomplished through more generalized associations with vertebrate frugivores. Figs are eaten by birds, bats, and marsupial mammals in New Guinea, which mostly pass the seeds undamaged through their digestive systems, resulting in dispersal. Most ripe fig crops are visited by more than one animal species and different fig species may attract different guilds of animals (Shanahan et al. 2001). Feeding preferences appear to be influenced by traits including the position of the figs on the plant as well as fig color, size, and nutritional content. The distinctive geocarpic species that bear figs on leafless runners at or just below the soil surface appear to be dispersed by bandicoots, pigs, and rats. Cauliflorous trees are often visited by bats (Dumont et al. 2004), while the hemiepiphytic stranglers have axillary figs consumed by birds including cuckoo shrikes, birds of paradise, bowerbirds, pitohuis, hornbills, and parrots. The endemic Vulturine Parrot appears to be a particular specialist on the distinctive figs of section Malvanthera, where the seeds are embedded in a lignified endocarp and receptacle (Mack and Wright 1998). The hemiepiphytic growth habit of figs in subgenus Urostimga (23 species in New Guinea) is also noteworthy. The seeds of all members in this group germinate in the forest canopy and become established as epiphytes, later sending aerial roots to the ground on the way to reproductive maturity. Some species strangle and kill their host tree to become free-standing trees with aerial roots while others remain entirely dependent on their hosts for structural support. This growth habit contrasts sharply with the climbing figs (subgenus Synoecia) which recruit from the forest understory and reach the canopy by means of adventitious roots adhering to host tree trunks. Moraceae are also economically important in subsistence agriculture with Ficus playing a major role in forest regeneration following gardening. The edible leaves and fruits of secondary forest species including F. copiosa, F. wassa, and F. dammaropsis are used throughout New Guinea in traditional cooking. Wild and cultivated breadfruit (Artocarpus camansi and A. altilis, respectively) are an important source of starch in Melanesia. In addition, the fibrous bark of many arborescent Moraceae is the source of tapa cloth. Boiled leaves of Ficus pachyrrhachis are used

................. 16157$

CH26

02-08-07 10:31:46

PS

PAGE 406

Angiosperms: Moraceae of Papua / 407

to dye traditional grass skirts and the latex of many species, especially Ficus septica and Maclura cochinchinensis, is employed in traditional medicine.

Literature Cited Berg, C.C. 1990. Differentiation of flowers and inflorescences of Urticales in relation to their protection against breeding insects and pollination. Sommerfeltia 11: 13–34. Cook, J.M., and J.-Y. Rasplus. 2003. Mutualists with attitude: coevolving fig wasps and figs. Trends in Ecology and Evolution 18: 241–248. Datwyler, S.L., and G.D. Weiblen. 2004. On the origin of the fig: Phylogenetic relationships of Moraceae from ndhF sequences. American Journal of Botany 91: 767–777. Dumont, E.R., G.D. Weiblen, and J.R. Winklemann. 2004. Preferences of fig wasps and fruit bats for figs of functionally dioecious Ficus pungens. Journal of Tropical Ecology 20: 233–238. Mack, A.L., and D.D. Wright. 1998. The Vulturine Parrot, Psittrichas fulgidus, a threatened New Guinea endemic: notes on its biology and conservation. Bird Conservation International 8: 185–194. Shanahan, M., S. So, S.G. Compton, and R. Corlett. 2001. Fig-eating by vertebrate frugivores: a global review. Biological Reviews 76: 529–572. Sytsma, K.J., J. Morawetz, J.C. Pires, M. Nepokroeff, E. Conti, M. Zjhra, J.C. Hall, and M.W. Chase. 2002. Urticalean Rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89: 1531–1546. Weiblen, G.D. 1998. Forest composition and structure of a one hectare plot in the Crater Mountain Wildlife Management Area, Papua New Guinea. Science in New Guinea 24: 23–32. Weiblen, G.D. 2000. Phylogenetic relationships of functionally dioecious Ficus (Moraceae) based on ribosomal DNA sequences and morphology. American Journal of Botany 87: 1342–1357. Weiblen, G.D. 2004. Correlated evolution in fig pollination. Systematic Biology 53: 128–139. Weiblen, G.D., and G.L. Bush. 2002. Speciation in fig pollinators and parasites. Molecular Ecology 11: 1573–1578. Weiblen, G.D., D.W. Yu, and S.A. West. 2001. Pollination and parasitism in functionally dioecious figs. Proceedings of the Royal Society of London Series B 268: 651–659. Wiebes, J.T. 1982. Fig wasps (Hymenoptera). Pp. 735–755 in J.L. Gressitt, (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague.

................. 16157$

CH26

02-08-07 10:31:46

PS

PAGE 407

Myristicaceae of Papua . . . Number of Genera and Species Myristicaceae is a well-defined middle-sized family, with 20 genera and ca 500 species worldwide. The number of genera is about equally divided over the tropics, with six genera (ca 350 species) in Southeast Asia, including New Guinea. The largest genus is Myristica, which is the most speciose in New Guinea and contains many endemics. The next most speciose genus is Horsfieldia. The genus Paramyristica is monotypic to New Guinea.

T

Distribution and Habitat The Myristicaceae are distributed in lowland rainforests throughout the tropics. They are normally limited to the lowlands, and only in New Guinea do several endemic species of Myristica penetrate the cooler environments of montane forests. The highest altitude at which this family has been reported in the world is 2,200 m, where a subspecies of Myristica subalulata was seen. A few species are widespread in Southeast Asia, including Horsfieldia irya, distributed in mostly riparian habitat from Sri Lanka to New Guinea. Papuan Myristicaceae are typically members of primary rainforest vegetation, including kerangas (heath) and marshy forest. The few exceptions are found in the genus Horsfieldia, which contains some species that seem to thrive in (wet) disturbed forest (e.g., H. laevigata). Most species are understory or midstory trees, but some do reach the high canopy. Others are essentially understory small trees or shrubs, notably a separate group of species of Horsfieldia (e.g., H. crux-militensis and allies) and some species in Myristica.

Family Classification The Myristicaceae are placed within a distinct suborder within the Magnoliales, which is an order next to the order Laurales. The family is well defined both macromorphologically and molecularly. The genera are rather different in the number of species that they contain, but can easily be told apart on the basis of inflorescence morphology, the presence or absence of bracteoles, the construction of the androecium (in the male flowers), and the condition of the showy aril of the seed. These characters can be used to differentiate the six New Guinea genera as follows:

Practical Key to the Genera The male inflorescence can be either a densely flowered (sub)sessile mostly unbranched, woody, brachyblast with the flowers fascicled at the apex, or Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

408

................. 16157$

CH27

02-08-07 10:31:51

PS PAGE 408

Angiosperms: Myristicaceae of Papua / 409

distinctly peduncled and branched, non-woody, of two types, simple or compound: Male inflorescence an unbranched brachyblast in Knema and Myristica (partly) Male inflorescences peduncled and/or branched (paniculate) in Endocomia, Gymnacranthera, Horsfieldia, Myristica (partly), and Paramyristica

Presence or absence of a bracteole on the male pedicel: Bracteole present in Knema and Myristica Bracteole absent in Endocomia, Gymnacranthera, Horsfieldia, and Paramyristica

Morphology of the androecium (see Figure 3.6.9): Androecium peltate, with separate sessile anthers around a stalked disc in Knema

The other genera have largely fused anthers into a globose or cylindrical body with the following synadrium: Synandrium (sub)sessile in Gymnacranthera and Horsfieldia Synandrium elongate, stalked, with a prolonged sterile apex in Myristica Synandrium globose, stalked, without a sterile apex in Endocomia and Paramyristica

The aril is entire or divided into narrow segments: Aril entire or aril only divided at the apex in Horsfieldia and Knema Aril divided to ca one third or halfway from the apex in Endocomia Aril divided to the base in Gymnacranthera, Myristica, and Paramyristica

Features of the Family Typical for the Myristicaceae are the monopodial growth form (model of Massart) with more or less tiered lateral side branches; leaves simple, pinnate, exstipulate, mostly distichously arranged, the twigs ending in a cylindrical leaf-bud; flowers simple, with a uniserate perianth (2- or) 3- or 4-lobed (no sepals or petals), uni-

Figure 3.6.9. Schematic drawings of the androecium of: a. Endocomia; b. Gymnacranthera; c. Horsfieldia; d. Knema; e. Myristica; f. Paramyristica. Drawing: Jan van Os.

................. 16157$

CH27

02-08-07 10:32:06

PS PAGE 409

. .

410 /

.

sexual (mostly dioecious). The large fruit opens lengthwise with a long slit exposing one showy arillate seed. The endosperm is ruminate. The slashed bark exudes a red sap. The precise details of the indumentum and whether the leaves are punctuate underneath or not, have great value in species delimitation in all genera. Habit: dioecious or monoecious (Endocomia) monopodial shrubs or trees with aromatic oil-cells, (2–)5–15(–40) m high (Virola sessilis, Brazil, 30 m high); stiltroots sometimes present; branching whorled, the crown often pyramidal. Bark and wood exuding a watery pink or red sap on cutting. Wood white or soon brownish on cutting, soft; rays narrow, tanniferous tubes present; growth rings usually distinct and visible to the naked eye. Indumentum: twigs and leaves hairy, glabrescent, hairs unseriate, often with armed cells, appearing as stellate or dendroid. Stipules: absent. Leaves: simple, entire, alternate, usually distichous in plagiotropic shoots, indumentum on lower surface remaining or not; lateral nerves distinct, interarching at the margin; reticulation sometimes faint; vernation conduplicate (Asian genera); dark dots (corky hair bases) on lower surface present or absent; glands absent. Male inflorescences: from foliar axils to ramiflorous (cauline), pedunculate, compound or not, with or without bracts at base, the flowers variably arranged at the end of the branches, or a short, woody (sub)sessile simple or 2–4fid scar-covered wart- or worm-like brachyblast with the flowers in (sub)umbels at the end. Bracts: caducous. Female inflorescences: similar to male, usually less branched. Flowers: unisexual, mostly pedicellate; bracteole present or absent. Perianth: gamophyllous, globose to rotate, buds 2-lobed (Horsfieldia, p.p.) to 5-lobed (Endocomia, p.p.), cleft to variable depths, lobes little or much recurved; female flowers usually urceolate, larger, more swollen than the male; disk rarely present. Androecium with synandrium sessile or stalked (androphore), in Myristica mostly with a short sterile apex; anthers dorsally adnate to the column and laterally to each other or free, in Knema 2–45 in number stellately arranged around a disc, ellipsoid to linear, extrorse, opening by longitudinal slits, 2-loculed, tetrasporangiate; immature pollen sacs septate. Ovary monocarpellate, sessile, superior, ovoid; style absent or short, stigma 2-lobed or rarely peltate with few to many laciniations; ovule one, mostly anatropous, basal. Fruits (sub)globose to oblong, rarely transversely elongate, dehiscing by a longitudinal circumferential suture into two halves; pericarp leathery, carnose or subligneous, glabrous or hairy. Aril orange or red, completely covering the seed, divided into narrow segments to varying depths or entire, attached to the base of the seed. Seeds with a testa of three layers; albumen hard, ruminate, containing oil, sometimes starch. Embryo small, near the base of the seed or a little above; cotyledons connate at base (peltate or cup-shaped) or free; radicle basal, cylindric.

Overview of Genera A widespread genus with four species in Southeast Asia, one of which, E. macrocoma subsp. prainii, is found in New Guinea. It is unique in that the seeds are

................. 16157$

CH27

02-08-07 10:32:08

PS

PAGE 410

Angiosperms: Myristicaceae of Papua / 411

variegated, as in seeds of Ricinus or Hevea (Euphorbiaceae). Endocomia is largely monoecious, with in each paniculate inflorescence many male flowers mixed with few female flowers. The perianth is hairy inside. The aril is dissected to about halfway.

A genus of seven species, of which one species, G. paniculata, is widespread in New Guinea. Fruits are smallish, with the seed aril deeply divided as in Myristica, but the fruits are several together in a paniculate infructescence.

This genus is widespread in the whole Malesian area, with some 100 species, of which about 30 are found in New Guinea. Those east of Wallace’s Line (including most New Guinea species) have a predominantly 2-lobed perianth (section Irya). The majority of species are rather restricted in their distribution or subendemic. One of the most common species is H. irya, a riverine species found from Sri Lanka to the Solomon Islands and characterized by globose fruit. Its seeds have a cavity that facilitates floating, possibly the reason for its wide distribution. A salient group of five species comprises particularly small undershrubs or mini-trees reaching only ca 1 m in height. They have a particular club-shaped perianth and a club-shaped androecium, with the anthers arranged in a star at apex (e.g., H. crux-melitensis). H. sylvestris may grow into a beautiful tall tree with pendulous branches, and when planted solitarily it has a potential as an ornamental.

A genus of nearly 100, mostly Malesian, species. Only one species, K. tomentella, reaches Papua (the western Vogelkop).

This is the most prominent genus of New Guinean Myristicaceae. In Asia it is the most widespread genus with some 175 species, distributed from south India eastward to far into the Pacific (Fiji). Some hundred species occur in New Guinea, including comparatively many endemics. The genus is publicly well known through the cultivated nutmeg of commerce (M. fragrans). Nutmeg (pala) is now known only from cultivation on a commercial scale in the Moluccas and West Indies, but originally it was a very local endemic of the nearby Banda Islands, Moluccas, west of Papua in the Banda Sea. Formerly a second species, M. argentea, from western Papua, was commercially cultivated for spice as well, but at present it is used only locally. New Guinean Myristica exhibits substantial morphological diversity, much more than all other Myristica species outside the area. During recent revisions by de Wilde many new species were discovered among the abundant new herbarium accessions of the last decades. It is likely that more exploration will yield more as yet undescribed species.

................. 16157$

CH27

02-08-07 10:32:08

PS

PAGE 411

. .

412 /

.

It has proved to be useful to divide the New Guinean species into groups and recognize them according to: (1) leaf size (i.e., those with leaves smaller or larger than ca 15 cm length); (2) the presence or absence of dark dots on the lower leaf surface; (3) male inflorescences, either peduncled or not; and (4) the presence of swollen (ant inhabited) parts in the (winged) twigs or ‘‘ant swellings.’’ Various characters of the male flower (bud) and fruit, including size and characteristics of the indumentum, ultimately define the species. Several distinct groups can be defined on technical characters of the flowers. A few small-leaved species have conspicuously yellow-green leaves on drying, similar to the well-known green-drying leaves of species of Symplocos (Symplocaceae) or Xanthophyllum (Polygalaceae) in which aluminum accumulation is known. These Myristica species are possibly restricted to ultramafic soils, but nothing is known with certainty.

A monotypic New Guinean genus whose morphology is clearly distinct from Myristica. However, in phylogenetic analysis, using molecular data and pollen characters, it appears nested within Myristica (Sauquet et al. 2003). Its pollen is identical to that of Myristica. Paramyristica could be regarded as an advanced ingroup in Myristica. However, for practical reasons, it cannot be included formally because it would severely blur the well established genus delimitation on current morphological criteria. The area of the species concerned is in northern Papua New Guinea approaching Papua. Additional collecting for further study is urgently needed.

Gaps in Floristic Documentation or Knowledge As with most plant groups, the whole New Guinea region is still undercollected. In Myristicaceae the recent revision of the New Guinean material yielded a comparatively large number of new taxa (ca 80), most of them known by one or a few collections, which indicates that more new species will be discovered if new collections are made.

Natural History Myristicaceae are found in all types of forest. Sometimes they constitute a considerable component of the forest, especially in the midstory of lowland rainforest, but they are not gregarious. Some species (notably some Horsfieldias) are ‘‘sciaphilous nomads’’ (i.e., fast growing, shade tolerant) and a few of these are found in secondary forest.

................. 16157$

CH27

02-08-07 10:32:09

PS

PAGE 412

Angiosperms: Myristicaceae of Papua / 413

Flowering and fruiting generally occur throughout the year. The usually carnose, yellow or brown, inside creamy pink or red flowers of several general have repeatedly been reported as being fragrant (e.g., Horsfieldia irya and Myristica fragrans). Anthesis is presumably mainly nocturnal, and small beetles may effect pollination, a plausible supposition given the tough structure of the perianth. Nectar is not reported for any species. Male plants of Myristica produce over 50 times as many flowers than do females (Armstrong and Drummond 1986; Armstrong and Tucker 1986; Armstrong and Irvine 1989a,b). Myristica subalulata and some related species are myrmecophilous, the ants possibly involved in pollination (De Wilde 1998). In addition to the colored inside of the perianth, in Knema the staminal disc may be bright purple-red. The contrast with the creamy-white pollen possibly heightens the attraction of pollinators in this genus.

The brown-black seeds that contrast sharply with the orange or red aril and the inner surface of the (red, pink or white) opened pericarp, attract birds and suggest dispersal by fruit pigeons, hornbills, and birds of paradise (see Beehler and Dumbacher 1996). Rodents may disperse seeds as well. Dissemination by water may occur in Horsfieldia irya (Figure 3.6.10), a widespread riverine species with cavities in the endosperm that appear to increase buoyancy of seeds. The seeds of the marshy nutmeg, Myristica elliptica, are also reported to float when the aril is removed.

Figure 3.6.10. Photo of Horsfieldia irya (Gaertn.) Warb. Photo: W.J.J.O. de Wilde.

................. 16157$

CH27

02-08-07 10:32:16

PS

PAGE 413

. .

414 /

.

Figure 3.6.11. Myristica subalulata Miq. var. subalulata. Male twig with immature flowers, ant swelling and ant-openings; stem partly opened to show ant cavities with two coccids. After Polak MP 750; drawn by Jan van Os. Scale bar

................. 16157$

CH27

2 cm.

02-08-07 10:33:08

PS

PAGE 414

Angiosperms: Myristicaceae of Papua / 415

Seeds remain viable for a restricted period (i.e., a few weeks) and germinate only in a damp, shady environment. Therefore the natural reintegration of Myristicaceae in secondary forest is impossible. Germination is (mostly) hypogeal. The cotyledons remain within the testa, the taproot and hypocotyls emerge. The shoot is erect, initially with reduced leaves (cataphylls), borne spirally (the Horsfieldia type and subtype (de Vogel, 1979), a common type in tropical woody dicotyledons).

Myrmecophily in Myristicaceae is known only for Myristica species in New Guinea, where it is quite apparent (de Wilde 1998). Most likely the relationship with the ants is symbiotic and apparently specific. Whether the typical ant swellings in the twigs are also formed when ants are absent is not known. Within the swellings in Myristica subalulata bugs (coccids) are always found, obviously an essential element of the symbiosis (Fig. 3.6.11).

Literature Cited Armstrong, J.E., and B.A. Drummond. 1986. Floral biology of Myristica fragrans Houtt. (Myristicaceae), the nutmeg of commerce. Biotropica 18: 32–38. Armstrong, J.E., and A.K. Irvine. 1989a. Floral biology of Myristica insipida (Myristicaceae), a distinctive beetle pollination syndrome. Am J Bot 76: 86–94. Armstrong, J.E., and A.K. Irvine. 1989b. Flowering, sex ratios, pollen-ovule ratios, fruit set, and reproductive effort of a dioecious tree, Myristica insipida (Myristicaceae), in two different rain forest communities. Am J Bot 76: 74–85. Armstrong, J.E., and S.C. Tucker. 1986. Floral development in Myristica (Myristicaceae). Am J Bot 73: 1131–1143. Beehler, B.M., and J.P. Dumbacher. 1996. More examples of fruiting trees visited predominantly by birds of paradise. Emu 96: 81–88. de Vogel, E.F. 1979. Seedlings of Dicotyledones. Pudoc, Wageningen. de Wilde, W.J.J.O. 1991. The genera of Myristicaceae as distinguished by their inflorescences, and the description of a new genus, Bicuiba. Beitr Biol Pflanzen 66: 95–125. de Wilde, W.J.J.O. 1998. The Myrmecophilous species of Myristica (Myristicaceae) from New Guinea. Blumea 43: 165–182. de Wilde, W.J.J.O. 2000. Myristicaceae. Flora Malesiana ser. I, 14: 1–634. Ku¨hn, U., and K. Kubitzki. 1993. Myristicaceae. Pp. 457–467 in Kubitzki, K., J.G. Rohwer, and V. Bittrich (eds.) The Families and Genera of Vascular Plants 2. Springer-Verlag, Berlin. Sauquet, H., J.A. Doyle, T. Scharaschkin, T. Borsch, K.W. Hilu, L.W. Chatrou, and A. Le Thomas. 2003. Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution. Bot J Linnean Soc 142: 125–186.

................. 16157$

CH27

02-08-07 10:33:09

PS

PAGE 415

Myrsinaceae of Papua . , the Myrsinaceae (coral berry or Myrsine family, including Maesaceae) in the narrowest sense contain 35–41 genera and over 2,000 species, with over 100 as yet undescribed (Chen and Pipoly 1996). Here, the Maesaceae (A. DC.) (Anderberg, Sta˚hl, and Ka¨llersjo¨ 2000) are included, but only as a matter of convenience, as I do agree with the segregation of Maesa into its own family. In the neotropics, there are approximately 14 currently recognized genera (Pipoly and Ricketson, in press) with two more as yet undescribed, while in the Paleotropics, there are 21–24 genera, with possibly one new genus. The genera Myrsine (including Suttonia and Rapanea), Ardisia, and Hymenandra are pantropical. In New Guinea there are 11 genera and approximately 131 species that do not include another 30 to be described when better collections are obtained.

W

Distribution and Habitat The Myrsinaceae occur in subtropical and tropical forests worldwide. In New Guinea and the Bismarck Archipelago, they occur in mangroves, coastal forest, swamp forest, mixed dipterocarp forest, lowland humid, wet, and rainforest, submontane, montane, and cloud forest, and in montane scrub and mossy forests. Only the smallest subshrubs occur in the middle of open forest; the majority of woody species occur at the margins of habitats, near rocks in areas with high humidity and just above the flood line along watercourses. Members of the genus Maesa are gap species, found frequently in recovering landslides and treefall areas. The greatest diversity in Myrsinaceae species is found in the submontane to montane habitats.

Family Classification Since the worldwide monograph of Mez (1902), no critical evaluation of the family has been presented until it was redelimited by Sta˚hl and Anderberg (2003), when the Maesaceae was segregated, and ten genera traditionally included in the Primulaceae were added to the Myrsinaceae. Calculations for family size were made by Pipoly and Stone and reported in Pipoly and Chen (1996) as ca 42 genera and 2,200 species. These calculations included information provided by all researchers working on the family, and took into account the many as yet unpublished new species and genera. For the taxa in New Guinea, Mez (1902, 1922) provided the most complete account, with significant additions by Moore (1916), Kanehira and Hatusima (1943), and a partial treatment in van Royen (1982). A series of summaries by Sleumer (1986, 1987a,b, 1988a,b,c) did much to assemble herbarium mateMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

416

................. 16157$

CH28

02-08-07 10:33:32

PS

PAGE 416

Angiosperms: Myrsinaceae of Papua / 417

rials, bibliography, and working treatments, but did not solve the problems caused by lack of fieldwork. Perusal of his writings indicate that few collections are known from Papua outside of the Vogelkop Peninsula, while in PNG few collections were made outside Morobe, Eastern Highlands, New Britain, Madang, and the Sepik, leaving large expanses without collections. Subsequent work by Stone (1988, 1990, 1991a,b, 1994) in Malesia necessarily treated some of the New Guinean taxa and their floristically-related sisters from Sulawesi and the Solomon Islands, while pointing out the many voids in the state of our knowledge, such as the absence of any records for the Indo-Malesian ‘‘boreotropical’’ genus Hymenandra from New Guinea (Stone 1991b). Given the dearth of our knowledge of the family in New Guinea, a practical key, based on the most obvious characters, is provided, as are short summaries for each genus.

Key to the Genera of Myrsinaceae in New Guinea 1. Ovary and fruit semi-inferior to inferior; calyx subtended by two bracteoles; seeds numerous, angular. .............................................................................Maesa 1. Ovary and fruit superior, calyx and pedicel without bracteoles; seeds single, globose or elongated and curved. ........................................................................ 2 2. Lianas, branches plagiotropic, leaves distichous, petals free or rarely partly fused. ...................................................................................................... Embelia 2. Herbs, shrubs, or trees; branches orthotropic or mixed, leaves spiral, petals fused. ................................................................................................................. 3 3. Mangrove trees; anthers versatile, transversely septate; fruit curved and elongate, viviparous. ........................................................................Aegiceras 3. Herbs, shrubs, and trees, not of mangroves; anthers basifixed or dorsifixed but not versatile, not transversely septate; fruit a globose, depressed-globose, ellipsoid, obovoid or obloid drupe, not viviparous. 4 4. Herbs or suffrutices with succulent stems; petals conduplicate, essentially concealing the anthers. ...............................................Labisia 4. Woody shrubs or trees, the stems not succulent; petals planar or reflexed, not concealing the anthers. .................................................... 5 5. Inflorescence shorter than the petioles, fasciculate or umbellate, sessile or on short shoots formed by persistent peduncles girdled by persistent floral bracts thus forming ‘‘short shoots’’; stigma subcapitate or sausage-like. .................................................. Myrsi ne 5. Inflorescence longer than the petioles, paniculate or racemose, not on ‘‘short shoots’’; stigma punctiform. ........................................... 6 6. Branches bearing prominent cataphylls, in spirals alternate with the leaves; inflorescence on special leafless branchlets, or axillary to the leaves, the branchlets or peduncles with reduced leaves transitional to large caducous bracts. ............................... Loheria 6. Branches without prominent cataphylls; inflorescences axillary but peduncles without reduced leaves transitional to bracts; inflorescence bracts minute. ........................................................ 7

................. 16157$

CH28

02-08-07 10:33:32

PS

PAGE 417

.

418 /

7. Corolla lobes contorted in bud, rarely dextrorsely imbricate; anthers punctate on the thecae and on apiculum when present. ..................................................................................... 8 8. Pedicels discrete, not swollen apically into the calyx; calyx campanulate or cotyliform; anthers basifixed, deshiscent by longitudinal slits. ........................................................... 9 9. Flowers bisexual; calyx campanulate, the lobes shortconnate; filaments free, short, the anthers longer than wide, connate at least basally and connivent to the apex, united over the ovary at anthesis; stigma punctate, the ovules numerous; pluriseriate, buried in the placenta. .................................................... Connandrium 9. Flowers unisexual; calyx cotyliform, the lobes connate nearly 2/3 their length; filaments basally united into an inconspicuous tube, the anthers free, as wide or wider than long; stigma obviously peltate to discoid; ovules few, uniseriate, exposed over most of their surface. ...... .......................................................................... Discocalyx 8. Pedicels distally swollen into the calyx; calyx prominently cupuliform; anthers dorsifixed near base, dehiscent by longitudinal slits starting as apical pores. ...Tapeinosperma 7. Corolla lobes imbricate in bud (including quincuncial); anthers punctate only on the connective, never on apiculum when present. ......................................................................... 10 10. Flowers unisexual; corolla campanulate to urceolate; stigma capitate to discoid; fruit with exocarp thick, spongy, often with a crest. ....................................Fittingia 10. Flowers bisexual; corolla rotate; stigma subulate and punctate; fruit with exocarp thin, not spongy. .....Ardisia

Features of the Family Habit: trees, shrubs, climbers (or rarely herbs). Stipules absent. Leaves simple, alternate, distichous, rarely opposite or whorled, venation pinnate, often glandular. Inflorescences terminal, axillary, or at apices of lateral branches, racemose (often paniculate), spicate, corymbose, cymose, umbellate, or fasciculate on scaly spur branches in leaf axils; the pedicels subtended by a floral bract or rarely (Maesa) with prophylls just below the calyx. Flowers bisexual or unisexual, the plants polygamodioecious, androdioecious, or dioecious, 4- or 5-(or 6)-merous, actinomorphic. Sepals basally connate or free, imbricate, quincuncial, contorted or valvate in bud, persistent, usually glandular. Petals basally connate or rarely free, imbricate, quincuncial, contorted or valvate in bud, usually glandular. Stamens as many as and opposite to petals, usually adnate to base or throat of corolla tube, sometimes free (Ardisia), often united into a tube with or without alternating

................. 16157$

CH28

02-08-07 10:33:33

PS

PAGE 418

Angiosperms: Myrsinaceae of Papua / 419

lobes on the tube; anthers 2-celled, dorsifixed or basifixed, dehiscing introrsely or laetrorsely by longitudinal slits or apical pores, or by apical pores opening into longitudinal slits, or rarely transversely septate, the filaments obviously present or absent. Ovary superior, rarely inferior to half-inferior (Maesa), 1-celled; placentation free-central, sometimes basal; ovules one to several in one to many rows, usually embedded in placenta, anatropous or semicampylotropous. Style one; stigma simple or lobed, punctate, discoid, morchelliform, or capitate. Fruit drupes with fleshy, rarely a spongy exocarp, one- or many-seeded capsules. Seeds one to many; endosperm fleshy or horny; embryo x 10–13, 23.

Genera of Myrsinaceae (sensu lato) in New Guinea and the Bismarck Archipelago Shrubs or small trees. Leaves alternate or subopposite. Inflorescences terminal or rarely axillary, umbellate. Flowers bisexual, 5-merous. Corolla campanulate, united into a tube; lobes ovate or ovate-lanceolate, imbricate, overlapping to right in bud, recurved or reflexed at anthesis, not glandular. Basal part of filaments united into a tube as long as corolla tube, distal part free, exserted; anthers ovate, two-celled, dehiscing longitudinally, transversely septate. Ovary superior; ovules numerous, within a globose placenta. Style elongated; stigma apiculate. Fruit elongated, terete, curved, one-seeded capsules; exocarp dry, crustaceous, dehiscing by a longitudinal fissure or separating into two fragments along back and front; endocarp somewhat fleshy; persistent calyx compactly enclosing fruit base. Seeds occupying whole cavity; embryo terete, curved. Two species: India, Malesia, China, and New Guinea. Aegiceras corniculatum (Linnaeus) Blanco is the most common, with Aegiceras floridum Roemer & Schultes reported from the northwest section of the island by Kaneheira and Hatusima (1943).

Trees, shrubs, suffruticesent (or rarely herbs). Leaves alternate or pseudoverticillate, usually punctate or punctate-lineate. Inflorescences paniculate, cymose, corymbose, or umbellate, rarely racemose. Flowers bisexual, often punctate, 5- or rarely 4-merous. Calyx campanulate or cupular; sepals free or barely united at base, imbricate or quincuncial, usually punctate or punctate-lineate. Corolla campanulate, often punctate; lobes united at base, overlapping to right or very rarely to left, imbricate, or quincuncial, often conical in bud. Stamens attached at base or middle of corolla tube; filaments long or very short, broad at base; anthers free, dehiscing longitudinally or by apical pores. Ovary ovoid or subglobose, ovules three to many, pluriseriate; style base persistent; stigma minute, apiculate. Fruit drupaceous, one-seeded, punctate, sometimes longitudinally ribbed, with somewhat fleshy exocarp and crusty or slightly bony endocarp. Seeds covered by membranous remnants of placenta, the endosperm smooth or ruminate. About 400–500 species: primarily tropical east and Southeast Asia, Americas, Australia, and Pacific

................. 16157$

CH28

02-08-07 10:33:33

PS

PAGE 419

.

420 /

Islands; in New Guinea, Sleumer (1988a) reported ca 36 species, of which five are incompletely known and several remain undescribed. A number of new species were described for New Guinea by Kaneheira and Hatusima (1943). It is interesting to note that the New Guinea species of subgenus Pimelandra are shrubs or small trees, whereas in western Malesia, they are large trees. The best developed subgenera are Tinus and Tinopsis, two subgenera that otherwise have their centers of diversity in the Philippines.

Shrubs or small trees. Leaves alternate, petiolate, prominently punctate and/or punctate-lineate. Inflorescences axillary panicles or racemes with long, slender pedicels. Flowers bisexual, 5-merous. Calyx campanulate; sepals dextrorsely contorted, barely united at base, densely punctate. Corolla broadly rotate, petals dextrorsely contorted, asymmetrical, broadly conical in bud. Stamens attached at base of tube, the filaments free, very short, terete; anthers basifixed, connate at least basally connate, introrsely longitudinally dehiscent. Ovary obturbinate, ovules many, 2–3-seriate; style base persistent; stigma punctate to discoid. Fruit drupaceous, one-seeded, densely and prominently punctate, globose, without ribs, the exocarp thin, the endocarp crusty. The genus, with two currently recognized species and perhaps a third undescribed, is endemic to New Guinea, the Moluccas, and the Bismarck Archipelago. The genus was treated by Mez (1902) and Sleumer (1988b).

Shrubs or small trees. Leaves alternate, long or rarely short-petiolate but always with pulvinus, prominently punctate and/or punctate-lineate. Inflorescences axillary racemes or panicles of racemes. Flowers unisexual, 4–5-merous. Calyx cotyliform, the lobes imbricate in bud, connate nearly two-thirds their length; corolla urceolate, campanulate or rotate, the lobes dextrorsely contorted in bud, variously connate; staminate flowers with filaments basally united into an inconspicuous tube, the anthers basifixed, free, as wide or wider than long, dehiscent by longitudinal slits; pistillate flowers with stamens reduced to fleshy protruberances; staminate flowers with ovary rudimentary, conic, hollow or with a few abortive ovules; pistillate flowers with ovary obturbinate, the ovules few, uniseriate, exposed, the style distinct but often short, the stigma obviously peltate to discoid. Fruit mostly obloid, wider than long, or rarely subglobose, the exocarp thin. The genus has been treated by Mez (1902) and by Sleumer (1988c) for New Guinea, but A. C. Smith (1973) provides important insights into Discocalyx and its relationship to Tapeinosperma. New species have also been described by Pipoly and Takeuchi (2004), but a revision of this genus, with Tapeinosperma, is urgently needed. There are currently 13 species recognized from New Guinea, with an estimated three other entities as yet undescribed owing to lack of suitable specimens.

................. 16157$

CH28

02-08-07 10:33:33

PS

PAGE 420

Angiosperms: Myrsinaceae of Papua / 421

. .

(

)

Shrubs, becoming lianous at maturity, or lianas. Leaves alternate, distichous, the branches plagiotropic. Inflorescences axillary or terminal, [paniculate, racemose] umbellate or corymbose, bracteate at base. Pedicel subtended by one persistent floral bract. Flowers 4- or 5-merous, bisexual or functionally unisexual and plants dioecious, polygamous, or monoecious. Sepals united at base. Petals free or united at base, glandular granulose inside and often along margin. Stamens adnate to base of petals, rarely free, normally exserted in staminate flowers, reduced in pistillate ones; filaments free; anthers dehiscing longitudinally, connectives usually punctate abaxially. Pistillode reduced and style included in staminate flowers; ovary globose or ovoid; ovules ca four, uniseriate. Style exserted; stigma discoid or capitate, sometimes slightly lobed. Fruit drupaceous, prominently punctate, 1-seeded; endocarp crusty or rarely bony. Seeds subglobose, covered by membranous remnants of placenta; the endosperm ruminate; embryo terete, transverse. About 140 species: Africa, Southeast Asia, Malesia, Australia, and the Pacific Islands. Sleumer (1987) lists ten species in New Guinea, but our data show there are perhaps an additional three undescribed species. Kanehira and Hatusima (1943) described several new species. Sleumer (1987) as well as Sta˚hl and Anderberg (2003) treated Grenacheria as distinct from Embelia, following Mez (1902), based on the slight fusion of the corolla lobes in the former, versus the free petals in the latter. However, the author has seen that there is greater fusion in the pistillate flowers and nearly free petals in the staminate ones, thus making their separation impractical. It is notable that the same corolla sexual dimorphism has been observed in numerous species of Embelia’s sister genus from the neotropics, Cybianthus (Pipoly 1987, 1992).

(

)

For illustration, see Figure 3.6.12. Few-branched shrubs or monoaxial trees. Leaves alternate to pseudoverticillate, rarely long-petiolate but never with pulvinus. Inflorescences axillary, long-pedunculate, racemose, the bracts small, without bracteoles. Flowers unisexual, 4- or 5-merous. Calyx campanulate, deeply lobed, the lobes imbricate in bud. Corolla campanulate to urceolate, the lobes imbricate; stamens in staminate flowers on short, free filaments basally adnate to the corolla but free from each other, the anthers ellipsoid to subsagittoid, basifixed or rarely dorsifixed above the base, the connective punctate; staminodes resembling stamens but reduced in size; pistillode in staminate flowers stipitiform, hollow; pistil obturbinate, the style short, the stigma capitate to discoid or peltate; the ovules unknown. Fruit with exocarp thick, spongy, often with a crest; seed with smooth endosperm. Sleumer (1988c) recognized five species, and lectotypified the genus, endemic to New Guinea and perhaps New Britain of the Bismarck Archipelago. Stone (1994) posthumously described an additional species, Fittingia mariae, based on Mary Clemens’s collections from Morobe Province.

................. 16157$

CH28

02-08-07 10:33:35

PS

PAGE 421

.

422 /

Figure 3.6.12. Fittingia tubiflora in its characteristic understory habitat. White arrow: the pendulous infructescence.

Perennial herbs or monoaxial subshrubs. Leaves long- or short-petiolate, but never with a pulvinus, strictly spiral. Inflorescences axillary, a columnar panicle with reduced side branches. Flowers 5-merous, bisexual. Calyx cotyliform, the lobes valvate. Corolla campanulate, the lobes valvate, short connate basally, conduplicate and concealing the anthers; stamens basally connate by their filaments, the anthers sessile on the staminal tube, oblong to narrowly ovate, dehiscent by broad longitudinal slits, the connective longer than the thecae, forming a long apiculum, densely punctate and punctate-lineate basally; pistil obturbinate, the style long, thin, the stigma punctate, ovules few, uniseriate, exposed on the placenta. Fruit globose, densely punctate, the exocarp thin; seed one, with smooth, crusty endosperm. Stone (1988, 1989) lists six species of which only one, Labisia sessilifolia, is known from New Guinea.

Small monoaxial or rarely few-branched trees. Branches bearing prominent cataphylls, in spirals alternate with the leaves. Leaves short-petiolate or subsessile, often large, pseudoverticillate, often with a pulvinus. Inflorescences on special branchlets, or axillary to the leaves, the branchlets or peduncles with reduced leaves transitional to large caducous bracts, the flowers arranged in racemes or spikes or rarely (subgenus Longicorona Stone) in globose corymbs, long-peduncu-

................. 16157$

CH28

02-08-07 10:34:07

PS

PAGE 422

Angiosperms: Myrsinaceae of Papua / 423

late. Flowers (3–)4- or 5-merous; functionally unisexual the plants dioecious. Calyx cotyliform to campanulate, the lobes nearly free, dextrorsely imbricate or quincuncial in bud, prominently punctate. Corolla rotate or slightly tubular, the tube very short, the lobes dextrorsely imbricate or quincuncial in bud; stamens on long, free filaments nearly free and adnate only at base, the anthers basifixed, mostly broader than long, dehiscent by latrorse longitudinal slits; staminodes similar to stamens but reduced in size; pistil obturbinate with a stout style, the stigma discoid to capitate stigma, the ovules few, uniseriate, exposed; pistillode hollow, conic when present or absent. Fruit subglobose, the exocarp thin, the endocarp thin, smooth, the endosperm ruminate or smooth. A Malesian genus of six species, with two endemic to New Guinea, last revised by Stone (1991a). The entire genus is exceedingly rare, as it is known from only 35 collections. Much more study of the group will be necessary to better determine its affinities.

˚ Shrubs or rarely small trees. Leaves entire or dentate, usually glandular. Inflorescences axillary or terminal, racemose or paniculate; pedicels subtended by a brach and bearing two prophylls just below the calyx, termed bracteoles. Flowers (4–)5(–6)-merous, bisexual or polygamous, perigynous. Calyx funnelform, adnate to ovary, lobes persistent, valvatein bud. Corolla white or yellowish, campanulate, rarely urceolate, the lobes imbricate or quincuncial in bud, often longitudinally glandular striate. Stamens distinct, included or exserted, the filaments free, adnate to the corolla tube, the anthers as wide or wider than long, introrsely dehiscent by longitudinal slits; Pistil semi-inferior or inferior, aborted in staminate flowers; ovules numerous, on a globose free-central placenta; the style as long as or longer than stamens, the stigma entire or 3–5-lobed. Fruit a globose or ovoid indehiscent berry or drupe with a crusty endocarp, apex covered by persistent calyx, often longitudinally glandular striate. Seeds small, numerous, angular, embedded in a hollow placenta. About 200 species of the paleotropics; 27 species recognized in New Guinea. Sleumer (1987) provided a revision of Maesa for New Guinea, the Moluccas, and the Solomon Islands, which may be the most natural geographic area; this was followed by two papers describing additional species (Utteridge 2000, 2001). This large and complicated genus requires a complete revision throughout its range. Once again, specimen citations in the Sleumer treatments point to large portions of New Guinea for which no specimens of this genus, frequent in forest gaps and along margins, have been collected.

(

)

Shrubs or small trees. Leaves alternate. Inflorescences axillary, lateral, umbellate or fasciculate, sessile or on short, perennating peduncles girdled by persistent floral bracts (thus forming ‘‘short shoots’’). Flowers 4–5(–6)-merous, bisexual or unisexual (plants then monoecious, dioecious or polygamous); sepals nearly free or united to length, imbricate or valvate, usually ciliate, punctate, persistent;

................. 16157$

CH28

02-08-07 10:34:07

PS

PAGE 423

.

424 /

petals nearly free or rarely united to their length, usually ciliate, glandulargranulose at least along margin and often throughout, punctate. Stamens and staminodes similar, subequalling corolla length, the filaments free or connate basally to form a tube, the tube with or without serile appendages alternating with the filaments and all merely adnate to the corolla tube, or developmentally fused throughout, the anthers thus appearing epipetalous, the anthers ovate or reniform, rarely sagittate, two-celled, dehiscing by introrse longitudinal slits, rarely by subterminal pores opening later into wide longitudinal slits. Pistil and pistillode similar; obconic, obturinate, obnapiform or variously subglobose; ovary globose, costate or not, ovules few, uniseriate or rarely biseriate, completely immersed in the placenta or seated below apical pores in placenta or variously projecting; style obsolete to present, tapering into stigma; stigma morchelliform, liguliform, sinutate to lobate, prismatic and three-lobed, or rarely, conical. Fruit a globose, subglobose, ellipsoid, ovoid or subovoid drupe with a somewhat fleshy exocarp and crusty or leathery endocarp, one-seeded. Seeds occupying cavity; endosperm horny, ruminate; embryo cylindrical, transverse. The genus, as here delimited, is pantropical, and contains over 300 species of which many remain undescribed (Pipoly 1996). Aside from the comprehensive treatment of Mez (1902), Kanehira and Hatusima (1943) described a number of new species, as did van Royen (1982) from the montane regions. The latest treatment of the genus (as Rapanea) was by Sleumer (1986) in which he recognized 22 species. However, a study currently underway has revealed that there may be as many as four new entities, and that perhaps six of the 22 species recognized heretofore might be reduced to synonymy, leaving a total of 20 for the island, and an additional three if the Bismarck Archipelago and Bougainville Island are included. Because of its dioecy and its minute flowers, the genus needs more thorough study.

.

.

Monaxial trees or rarely few-branched shrubs. Leaves pseudoverticillate, shortor long-petiolate, but always with a pulvinus. Inflorescences axillary or terminal, paniculate with the flowers in corymbs. Flowers mostly 5-merous, bisexual, calyx cupuliform, united up to their length, the lobes imbricate in bud, corolla rotate or campanulate, the lobes dextrorsely contorted in bud; stamens with filaments basally connate and adnate to the corolla tube, the anthers free, oblong-ovoid, dorsifixed near the base, opening by longitudinal slits starting as subapical pores; ovary globose or ovoid, style thick, longer than the ovary, the stigma truncate, capitate or somewhat discoid; ovules few, uniseriate. Fruits large, mostly depressed-globose to obloid, the exocarp thick and juicy but not spongy, endocarp crusty, one-seeded. Seeds with endosperm smooth or ruminate. A genus of over 60 species, with its center of diversity in Fiji, New Caledonia, and Vanuatu, but with others in Borneo, New Guinea, and new taxa in the Philippines and eastern Malesia. There are currently five species recognized from New Guinea, including

................. 16157$

CH28

02-08-07 10:34:08

PS

PAGE 424

Angiosperms: Myrsinaceae of Papua / 425

those treated by Sleumer (1988c) and Pipoly and Takeuchi (2004; Figures 3.6.13, 14).

Gaps in Floristic Documentation or Knowledge Despite a broad overview for conservation needs provided by Swartzendruber (1993), in which 16 terrestrial habitats in Papua New Guinea were highlighted as ‘‘major terrestrial unknowns,’’ little has been done to solve the problem because of the remoteness of many areas and lack of adequate numbers of trained incountry plant systematicists, and experts in the related fields of dendrology and forest mensuration. While the PNG National Herbarium (LAE) is one of the few national herbaria managed by a forest department, funding for major expeditions has been lacking, and expeditions are more difficult to implement because of aspects of the land tenure system. Outside of Papua New Guinea, progress on the Papua (Irian Jaya) side has been made through several floristic programs, but is moving at a very slow pace. The majority of species of the family are known from less than several dozen gatherings, save the upper montane taxa near roads, which have a reasonable number of collections, but taken from the same place. Collecting trips to remote areas accessible only by helicopters are needed to get a realistic estimate of the island’s diversity.

Natural History The Myrsinaceae are most speciose in the submontane habitat, and most common within a species in montane areas, where a species can form large populations. Observations from members of the family in cultivation among a variety of genera and geographic origins indicate that they are not particularly aggressive (except for some species of Ardisia subgenus Tinus). Myrsinaceae are normally slow to germinate (over two months), develop slowly into seedlings over 6–8 months, then begin to show rhythmic growth with sylleptic branching. Depending on the taxon, once flowering is triggered, each flower usually lasts less than two full days, and if pollinated, the fruit will take up to four months to produce. Consequently, many taxa are known only from fruiting specimens. Monoaxial taxa are most frequent in the lowlands, in alluvial flood plains, where they are capable of being submerged for short periods, or they are gap species in environments subject to periodic blow-downs, landslides, and other related events. Despite this resiliency, very few Myrsinaceae can withstand soil compaction. My fieldwork has shown that striking polymorphisms between seedlings, saplings, and even within trees (due to reiteration phenomena), concomitant with precocious flowering among some juveniles, make identifications difficult. Myrsinaceae wood ranges from brittle to strong. Among the monoaxial taxa, it is frequent to see rather thin-walled vessels and rays with variously shaped starch grains. Most members of the family are pollinated by small bees or dipterans, and

................. 16157$

CH28

02-08-07 10:34:08

PS

PAGE 425

.

426 /

Figure 3.6.13. Tapeinosperma magnifica, a recently described endemic from northeast New Guinea. White arrow: the polelike stem with umbelliform branching at the top.

................. 16157$

CH28

02-08-07 10:34:55

PS

PAGE 426

Angiosperms: Myrsinaceae of Papua / 427

Figure 3.6.14. Tapeinosperma magnifica. The flowers occasionally have ten wellformed and fertile stamens in otherwise normal flowers. are not very specific. Ardisia subgenus Stylardisia is defined by its protogynous behavior. Myrsinaceae are used worldwide for fish poison because of the triterpenoid saponins found in their lyso-schizogenous resin canals, punctations, and punctate lineations. The many punctations found on the plants also render them attractive for decorating community events, religious services, and so on, so the flowering branches are frequently harvested for those purposes.

Literature Cited Kaneheira, R., and S. Hatusima. 1943. The Kanehira and Hatusima 1940 Collection of New Guinea Plants XXI. Botanical Magazine (Tokyo) 57: 215–236. Mez, C. 1902. Myrsinaceae. Pp. 1–437 in Engler, A. (ed.) Das Pflanzenreich IV. 236 (Heft 9). Leipzig. Moore, S. 1916. Myrsinaceae. In Ridley, H.N., Report on the botany of the Wollaston Expedition to Dutch New Guinea, 1912–1913. Transactions of the Linnaean Society London, Botany 9: 1–269. Pipoly, J.J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia (Myrsinaceae). Mem. New York Bot. Gard. 43: 1 –76. Pipoly, J.J. 1992. The genus Cybianthus subgenus Conomorpha (Myrsinaceae) in Guayana. Ann. Missouri Bot. Gard. 79: 908–957.

................. 16157$

CH28

02-08-07 10:35:13

PS

PAGE 427

.

428 /

Pipoly, J.J. 1996. Contributions toward a new flora of the Philippines: I. A synopsis of the genus Myrsine (Myrsinaceae). Sida 17 (1): 115–162. Pipoly, J.J., and W. Takeuchi. 2004. New species of Tapeinosperma and Discocalyx (Myrsinaceae) from Morobe Province, Papua New Guinea. Harvard Papers in Botany 8: 153–159. Sleumer, H. 1986. A revision of the genus Rapanea Aubl. (Myrsinaceae) in New Guinea. Blumea 31: 245–269. Sleumer, H. 1987a. A revision of the genus Maesa Forsk. (Myrsinaceae) in New Guinea, the Moluccas, and the Solomon Islands. Blumea 32: 39–45. Sleumer, H. 1987b. The genera Embelia Burm. f. and Grenacheria Mez (Myrsinaceae) in New Guinea. Blumea 32: 385–396. Sleumer, H. 1988a. A revision of the genus Ardisia Sw. (Myrsinaceae) in New Guinea. Blumea 33: 115–140. Sleumer, H. 1988b. The genus Connandrium Mez (Myrsinaceae). Blumea 33: 109–113. Sleumer, H. 1988c. The genera Discocalyx Mez, Fittingia Mez, Loheria Merrill, and Tapeinosperma Hook f. in New Guinea. Blumea 33: 81–107. Smith, A.C. 1973. Studies of Pacific Island plants, XXV. Myrsinaceae of the Fijian Region. Journal of the Arnold Arboretum 54 (1): 1–40; 54 (2): 228–292. Sta˚hl, B., and A. Anderberg. 2003. Myrsinaceae. Pp. 266–281 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. VI. Flowering Plants: Dicotyledons. Springer, Berlin. Stone, B. 1988. Notes on the genus Labisia Lindl. (Myrsinaceae). Malayan Nature Journal 42: 43–51. Stone, B. 1990. Studies in Malesian Myrsinaceae, V. Additional new species of Ardisia Sw. Proceedings of the Academy of Natural Sciences of Philadelphia 142: 21–58. Stone, B. 1991a. The genus Loheria Merrill (Myrsinaceae). Micronesica 24: 65–80. Stone, B. 1991b. New and noteworthy Myrsinaceae, VI. Revision of the genus Hymenandra A. DC. The Garden’s Bulletin, Singapore 43: 1–17. Stone, B. 1994. New and noteworthy Malesian Myrsinaceae, VIII. Sida 16: 263–272. Utteridge, T. 2000. Two new species of Maesa (Myrsinaceae) from Puncak Jaya, New Guinea. Kew Bulletin 55: 443–449. Utteridge, T. 2001. Two new species of Maesa (Maesaceae) from New Guinea. Kew Bulletin 56: 677–683.

................. 16157$

CH28

02-08-07 10:35:14

PS

PAGE 428

Myrtaceae of Papua lyn a. craven Number of Genera and Species o r ld w i d e, the Myrtaceae comprises about 4,500 to 5,000 species and about 130 genera. In New Guinea there are 28 wild-occurring genera, one of which is represented by a single naturalized species. Basisperma is presently known only from Papua New Guinea, but the myrtle flora is much undercollected and it is expected that it will eventually be recorded from Papua. The genera occurring in New Guinea are: Asteromyrtus (7 species in genus/3 species in New Guinea), Baeckea (14/1), Basisperma (1/1), Decaspermum (ca 30/ca 15), Eucalyptopsis (2/2), Eucalyptus sensu lato (800/ca 15), Eugenia (ca 1000/1), Kania (6/4), Kjellbergiodendron (ca 4/2), Leptospermum (ca 85/1), Lindsayomyrtus (1/1), Lophostemon (4/1), Melaleuca (ca 270/7), Metrosideros (ca 50/ca 4), Myrtella (2/2), Octamyrtus (6/5), Osbornia (1/1), Pilidiostigma (6/1), Psidium (ca 100/1 naturalized), Rhodamnia (ca 26/10), Rhodomyrtus (10/5), Syzygium (incl. Acmena, Acmenosperma, Cleistocalyx; ca 1,200/ca 200Ⳮ), Thaleropia (3/2), Tristaniopsis (ca 40/ca 8), Uromyrtus (ca 20/ca 5), Welchiodendron (1/1), Xanthomyrtus (23/18), Xanthostemon (45/5).

W

Distribution and Habitat Myrtaceae species occur widely and are found in North and South America, Europe, Africa, the Indian Ocean islands, Asia and Southeast Asia, Malesia, Australasia, and the Pacific Ocean islands. They are predominantly tropical to subtropical plants and are particularly plentiful in South America, Southeast Asia, Malesia, and tropical Australia, although there is a major center of development in the subtemperate regions of Australia. Within New Guinea, species of Myrtaceae occur in many habitats, including the littoral zone, savanna, gallery forests, swamp forests, rainforest (both lowland and montane), and subalpine shrubberies. In these habitats, myrtle genera may be rare to occasional or they may be common, and in some cases, such as the Melaleuca and eucalypt savannas, may be the dominant plants.

Family Classification Myrtaceae (incl. Heteropyxidaceae and Psiloxylaceae) is conventionally classified in the order Myrtales, along with eleven other families: Combretaceae, Crypteroniaceae, Lythraceae, Melastomataceae, Oliniaceae, Onagraceae, Penaeaceae, Punicaceae, Sonneratiaceae, Thymelaeaceae, and Trapaceae. Recently it has been demonstrated that the sister family to Myrtaceae may be Vochysiaceae, which had Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

429

................. 16157$

CH29

04-02-07 11:57:43

PS

PAGE 429

.

430 /

previously been placed in Polygalales. Despite some changes in family circumscription, such as the inclusion of Punicaceae, Sonneratiaceae, and Trapaceae in Lythraceae, and the erection of two new families from Lythraceae, Alzateaceae, and Rhynchocalycaceae, the order essentially remains as circumscribed by Cronquist (1981), but with the addition of Vochysiaceae and the removal of Thymelaeaceae to Malvales (Angiosperm Phylogeny Group 1998; Conti et al. 1998). Less agreed upon is the recognition of Heteropyxidaceae and Psiloxylaceae as families separate from Myrtaceae, and of Memecylaceae separate from Melastomataceae. Characteristic features of Myrtaceae include: cavities containing essential oil in most of the unlignified tissues of the shoots (the aromatic oils are often useful field characters for identifying sterile plants); leaves opposite or alternate and often with a distinct intramarginal vein or veins; inflorescences based upon cymes; flowers inferior; fruit a berry or drupe. A more complete description of the family is given in the next section. The genera of the family traditionally have been separated into two groups (whether fruits fleshy or dry) but there are many exceptions, with dryfruited species in otherwise fleshy-fruited genera and vice versa. A new suprageneric classification of Myrtaceae based upon recent molecular and morphological studies has been prepared (Wilson et al. 2005).

Features of the Family Habit trees or shrubs, usually with cavities containing essential oil in the foliage, branchlets and inflorescences. Leaves decussate or alternate, rarely pseudowhorled, simple, often with one or more intramarginal veins, stipules obsolete or sometimes absent. Inflorescences with the basic floral unit a cyme and often elaborated into a paniculate or corymbose structure or rarely the cyme reduced to a 1- or 2flowered unit, the inflorescence axillary, terminal or below the leaves (sometimes on the trunk and/or major branches). Flowers usually bisexual, with the hypanthium adnate to, or free from, the ovary; sepals or calyx lobes (3–)4–5(–6) or calyptrate or rarely absent; petals (3–)4–5(–6) or calyptrate or rarely absent, stamens definite or indefinite in number, often very numerous, free or with the filaments fused at the base and the stamens then usually in bundles, the anthers versatile or not, dorsifixed or basifixed, the connective usually glandular, the thecae dehiscing by slits or pores; ovary semi-inferior, superior or inferior, 1–5(–16)locular, the style terminal, the stigma usually punctate or sometimes capitate or peltate, the placentation axile or parietal, the ovules two to many per locule, anatropous or campylotropous. Fruits a berry, drupe, capsule, or nut, 1- to multiseeded; dehiscent or indehiscent; seeds uni- or rarely polyembryonic, testa present or absent, embryo with the cotyledons relative to the hypocotyl small to very large, sometimes curved or folded.

Gaps in Floristic Documentation or Knowledge Many of the New Guinean genera have been revised in the last three decades. The taxonomic quality of the completed revisions is variable and realignments in sev-

................. 16157$

CH29

02-08-07 10:34:00

PS

PAGE 430

Angiosperms: Myrtaceae of Papua / 431

eral genera are expected as work proceeds on the Flora Malesiana project. The indigenous genera occurring in Papua are included in a key to the genera of Indonesian Myrtaceae (Craven et al. 2003) but treatment of the generic group allied to Myrtus therein is inadequate, reflecting the state of knowledge at the time. Although revisionary work on the large genus Syzygium has commenced, its taxonomy will likely continue to be difficult due to the rarity and undercollection of many of its species and the consequent paucity of morphological information. In fact, the lack of high quality collections from across the whole island is a serious impediment to achieving comprehensive knowledge of its rich myrtle flora. Several genera in New Guinea contain species of present or potential economic uses, such as Asteromyrtus (essential oils) and Syzygium (fruit and timber). Adequate documentation for identification and distribution is required for user groups such as industry, conservation agencies, and land managers. Additionally, in species such as Eucalyptopsis papuana there is potentially significant taxonomic variation that cannot be evaluated due to the lack of comparative stages of growth from presently sampled populations. Specimens with buds, flowers, fruits, and foliage are required from a series of populations for the taxonomic significance of character variation to be properly assessed. This will require monitoring populations over time as the plants undergo a reproductive cycle. Genera for which taxonomic revisions, identification keys, or other systematic information are available include: Asteromyrtus (Craven 1989; Craven et al. 2002), Baeckea (Bean 1998), Basisperma (Foreman 1978), Decaspermum (Scott 1985), Eucalyptopsis (Craven 1990), Gossia (Snow et al. 2003), Kania (Scott 1983; Wilson 1982), Leptospermum (Bean 1992), Lindsayomyrtus (Craven 1990; Hyland and Van Steenis 1973), Lophostemon (Wilson and Waterhouse 1982), Melaleuca (Craven et al. 2002), Myrtella (Scott 1978a), Octamyrtus (Craven and Sunarti 2004), Osbornia (Johns 1980), Rhodamnia (Scott 1979a), Rhodomyrtus (Guymer 1991; Scott 1978b), Syzygium sensu lato (Craven 2001; Craven et al. 2006; Hartley and Craven 1977; Hartley and Perry 1973), Thaleropia (Wilson 1993), Tristaniopsis (Wilson and Waterhouse 1982), Uromyrtus (Scott 1979c), Welchiodendron (Wilson and Waterhouse 1982), and Xanthomyrtus (Scott 1979b).

Comparison with Adjacent Regions The generic links with Australia are stronger than the links with areas to the west or east, not surprisingly in view of the geological connection between the two land surfaces. As far as is known, Basisperma is the only endemic genus of the family in New Guinea, although there are several that have their primary taxic or geographic diversity there (e.g., Eucalyptopsis, Kania, Lindsayomyrtus, Myrtella, Octamyrtus, Thaleropia, Xanthomyrtus). The New Guinean species of Myrtaceae are mainly endemic to the island and its immediate offshore islets. In the case of Syzygium sensu lato, there are over 200 species in New Guinea and 73 in Australia, of which 14 occur on both landmasses. The exact number of species of Syzygium occurring in common on islands to the west of New Guinea will not be known until the

................. 16157$

CH29

02-08-07 10:34:01

PS

PAGE 431

.

432 /

necessary revisionary work is conducted, but overlap is not expected to be high. The shared supraspecific diversity in Syzygium is greater between New Guinea and Australia than between New Guinea and its other neighbors.

Natural History The New Guinean Myrtaceae have a wide diversity in habit. All the species are woody and they are primarily understory, canopy, or emergent trees. When shrubs, they often are occurring in situations in which they form the ‘‘canopy,’’ such as in subalpine shrubberies. Other life forms occurring in New Guinea include treelets, rheophytes, and epiphytes. In the Trans-Fly region, Melaleuca often occurs as the shrubs forming shrub-savannas; it is not known if this condition is a consequence of the strong seasonal inundation in these regions, of the dry season fires, or of both environmental phenomena. In most species of the family, the flowers are small and pollinators are attracted by the flowers being massed in often large inflorescences or compound inflorescences. These species are probably generalists with regard to pollinators and it is believed that their pollinators include insects, birds, and bats. Species with small, solitary, axillary flowers are probably pollinated by insects. An apparently rare example of specialization is found in Octamyrtus. This genus has an increased number of petals relative to most myrtles and the petals are much larger and often brightly colored, usually red; this may represent a shift to bird pollination. Genera with fleshy fruits undoubtedly are zoochorous. Fruit size is probably an indicator of frugivore size, the small- to medium-sized berries and drupes being consumed whole by many birds and mammals and the fruit of the larger-fruited species (especially large-fruited Syzygium species) being consumed whole only by cassowaries, hornbills, and the larger fruit pigeons. Rodents probably also contribute to dispersal of the larger-fruited species by carrying the fruit away before its consumption. The thick testa, characteristic of Decaspermum and Uromyrtus for instance, affords protection against digestion while passing through the gut of animals, but some of the larger-fruited species have thin or no testa, as in Eugenia and many species of Syzygium. In the case of species without an adequate testal defense, it is not known how the embryo is protected; perhaps it does not remain long enough in the gut for damage to occur. Syzygium seeds are known to be capable of germinating even after they have lost much of the cotyledons through herbivory, indicating a considerable degree of robustness. Dispersal of seeds of the dry-fruited genera is believed to be either wind-facilitated (the seeds of Asteromyrtus and of many Tristaniopsis, for instance, are winged) or subject to chance movement by storm winds or chance adhesion to vertebrates. Dispersal of the propagule in genera such as Lindsayomyrtus is unknown; the fruit of Lindsayomyrtus is recorded as fleshy but it may be dehiscent (and thus sheds the seed) or indehiscent.

Acknowledgments Kipiro Damas provided information on the status of two species in Papua New Guinea, and Neil Snow and Peter Wilson provided information on genera for

................. 16157$

CH29

02-08-07 10:34:01

PS PAGE 432

Angiosperms: Myrtaceae of Papua / 433

which they hold unpublished data. Peter Wilson is also acknowledged for reading the manuscript, as is Anna Monro.

Literature Cited Angiosperm Phylogeny Group. 1998. An ordinal classification for the families of flowering plants. Ann Missouri Bot Gard 85: 531–553. Bean, A.R. 1992. The genus Leptospermum Forst. et Forst. (Myrtaceae) in northern Australia and Malesia. Austrobaileya 3: 643–659. Bean, A.R. 1998. A revision of Baeckea (Myrtaceae) in eastern Australia, Malesia and south-east Asia. Telopea 7: 245–268. Conti, E., A. Litt, P.G. Wilson, S.A. Graham, B.G. Briggs, L.A.S. Johnson, and K.J. Sytsma. 1998. Interfamilial relationships in Myrtales: molecular phylogeny and patterns of morphological evolution. Syst Bot 22: 629–647. Craven, L.A. 1989. Reinstatement and revision of Asteromyrtus (Myrtaceae). Austral Syst Bot 1: 373–385. Craven, L.A. 1990. One new species each in Acmena and Eucalyptopsis and a new name in Lindsayomyrtus (all Myrtaceae). Austral Syst Bot 3: 727–732. Craven, L.A. 2001. Unravelling knots or plaiting rope: what are the major taxonomic strands in Syzygium sens. lat. (Myrtaceae) and what should be done with them? Pp. 75–85 in Saw, L.G., L.S.L. Chua, and K.C. Khoo (eds.) Taxonomy: The Cornerstone of Biodiversity. Proc. Fourth Fl. Mal. Symp. Eds. Inst. Pen. Perhutanan Malaysia, Kuala Lumpur. Craven, L.A., E. Biffin, and P.S. Ashton. 2006. Acmena, Acmenosperma, Cleistocalyx, Piliocalyx and Waterhousea formally transferred to Syzygium (Myrtaceae). Blumea 51: 131–142. Craven, L.A., and S. Sunarti. 2004. A new species of, and reinstatements in, Octamyrtus (Myrtaceae). Gard Bull Singapore 56: 147–152. Craven, L.A., S. Sunarti, D. Mudiana, T. Yulistiarini, and M. Wardani. 2003. Identification key to the indigenous Indonesian genera of Myrtaceae. Floribunda 2: 89–94. Craven, L.A., S. Sunarti, M. Wardani, D. Mudiana, and T. Yulistiarini. 2002. Kayu putih and its relatives in Indonesia (Myrtaceae, Melaleuca and Asteromyrtus). Floribunda 2: 16–26. Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia University Press, New York. Foreman, D.B. 1978. Notes on Basisperma lanceolata C.T. White (Myrtaceae). Brunonia 1: 95–101. Guymer, G.P. 1991. Revision of the Rhodomyrtus trineura (F.Muell.) F.Muell. ex Benth. (Myrtaceae) species complex. Austrobaileya 3: 377–387. Hartley, T.G., and L.A. Craven. 1977. A revision of the Papuasian species of Acmena (Myrtaceae). J Arnold Arb 58: 325–342. Hartley, T.G., and L.M. Perry. 1973. A provisional key and enumeration of species of Syzygium (Myrtaceae) from Papuasia. J Arnold Arb 54: 160–227. Hyland, B.P.M., and C.G.G.J. van Steenis. 1973. The generic identity of Xanthostemon brachyandrus C.T. White: Lindsayomyrtus novum genus (Myrtaceae). Blumea 21: 189–192. Johns, R.J. 1980. Notes on New Guinea Myrtaceae. 3: the genus Osbornia F. von Mueller. Klinkii 1: 63–67. Scott, A.J. 1978a. A new species of Myrtella (Myrtaceae) from Australia and a synopsis of the genus. Kew Bull 33: 299–302.

................. 16157$

CH29

02-08-07 10:34:02

PS PAGE 433

.

434 /

Scott, A.J. 1978b. A revision of Rhodomyrtus (Myrtaceae). Kew Bull 33: 311–329. Scott, A.J. 1979a. A revision of Rhodamnia (Myrtaceae). Kew Bull 33: 429–459. Scott, A.J. 1979b. A revision of Xanthomyrtus (Myrtaceae). Kew Bull 33: 461–484. Scott, A.J. 1979c. New species and combinations in Myrtaceae from Malesia and Australia. Kew Bull 33: 511–515. Scott, A.J. 1983. Two new species of Kania (Myrtaceae) from New Guinea. Kew Bull 38: 309–310. Scott, A.J. 1985. Decaspermum (Myrtaceae) in New Guinea. Kew Bull 40: 149–165. Snow, N., G.P. Guymer, and G. Sawvel. 2003. Systematics of Austromyrtus, Lenwebbia, and the Australian species of Gossia (Myrtaceae). Syst Bot Mon 65: 1–95. Wilson, P.G. 1982. Additions to the genus Kania (Myrtaceae) in Malesia with notes on Cloezia. Blumea 28: 177–180. Wilson, P.G. 1993. Thaleropia, a new genus for Metrosideros queenslandica (Myrtaceae) and its allies. Austral Syst Bot 6: 251–259. Wilson, P.G., and J.T. Waterhouse. 1982. A review of the genus Tristania R.Br. (Myrtaceae): a heterogeneous assemblage of five genera. Austral J Bot 30: 413–446. Wilson, P.G., M.M. O’Brien, M.M. Heslewood, and C.J. Quinn. 2005. Relationships within Myrtaceae sensu lato based on a matK phylogeny. Pl Syst Evol 251: 3–19.

................. 16157$

CH29

02-08-07 10:34:02

PS

PAGE 434

Orchidaceae of Papua ´

.

recent estimates (Hassler 2001; Royal Botanic Gardens, Kew, 2003), the Orchidaceae comprise about 25,000 known species worldwide and thereby represent the largest family of flowering plants on earth. The most prolific author on New Guinea orchids, Rudolf Schlechter (1914), believed that New Guinea was the richest area in the world as far as orchids are concerned, but we now know that the most significant global orchid hotspot is to be found in the northern Andes (Colombia, Ecuador, Peru). There is no doubt, however, that New Guinea is a very close second. Perhaps as many as 2,800 species occur here, accounting for about 11 percent of the world’s orchid flora (Schuiteman and de Vogel 2001). The total number of orchid genera is between 820 and 1,042, depending on which authority is used. Of these, 132 (or a few dozen more, according to authors who prefer to split up larger genera) have wild representatives in New Guinea.

A

Family Classification The now widespread use of DNA analyses in phylogeny reconstruction has had a significant impact on orchid classification. The picture that has emerged is that the family consists of five main clades, which are recognized at the rank of subfamily (Pridgeon et al. 1999; Chase et al. 2003): Apostasioideae, Vanilloideae, Cypripedioideae, Orchidoideae, and Epidendroideae. All have representatives in New Guinea. The small subfamily Apostasioideae, with only about 17 species in two genera, forms the sister clade to all other orchids. At first glance these are rather nondescript monocots that most people would not recognize as orchids, as they have very simple, seemingly almost regular flowers with two or three normal-looking anthers and powdery pollen. They do possess, however, the characteristic orchid column that is formed by fusion of the filaments and the style, although the fusion is incomplete here. The Apostasioideae are confined to Southeast Asia and Malesia, including Australia. In New Guinea only two species, Apostasia wallichii R.Br. and Neuwiedia veratrifolia Blume, occur as inhabitants of the forest floor in hill forest; both are widespread outside New Guinea as well. Surprisingly, the Vanilloideae, and not the Cypripedioideae as previously thought by taxonomists, are next in line, forming the sister clade to the remaining subfamilies. The Vanilloideae possess typical orchid flowers, in that they are highly zygomorphic (i.e., bilaterally symmetrical), while the single fertile stamen and pistil are completely fused to form the column. This is essentially a pantropical group, with some outliers in northeast North America, Japan, and Australia. It contains Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

435

................. 16157$

CH30

02-08-07 10:34:09

PS

PAGE 435

436 /

´

.

approximately 235 species in 15 genera. In New Guinea this subfamily is represented by five genera: Vanilla, Pseudovanilla, Galeola, Cyrtosia, and Lecanorchis, with about a dozen species in total, none of which depart to any significant extent from their extra-New Guinean congeners. The subfamily Cypripedioideae differs from the Vanilloideae and the two terminal subfamilies Orchidoideae and Epidendroideae, in that the species in this group possess not one, but two fertile anthers. In this respect it resembles the genus Apostasia of the Apostasioideae, but otherwise Apostasioideae and Cypripedioideae have little in common. The Cypripedioideae consist of the famous slipper orchids, so called for the shape of the lip (the median petal). This subfamily occurs in America and Eurasia, while it is absent from Africa, Australia, and New Zealand. It contains about 165 species in five genera. Only the genus Paphiopedilum is found in New Guinea, with four species belonging to two different clades, which are both much more speciose outside New Guinea. The shape of the lip is paralleled in the mainly New Guinean genus Pedilochilus (Figure 3.6.15; probably better regarded as a section within the polymorphic giant Bulbophyllum), which is a member of subfamily Epidendroideae. The Orchidoideae constitute an important subfamily in New Guinea, where it is second in number of species with 31 genera and some 185 species. Members of this subfamily are mainly terrestrials with soft, herbaceous, not plicate leaves, which are inrolled (convolute), not folded, in bud. They often possess underground tuberoids. This large and cosmopolitan subfamily (roughly 250 genera with 4,600 species) is further subdivided into various tribes and subtribes, which we will not describe here, but it is worth noting that the subtribe Goodyerinae is particularly well represented in New Guinea. There are several striking endemic species in this subtribe (e.g., Pristiglottis coerulescens (Schltr.) Cretz. & J.J.Sm., which has bluish green flowers, a highly exceptional color in this alliance), and even an endemic genus, Papuaea, is present. The most notable phenomenon in the subfamily Orchidoideae in New Guinea is the extensive radiation shown by the genus Corybas (Figure 3.6.16). These are conspicuous and yet elusive little plants, generally with a single, cordate leaf and only one, disproportionally large, pitcher-shaped flower. The genus is widespread from India to New Zealand, with about 125 species, of which ca 45 are found in New Guinea, all of which are endemic, as far as we know now. The final subfamily is Epidendroideae, by far the most speciose in New Guinea, with perhaps as many as 2,600 species in 93 genera. This is a huge and cosmopolitan clade, comprising an estimated 20,000 species in 550 genera. The New Guinea members of this subfamily belong to 21 subtribes, of which some, like the Bulbophyllinae, the Dendrobiinae, and the Thelasiinae, have proliferated in New Guinea more than anywhere else in the world. By far the most species-rich genus is Bulbophyllum, which may well contain some 600 species in New Guinea (about 1,700 worldwide), followed by Dendrobium (400), Phreatia (130), Glomera (100), Taeniophyllum (90), Malaxis (90), Oberonia (90), and Liparis (80). These figures are rather crude estimates because the taxonomy of most of these genera is insuffi-

................. 16157$

CH30

02-08-07 10:34:10

PS

PAGE 436

Angiosperms: Orchidaceae of Papua / 437

Figure 3.6.15. Pedilochilus sp. Photo: P. Jongejan.

ciently known. It is nevertheless instructive to compare them with the relevant numbers for the island of Borneo, which has by no means a depauperate orchid flora (about 1,400 known species): Bulbophyllum (215), Dendrobium (145), Phreatia (7), Taeniophyllum (12), Malaxis (32), Oberonia (27), Liparis (37), and Glomera (0). There is evidently not only a lower diversity in these genera, even if we take the smaller size of Borneo into account, but the spectrum of diversity is also rather

................. 16157$

CH30

02-08-07 10:34:33

PS

PAGE 437

438 /

´

.

Figure 3.6.16. Corybas sp. (probably an undescribed species). Photo: A. Schuiteman.

................. 16157$

CH30

02-08-07 10:34:57

PS

PAGE 438

Angiosperms: Orchidaceae of Papua / 439

different. For example, certain genera of the Coelogyninae are poorly represented in New Guinea, in particular Coelogyne (6 species; Figure 3.6.17), Dendrochilum (1), and Pholidota (3), each containing far fewer species than are found on Borneo: Coelogyne (66), Dendrochilum (83), and Pholidota (11). Recent DNA studies have unexpectedly shown that the genus Glomera belongs to the same subtribe; therefore, it can no longer be maintained that the Coelogyninae as a whole are poorly represented in New Guinea.

Features of the Family Habit: terrestrial or more frequently epiphytic herbs, sometimes vines, occasionally without chlorophyll, often with aerial roots. Stems very short to much elongated, often green and swollen (forming so-called pseudobulbs). Leaves simple, sometimes reduced to scales, alternate, rarely opposite, often in two alternating ranks or more or less spirally arranged, margin entire or rarely serrulate, stipules absent. Inflorescence terminal or lateral, a panicle or a raceme, often reduced to a single flower. Flowers usually resupinate, zygomorphic, bisexual, exceptionally unisexual

Figure 3.6.17. Coelogyne carinata Rolfe. Photo: E. F. de Vogel.

................. 16157$

CH30

02-08-07 10:35:08

PS

PAGE 439

440 /

´

.

(not in New Guinea), not rarely spurred. Tepals six, those of the outer whorl (sepals) mostly almost equal, but not rarely distinctly unequal, free or in various ways connate. Tepals of the inner whorl (petals) unequal, the median one (the labellum or lip) free, usually markedly different from the lateral ones; lateral petals (usually just referred to as the petals) free or rarely adnate to the sepals. Stamens three, two, or mostly one, in the latter case the filament usually entirely fused with the style into a structure called the column that carries the anther(s) and the stigma. Anther often with an easily detached anther-cap. Pollen usually forming tetrads (grains in cohering groups of four), in the three basal subfamilies in monads (single grains), usually arranged into two, four, six, or eight discrete bodies (pollinia), these mostly forming part of a specialized pollen transfer structure of frequently rather complicated structure (the pollinarium), sometimes pollen-mass gel-like or powdery and not forming discrete pollinia. Ovary inferior, 1-locular or rarely 3-locular. Fruit a dry capsule, very rarely a berry. Seeds many, usually minute, almost always lacking endosperm (food-storage tissue), usually consisting of a naked embryo surrounded by a loose seed coat of dead tissue which has a netlike surface structure, rarely with a tight and hard seed coat.

Distribution and Habitat Orchids occur from the North Cape to Tierra del Fuego and in almost all the land in between (Dressler 1981). They are, however, very unevenly distributed, both geographically and ecologically. The family is essentially a tropical one, as is evident from the very low percentage—probably less than five percent—of species that occur in temperate parts of the world. Within the tropics orchids are poorly represented in arid regions, such as much of Africa. Conversely, they may reach astounding levels of diversity in areas with an everwet climate where temperatures are moderate at all times, which is mainly in the mountains between ca 900 and 2,500 m above sea level. This is undoubtedly correlated with the epiphytic life style of the great majority of the orchids: the habitats hosting the greatest number of orchid species are those where conditions are most conducive to the growth of epiphytic vascular plants. Much of New Guinea is covered with precisely such moist, mild montane forests that are favored by epiphytic orchids, which may partly explain their abundance on this island. About eighty percent of the orchids of New Guinea are epiphytes (as against seventy percent worldwide). Some of the terrestrial species grow in swampy ground and may have ‘‘wet feet’’ for part of the year, but no orchid in New Guinea is truly aquatic. When considering the many different habitats in New Guinea it can be observed that the global patterns of orchid distribution are reiterated on a smaller scale: orchids shun the driest, hottest, or coldest areas. Consequently, they are very scarce or almost absent in seasonally dry savannas and grasslands, while in moist evergreen forests their diversity first increases when ascending from sea level upwards and then decreases again towards the subalpine zone. Of the evergreen lowland forest types, mangrove forest is the poorest in orchid

................. 16157$

CH30

02-08-07 10:35:08

PS

PAGE 440

Angiosperms: Orchidaceae of Papua / 441

species, though a few, such as the inconspicuous Dendrobium viridiflorum F.M. Bailey, appear to be restricted to this habitat. Freshwater swamp forest, while not as poor as mangroves, is likewise not rich in orchids, partly because terrestrial orchids are virtually absent: most of these cannot survive inundation for long. Beach forest is somewhat richer, especially on Calophyllum trees, where several species of Dendrobium sect. Latouria, Cleisostoma, and others occur. Lowland rainforest, in particular in hill country with its associated streams, rocky outcrops, slopes, and ridges, can even at low elevations already be quite rich in orchids, both terrestrial and epiphytic. Many species, and a limited number of genera, are not found above 800 m altitude, as for example the genera Renanthera and Trichoglottis. On the other hand, the number of species and genera not found below 800 m is far greater. With increasing altitude orchid species richness steadily increases, reaching a peak in the zone between 1,000 and 1,500 m. Above 2,000 m the tree genus Nothofagus is often dominant; this is clearly an excellent host tree for orchids, judging from the enormous masses of epiphytes that are often seen clothing the branches. This Nothofagus forest can be surprisingly tall given its occurrence at high altitudes, with individual trees up to 50 m high. Above 2,700 m the forest becomes increasingly open, everything is covered in cushions of dripping wet moss, and the borderline between epiphytes and terrestrials becomes blurred: many species appear equally at home in moss cushions on the ground and on the branches of trees and shrubs. At these high altitudes orchids are very common in terms of individuals, but diversity is clearly lower than it is at around 2,000 m. Among the most frequent orchids here are species of Glomera, which superficially resemble miniature ericaceous shrubs, but with totally different flowers (Figure 3.6.18). On most New Guinea mountains above 3,500 m the forest gives way to grassland and shrubland (the subalpine zone), and this in turn, on the highest summits, to bare rock with very sparse vegetation (the alpine zone). Only 25 orchid genera with some 200 species occur above 3,000 m (van Royen 1979, revised). Most of these are epiphytes in the lower part of the subalpine zone, or grow on mossy rocks and banks. Orchids may be found in New Guinea up to about 4,000 m; those at the upper limit belong to the genera Dendrobium, Octarrhena, Pedilochilus, and Thelymitra. It is noteworthy that only the last mentioned is a strictly terrestrial genus of open country—the three other genera comprise mainly epiphytic forest plants—so it seems likely that most subalpine orchids in New Guinea evolved from ancestors growing in the forest at lower altitudes. Grasslands in New Guinea, whether lowland or montane, are generally poor in orchid species. In grasslands at high altitudes, above about 2,700 m, it is fairly common to find orchids belonging to species that are predominantly epiphytic growing as opportunistic terrestrials in the peaty soil or in moss cushions. At the same time, relatively few truly terrestrial species, of genera such as Spathoglottis, Calanthe, and Peristylus, occur exclusively in montane grasslands, and they are rarely found in great numbers. This is in contrast to some other parts of the world,

................. 16157$

CH30

02-08-07 10:35:09

PS

PAGE 441

442 /

´

.

Figure 3.6.18. Glomera hamadryas (Schltr.) J.J.Sm. Photo: A. Schuiteman.

such as south and east Africa, where orchids can be highly diverse and abundant in this biome. A remarkable habitat in the mountains, apparently unique to New Guinea, is grassland with stands of tree ferns (tree fern savanna; see Paijmans 1976), which creates a striking, archaic looking landscape. This occurs mainly above 2,700 m. Although orchids are often cultivated on slabs of tree fern stem, which largely consists of the densely interwoven roots of the fern, only a relatively limited number colonize tree fern trunks in nature. These often appear to be specialists, with species of Pedilochilus being the most prominent in the tree fern savanna. Another special biome is that of karst landscapes, which are both extensive and virtually impenetrable in several parts of New Guinea, including in the Vogelkop of Papua, in the southern highlands of Papua New Guinea, and in the Bismarck Archipelago. Very little is known about their orchid flora. Our own observations around the village of Ayawasi in the Vogelkop indicate that limestone hills with their characteristic steep slopes and flat tops and generally rather open and low forest can be extremely rich in orchid species, even at the moderate altitude of 400–500 m. We suspect that it is not so much the limestone itself, but rather the varied topography and openness of the forest associated with it, that promotes an unusually high orchid diversity.

Endemism New Guinea is not only remarkable for the absolute number of orchid species it harbors; perhaps the extremely high percentage of endemic orchid species is even more impressive: 90%! To put this figure in perspective, it may be noted that the

................. 16157$

CH30

02-08-07 10:35:20

PS PAGE 442

Angiosperms: Orchidaceae of Papua / 443

second highest percentage in the Indo-Pacific region is 74% for the Philippines, while Borneo (55%) and Java (31%) score very much lower (Agoo et al. 2003). As a corollary one might, therefore, expect that most of the orchid species in New Guinea have very narrow ranges, reflecting the complex topography and geological history of this huge landmass. Surprisingly, this is in general not the case. Revisions of several orchid genera or sections of genera, such as Bulbophyllum (Vermeulen 1993), Dendrobium (Reeve and Woods 1989; Cribb 1983, 1986), and Mediocalcar (Schuiteman 1997), have all shown that the majority of the species are widespread on the island. This can probably be explained by the fact that most species occur in the central mountain range, which allows dispersal along most of its length, running almost from the western to the eastern tip of the island. Moreover, during Pleistocene glaciations, montane vegetation zones which are presently separated by valleys probably formed a more or less continuous range, since the belts in which they occur were situated at lower elevations. Consequently, such readily identified species as Bulbophyllum olivinum J.J.Sm., Dendrobium cuthbertsonii F.Muell., D. cyanocentrum Schltr., and Mediocalcar uniflorum Schltr. may be found almost throughout New Guinea wherever suitable habitats occur. This even includes the more isolated mountain ranges along the northern coast. Many other species are not quite so widespread, but still have ranges that span several hundreds of kilometers. Undoubtedly, highly localized endemics do exist, but due to our still scanty knowledge of the distribution of most orchids it is hard to be certain about particular cases. For lowland species one could conjecture that the central mountain range effectively blocks north-south dispersal and vice versa. So there might be widespread species that are only found north or south of this barrier, of which species it could then be hypothesized that they evolved after the uplift of the central range, which started about five million years ago (Axelrod and Raven 1982). As a matter of fact, there are only very few good examples of this, such as Dendrobium lineale Rolfe and D. convolutum Rolfe (both north). Most of the widespread lowland species are found both north and south of the central range, such as D. antennatum Lindl., D. spectabile Blume, and Cadetia transversiloba J.J.Sm., although several are either much more common in the north or in the south, such as D. lasianthera J.J.Sm. (Figure 3.6.19), D. mirbelianum Gaud. (both north), and D. discolor Lindl. (south). Of course, there are numerous ‘‘rare’’ species that are represented by a single dot on the distribution map. It is hard to draw conclusions from these, in particular since their taxonomic status is in many cases not clear. In complete contrast to endemism at species level, endemism at higher taxonomic levels in New Guinea is low. This is, however, to some extent almost certainly because clades that originated in New Guinea managed to disperse to neighboring regions. Truly endemic (and named) clades include Dryadorchis (5 species; Figure 3.6.20), Ophioglossella (1), Papuaea (1), Ridleyella (1), Bulbophyllum sect. Vesicisepalum (3), and others. Among the subendemic clades that most likely originated in New Guinea are Aglossorhyncha, Cadetia (Figure 3.6.21), Diplocaulobium, Epiblastus, Eurycentrum, Glomera, Hymenorchis, Mediocalcar, Microtatorchis, Pedilochilus, Pseuderia, Dendrobium sect. Biloba, Dendrobium sect.

................. 16157$

CH30

02-08-07 10:35:21

PS PAGE 443

444 /

´

.

Figure 3.6.19. Dendrobium lasianthera J.J.Sm. Photo: T. Roelfsema.

Grastidium, D. sect. Herpetophytum, D. sect. Latouria, Bulbophyllum sect. Coelochilus, B. sect. Lepanthanthe, B. sect. Peltopus, and B. sect. Polyblepharon, to list the more significant ones. They have in common that at least 90% of their species occur only in New Guinea, while the extra-territorial species either occur in New Guinea as well, or do not differ substantially from species indigenous in New Guinea.

Gaps in Floristic Documentation and Knowledge Our knowledge of the distribution patterns of New Guinea orchids is still highly fragmentary. Quite a few species are at present only known from the type collection, that is, from a single locality, such as many species of the genus Corybas (van Royen 1983). Whether or not these truly represent narrow-ranged endemics is hard to tell. There are several examples of species that have been collected only twice, but in localities hundreds of kilometers apart. To further complicate matters, the collecting density for Papua New Guinea is about five times higher than for Papua, where only about one percent of the entire area has ever been visited by a botanist. This is well illustrated by the known distribution of the easily recognized and conspicuous Dendrobium hellwigianum Kraenzl. While there are literally dozens of records from all over Papua New Guinea (Reeve and Woods 1989), only recently a single collection from the center of Papua turned up the first record from this part of New Guinea (Schuiteman and de Vogel 2002). It seems unlikely that it really is common throughout Papua New Guinea and at the same time very rare in Papua, especially so because this is by no means an isolated example. There

................. 16157$

CH30

02-08-07 10:35:30

PS

PAGE 444

Angiosperms: Orchidaceae of Papua / 445

Figure 3.6.20. Dryadorchis dasystele Schuit. & de Vogel. Photo: E. F. de Vogel.

are several others, such as D. aurantiroseum Royen ex T.M.Reeve and Cadetia chionantha (Schltr.) Schltr., which show a comparable pattern.

Natural History Compared with neighboring areas, the orchid flora of New Guinea shows some special features. We have already mentioned the unusually high radiation of certain clades within the family, the most dramatic example being the genus Bulbophyllum (Figure 3.6.22, 23). In Papua New Guinea, one of us (A.S.) once picked up a piece of a recently fallen branch on a trail through Nothofagus forest. It was about 40 cm long and 3 cm diameter and carried mature specimens of seven different species of Bulbophyllum! This not only nicely demonstrates the diversity of this genus, it also implies that these are often very small plants indeed. Small or even tiny orchids are certainly not unique to New Guinea, but it is fairly obvious that the proportion of what amateurs call ‘‘miniature orchids’’ is definitely higher in New Guinea than elsewhere in the Indo-Pacific region. Another observation,

................. 16157$

CH30

02-08-07 10:35:40

PS

PAGE 445

446 /

´

.

Figure 3.6.21. Cadetia arfakensis (J.J.Sm.) Schltr. Photo: A. Schuiteman.

already made by Schlechter (1914), is the extraordinarily large number of orchids in New Guinea that have pendulous stems, either because the stems are positively geotropous (as in the aptly named Dendrobium geotropum T.M.Reeve), or more frequently because they are flaccid and much elongated. In many species of Bulbophyllum that display this habit it is the string-like rhizome that is hanging down perpendicularly, often more than one meter long. Again, this phenomenon is not unique to New Guinea, but the frequency with which it occurs is. It may be related to the constantly humid climate and the absence of very strong winds over much of New Guinea. Similarly, bird-pollinated orchids are certainly not restricted to New Guinea, but it is probably true to say that no area in the world, with the possible exception

................. 16157$

CH30

02-08-07 10:35:59

PS

PAGE 446

Angiosperms: Orchidaceae of Papua / 447

Figure 3.6.22. Bulbophyllum macrorhopalon Schltr. Photo: E.F. de Vogel.

of Andean South America, has so many of these as New Guinea. It must be admitted that actual observations are few and far between, but it is well known which properties characterize bird-pollinated flowers (van der Pijl and Dodson 1966). Among those properties are: flowers opening during the day; weakly expressed zygomorphy (i.e., the flowers appear not strongly bilaterally symmetrical); tubular and hard-textured flowers; vivid colors (often including orange or red and often with stark contrasts); no scent; abundant nectar in relatively short and broad receptacles; nectar guide absent; flowers horizontal or hanging freely in space. Based on these properties it can be predicted that many dozens of New Guinea orchids, probably at least a hundred species, are ornithophilous (pollinated by birds). This includes all species of the genera Mediocalcar (Figure 3.6.24) and Epiblastus (Figure 3.6.25), and in addition many species of Glomera and Dendrobium (e.g., D. fulgidum Schltr.; Figure 3.6.26), as well as a few species of Calanthe (Schuiteman 1997). Most of these occur in mountain habitats that are generally cool and cloudy, with relatively few insects, and it has been conjectured in the case of Rhododendron that ornithophilous species evolved at least partly in response to the paucity of insects (Stevens 1976). The rhododendrons are mainly or even exclusively pollinated by Meliphagidae (honeyeaters), and the few existing observations on orchids implicate these birds as well. In this connection one of the most remarkable orchids of New Guinea warrants a separate discussion.

................. 16157$

CH30

02-08-07 10:36:12

PS

PAGE 447

448 /

´

.

Figure 3.6.23. Bulbophyllum tricanaliferum J.J.Sm. Photo: P. Jongejan.

................. 16157$

CH30

02-08-07 10:36:39

PS

PAGE 448

Angiosperms: Orchidaceae of Papua / 449

Figure 3.6.24. Mediocalcar bifolium J.J.Sm. Photo: A. Schuiteman.

Dendrobium cuthbertsonii F.Muell. (Figure 3.6.27) is a highly appealing miniature orchid. Even without flowers it is easily recognized, as the small, deep green leaves are normally covered with numerous crystalline warts, unlike any other orchid in the world. The solitary, short-stalked flowers are relatively enormous (up to 5 cm wide, but usually half that size), bell-shaped, and are held in a horizontal position, with the lip uppermost. Apart from its great beauty, the most striking aspect of this species is the color polymorphism it displays in at least some of the localities where it is found (generally above 2,000 m altitude). The most frequent color form has pure scarlet-crimson flowers, with a darker edging to the lip, which is common to nearly all color forms. But in some populations, especially those in more open habitats, these red forms are intermixed with individuals having pink, creamy yellow, or purple flowers, as well as particularly dazzling bicolored forms: red with yellow, purple with orange, pink with cream, and purple with white. In view of the fact that these flowers do not contain nectar, and that they occur sympatrically with Rhododendron species having superficially similar flowers, it is tempting to speculate that D. cuthbertsonii mimics Rhododendron flowers. As the latter are ornithophilous, the color-polymorphism may serve to prevent the pollinating birds from learning to avoid visiting Dendrobium cuthbertsonii altogether. It would be fascinating to unravel the population genetics behind this mechanism. If our hypothesis is correct it will probably be found that differ-

................. 16157$

CH30

02-08-07 10:36:50

PS

PAGE 449

450 /

´

.

Figure 3.6.25. Epiblastus sp. Photo: E. F. de Vogel.

ently colored forms are cross-pollinated more frequently than individuals with the same colors. Dendrobium cuthbertsonii is a widespread and locally common species, so its strategy is obviously successful. This in spite of the fact that fruit set is often low, even though the individual flowers last several months. In contrast, the often sympatric, ornithophilous orchids of the genus Mediocalcar do produce nectar, and frequently set many fruits. The individual flowers of these last at most about two to three weeks. There are clearly several tradeoffs involved here.

................. 16157$

CH30

02-08-07 10:37:10

PS

PAGE 450

Angiosperms: Orchidaceae of Papua / 451

Figure 3.6.26. Dendrobium fulgidum Schltr. Photo: A. Schuiteman.

We have no doubt that pollination studies on New Guinea orchids will, one day, lead to surprising discoveries. Some species, especially in the genus Bulbophyllum, have such bizarre flowers that it is hard to predict how they are pollinated. As one example we would like to single out B. cimicinum J.J.Verm. (Figure 3.6.28). This is the most insect-like, or more exactly, arthropod-like in appearance of all orchids that we know of. The sepals are unremarkable, but the petals and the lip are modified almost beyond recognition: the two petals each carry four black, hairy, leg-like appendages, which are suspended by minute threads, so that they wriggle in the lightest breeze. The lip is also black, covered with many shining, rod-like papillae, and looks for all the world like the abdomen of an arthropod. The whole contraption seems to mimic a little spider sitting in the middle of a flower. What can be the pollinator of this? We know, of course, many examples where the lip of an orchid mimics a female bee, deceiving male bees to the extent that they try to copulate with the flower. But it seems unlikely that this Bulbophyllum is aiming at other spiders, as these are bound to be very inefficient pollinators. One possibility we can think of is that the orchid mimics a prey to attract spidercatching insects, such as robber flies or certain wasps. On the other hand, the similarity of the flower to a spider may be entirely coincidental. The chances that we will unravel the pollinating mechanism in the near future seem remote, as Bulbophyllum cimicinum appears to be very rare in the wild, and the individual flowers last only one day. It is but one of several species of the section Epicrianthes, all of which possess bizarre insect-like flowers with highly diverse appendages,

................. 16157$

CH30

02-08-07 10:37:21

PS

PAGE 451

452 /

´

.

Figure 3.6.27. Dendrobium cuthbertsonii F.Muell. Photo: T. Roelfsema.

some leg-like, some balloon-shaped (Figure 3.6.22), and others resembling slender tentacles. This section has a center of diversity in New Guinea, where about 15 out of 32 species occur. In another section of Bulbophyllum, Monosepalum, which is subendemic in New Guinea, the petals are also provided with mobile appendages, but the structure of the flowers is entirely different from that in section Epicrianthes. Numerous other examples of unusual morphologies are evident, like that of Bulbophyllum inauditum Schltr. This rare species has inconspicuous flowers, less than 1 cm across, unremarkable except for the presence of a pair of 13 cm long flaccid appendages at the tips of the lateral sepals that dangle in the wind, no doubt to attract its as yet unknown pollinator, and that remind us of the disproportionate tail feathers of certain birds of paradise. In B. ustusfortiter J.J.Verm. the sepals are almost entirely fused to form a dark brown, papillose-hairy tube with only a small apical hole through which the pollinator can enter. Perhaps the most noteworthy fact in this case is the circumstance that in Panama a very distantly related orchid with almost identical flowers occurs: Dresslerella pertusa Luer.

Local Uses Few orchids in New Guinea are used by humans. There are however some notable exceptions in the genus Diplocaulobium (Figure 3.6.29). In several species strips of

................. 16157$

CH30

02-08-07 10:37:33

PS PAGE 452

Angiosperms: Orchidaceae of Papua / 453

Figure 3.6.28. Bulbophyllum cimicinum J.J.Verm. Photo: P. Jongejan.

the elongated pseudobulbs provide a very tough, shiny and yellow fiber, which is widely applied for decoration purposes (e.g., arm bands, small bags, and penis gourds). Plants of Diplocaulobium are often cultivated for this purpose in villages on roofs, posts, and even on top of carved wooden statues, where they resemble mops of hair. Dendrobium flowers are frequently used as head adornments, being stuck in the hair along with leaves, feathers, everlastings, and other items. But on the whole the role of orchids in the lives of the indigenous people is very limited.

Conservation Orchids are to the Plant Kingdom what birds and butterflies are in the animal world: a group of organisms that attract attention for their beauty and that are exposed to a variety of human-made threats (Dixon et al. 2003). The first threat that usually comes to mind is collecting for commercial purposes. While this is

................. 16157$

CH30

02-08-07 10:37:50

PS PAGE 453

454 /

´

.

Figure 3.6.29. Diplocaulobium hydrophilum (J.J.Sm.) Kraenzl. Photo: E. F. de Vogel.

also a problem in New Guinea, the governments of both Indonesia and Papua New Guinea have essentially banned the export of wild-collected orchids. This has evidently reduced these practices considerably, because only very limited numbers of wild collected orchids from New Guinea are now seen in the trade. Moreover, collecting will only become a problem if particular species are targeted. Most of the thousands of New Guinea orchid species are not so desirable that collectors would make a special effort to procure every last specimen. It is probably true to

................. 16157$

CH30

02-08-07 10:38:09

PS

PAGE 454

Angiosperms: Orchidaceae of Papua / 455

say that only species of Paphiopedilum and a limited number of Dendrobium species have such a high commercial value and such a low number of populations that they could be collected to extinction if collecting were not prohibited. The main threat to the future existence of wild orchids, as to all other wildlife, is clearly not the collecting of individual plants but the wholesale destruction of their habitats. This depressing subject hardly requires further comment, and we can only express our hope that New Guinea will forever remain the orchid paradise it still is.

Acknowledgments We thank the Cheng Kim Loke Foundation, the Hermon Slade Foundations, and Conservation International for supporting the NHN New Guinea orchid project, and the Netherlands Foundation for the Advancement of Tropical Research (WOTRO) as well as the Netherlands Organisation for Scientific Research (NWO) for supporting fieldwork in Papua New Guinea and Papua.

Literature Cited Agoo, E.M.G., A. Schuiteman, and E.F. de Vogel. 2003. Flora Malesiana: Orchids of the Philippines, vol. I; Illustrated Checklist and Genera. CD-ROM. ETI, Amsterdam; Nationaal Herbarium Nederland, Leiden. Axelrod, D.I., and P.H. Raven. 1982. Paleobiogeography and origin of the New Guinea flora. Pp. 919–949 in Gressit, J.L. (ed.) Biogeography and Ecology of New Guinea, vol. 2. Dr. W. Junk Publishers, The Hague. Chase, M.W., K.M. Cameron, R.L. Barrett, and J.V. Freudenstein. 2003. DNA data and Orchidaceae systematics: a new phylogenetic classification. Pp. 69–89 in Dixon, K.W., S.P. Kell, R.L. Barrett, and P.J. Cribb (eds.) Orchid Conservation. Natural History Publications, Kota Kinabalu, Sabah. Cribb, P.J. 1983. A revision of Dendrobium section Latouria (Orchidaceae). Kew Bull. 38: 229–306. Cribb, P.J. 1986. A revision of Dendrobium section Spatulata (Orchidaceae). Kew Bull. 41: 615–692. Dixon, K.W., S.P. Kell, R.L. Barrett, and P.J. Cribb (eds.). 2003. Orchid Conservation. Natural History Publications, Kota Kinabalu, Sabah. Dressler, R.L. 1981. The Orchids, Natural History and Classification. Harvard University Press, Cambridge, Massachusetts. ¨ berblick u¨ber die Familie Orchidaceae und eine weltweite Hassler, M. 2001. Statistischer U Checkliste der Orchideen. Pp. 2826–2898 in Schlechter, R. (ed.) Die Orchideen, vol. 1/ C. Parey Buchverlag, Berlin. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO, Canberra. Pridgeon, A.M., P.J. Cribb, M.A. Chase, and F. Rasmussen (eds.). 1999. Genera Orchidacearum, vol. 1. Oxford University Press, Oxford. Reeve, T.M., and P.J.B. Woods. 1989. A revision of Dendrobium section Oxyglossum (Orchidaceae). Notes Roy. Bot. Gard. Edinburgh 46: 161–305. Royal Botanic Gardens, Kew. 2003. Monocot Checklist. Available at http://www.kew .org/wcsp (accessed 2 February 2004).

................. 16157$

CH30

02-08-07 10:38:10

PS

PAGE 455

456 /

´

.

Schlechter, R. 1914. Die Orchidaceen von Deutsch-Neu-Guinea. Rep. Spec. Nov. Regni Veg., Beih. 1: i–lxvi, 1–1058. Schuiteman, A. 1997. Revision of the genus Mediocalcar. Orchid Monogr. 8: 21–77. Schuiteman, A., and E.F. de Vogel. 2001. Orchids of New Guinea, vol. I; Illustrated Checklist and Genera. CD-ROM. ETI, Amsterdam; Nationaal Herbarium Nederland, Leiden. Schuiteman, A., and E.F. de Vogel. 2002. Flora Malesiana: Orchids of New Guinea, vol. II; Dendrobium and allied genera. CD-ROM. ETI, Amsterdam; Nationaal Herbarium Nederland, Leiden. Stevens, P.F. 1976. The altitudinal and geographical distributions of flower types in Rhododendron section Vireya, especially in the Papuasian species, and their significance. Bot. J. Linn. Soc. 72: 1–33. van der Pijl, L., and C.H. Dodson. 1966. Orchid Flowers: Their Pollination and Evolution. University of Miami Press, Coral Gables. van Royen, P. 1979. The Alpine Flora of New Guinea, vol. 2. J. Cramer, Vaduz. van Royen, P. 1983. The Genus Corybas (Orchidaceae) in Its Eastern Areas. J. Cramer, Vaduz. Vermeulen, J.J. 1993. Taxonomic revision of Bulbophyllum, sections Adelopetalum, Lepanthanthe, Macrouris, Pelma, Peltopus and Uncifera. Orchid Monogr. 7: 1–324.

................. 16157$

CH30

02-08-07 10:38:11

PS

PAGE 456

Sapindaceae of Papua . Number of Genera and Species have 140 genera and 1,350 species worldwide. In Malesia the family is represented by 42 genera and 235 species. The majority of the species and genera are found in New Guinea (30 genera, 150 species); of these 25 genera and 67 species are known for Papua and 30 genera and 133 species for Papua New Guinea (52 species in both regions). The discrepancy in numbers across regions is probably caused mainly by insufficient sampling in Papua.

T

Distribution and Habitat The Sapindaceae are a pantropical family with few genera in the more temperate zones (or in Papua at the higher elevations). Two more speciose temperate groups are Acer and Aesculus. The Southeast Asian Sapindaceae have their main diversification in New Guinea, and these species are mainly endemic or extend to Australia or the Moluccas. Very few species are widespread; most of those are endemic in the whole of Malesia (e.g., all Papuan species of Lepisanthes). Most Papuan species are understory or midstory species in lowland primary rainforest, but some also occur in secondary vegetations, especially in habitats where the grass cover is not too high, permitting easy germination and establishment (vanWelzen 1989: 38). Pometia pinnata is the only dominant canopy species (Tristiropsis is also a canopy tree, but never dominant). Very few species reach high altitudes or are restricted to altitudes above 1,500 m (e.g., several species in Atalya, Cnesmocarpon, Guioa, Sarcopteryx, and Sarcotoechia). Also, a few species are adapted to more specialized habitats: Arytera litoralis, Harpullia leptococca, and Dodonaea viscosa occur mainly in coastal vegetations. Most species occur in everwet conditions, but can often also grow under monsoonal conditions. Similarly, the soil can be well drained to marshy. Sometimes, the species also occur at the edge of mangroves, but they are not really a part of the mangrove biome.

Family Classification The Sapindaceae is placed in the order Sapindales and forms one family with the temperate Aceraceae and Hippocastanaceae (Judd et al. 1999). Most typical for the Sapindaceae is the extrastaminal disk (and in Asia, usually the paripinnate leaves). The family was divided by Radlkofer into two subfamilies (Dodonaeiodeae and the Sapindoideae: two or more apotropous ovules per locule in the Dodonaeiodae or sometimes one, but then these epitropous; one apotropous ovule per locule in the Sapindoideae). The two subfamilies are subdivided into several tribes, Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

457

................. 16157$

CH31

02-08-07 10:35:42

PS

PAGE 457

458 /

.

but none of these seem to be natural groups (nor are the Dodonaeiodeae). Most Papuan genera are part of the tribe Cupanieae (Sapindoideae), a group with (relatively) small capsules as fruits. The economically most important timber and fruit tree in Papua, Pometia pinnata (called matoa locally), is classified in the tribe Nephelieae, a tribe closely related to or part of the Cupanieae, but with larger, indehiscent capsules (and usually edible arils or sarcotestas). The most recent revision of the family is by Adema, Leenhouts, and van Welzen (1994).

Features of the Family Habit: shrubs or trees, lianas or climbing herbs (Cardiospermum), usually functionally dioecious. Hairs usually simple, solitary, sometimes two-branched (Litchi) or stellately bundled (Dimocarpus, Harpullia); scale hairs present in a few genera, then inflorescences and young parts sticky. True stipules absent (except Cardiospermum), sometimes basal leaflets as pseudostipules. Leaves mainly paripinnate (no terminal leaflet), sometimes double pinnate (Tristiropsis), 3-foliolate (Allophylus), imparipinnate (some Lepisanthes) or simple (Dodonaea) or unifoliolate, rachis winged or not, leaflets opposite to alternate, base, margins, apex and indumentum variable, often domatia in nerve axils on lower surface, venation pinnate. Inflorescences (cauliflorous to) axillary to terminal, usually thyrsoid panicles; bracts and bracteoles present. Flowers seemingly bisexual, functionally unisexual, usually actinomorphic. Sepals usually four or five, free to almost totally connate, equal and sepaloid to distinctly unequal, then the outer one or two much smaller than the inner three and all often petaloid. Petals absent to 2–6, free, usually clawed, often with one or two scales or auricles ( inrolled margins), scales with inward small scale ( crest) or not. Disk extrastaminal, usually ringlike or horseshoe-like (open near 5th petal), usually lobed, rarely with appendages or a tubular rim. Stamens 5–10 (to many), usually eight, often exserted in staminate flowers; filaments glabrous or hairy; anthers basifixed, opening introrsely or latero-introrsely lengthwise, staminodal and not opening in pistillate flowers. Ovary superior, 1–3(–8)-celled, lobed or not, style usually apical, rarely inserted between the lobes, stigma entire with (1), 2, or 3 lines or grooves or lobed; rudimentary in staminate flowers; ovules one or two or more, apotropous and erect or ascending or epitropous and hanging. Fruits capsular or drupaceous or consisting of two or three samaras, when capsular usually opening loculicidally, rarely septicidally or septifragally. Seeds globose to obovoid, sometimes compressed, often with an arillode or a sarcotesta; endosperm absent; embryo usually thick, straight, sometimes sigmoid or convolute, cotyledons above each other (notorrhizal) to laterally besides each other (lomatorrhizal). Mainly understory trees in primary or secondary rainforest, forest edges, scrubland, savanna, coastal vegetation, often along rivers or roads; in everwet or seasonal conditions; mainly lowland (to up to 3,600 m); on all kinds of soil.

................. 16157$

CH31

02-08-07 10:35:43

PS

PAGE 458

Angiosperms: Sapindaceae of Papua / 459

Gaps in Floristic Documentation or Knowledge The Sapindaceae remain poorly known in Papua. Most collecting has been done in the Vogelkop, around Jayapura, and to a lesser extent, in the Star Mts; the rest of Papua remains virtually unsurveyed for this group. The expectation is that still quite a few Sapindaceae are undescribed and that more material would help to unravel a few species complexes.

Natural History Sapindaceae have been recorded as monoecious or dioecious except Dodonaea, which may have bisexual flowers. Most Sapindaceae show dichogamy: three stages of floral development appear within the same inflorescence. The first flowers to appear are functionally staminate (long filaments, dehiscing anthers, functional pollen, undeveloped pistil). After they have dropped, the second flush of flowers, pistillate, opens (short filaments, indehiscent anthers, nonfunctional pollen, welldeveloped pistil). After fruit set the third phase of flowers opens, organs of both sexes present but flowers functionally staminate, with intermediate to long filaments, dehiscing anthers, functional pollen, more or less well-developed ovary. (If fruit set in the former phase was poor, the flowers in this phase may also act as pistillate flowers.) In duodichogamy the second and third phase are usually repeated as the fourth and fifth phase. The consecutive phases may overlap. Dichogamous plants with synchronized flowering within the same plant are effectively dioecious too, which means that most Sapindaceae have to be cross-pollinated, though occasional self-pollination was reported (Appanah 1982; van Welzen 1989). There is apparently no self-incompatibility since interflower selfing occurs in several economically important plants. The Sapindaceae are mainly entomophilous; Dodonaea is the only genus within the family for which wind-pollination is recorded. The most important pollinators are bees, mainly of the genera Trigona (stingless bees) and Apis (honeybees); stingless bees seem to be the main pollinators. They may be attracted either by the nectar or by the pollen, and many plant species are reported to have fragrant flowers (the attractant may be different for the various species of Trigona). The trees seem to have three mechanisms to ensure crosspollination by bees. The most important is using the bee’s ability to memorize collecting sites. The first phase in dichogamous flowering, though functionally staminate, probably serves only to put the tree on the bee’s visiting list, after which the bee will remain visiting the less attractive pistillate flowers. (The staminate and bisexual flowers are visited much more often as they are more colorful due the exserted, colored stamens.) The second mechanism to ensure cross-pollination by bees is fluctuations in nectar productivity; staminate or bisexual flowers alternate with pistillate flowers in nectar production during the daytime (five nectar peaks,

................. 16157$

CH31

02-08-07 10:35:43

PS

PAGE 459

460 /

.

starting and ending with the staminate flowers). The third mechanism is differences in nectar composition between staminate and pistillate flowers. The staminate flowers offer monosaccharides in the nectar and proteins via the pollen; the pistillate flowers have nectar with disaccharides and protein (Appanah 1982; van Welzen 1989).

Fruits and dispersal are covered in van Welzen, Lamb, and Wong (1988). Young fruits of the Sapindaceae are usually green, hard, and full of tannin. When mature, the color changes, usually to red, and the fruit wall softens and the tannin level drops. Sapindaceae may fruit throughout the year, once or twice per year. They tend to have a good harvest one season, followed by a poor harvest the next season. Several fruit types occur in the Papuan Sapindaceae: winged fruits (Atalaya and Dodonaea); inflated fruits with a thin, papery wall (Cardiospermum); drupaceous fruits (Allophylus and Lepisanthes); dehiscent capsules, usually with (partial) arillodes around the seeds (most Cupanieae); and indehiscent ‘‘capsules,’’ also usually with rather thick arillodes around the seeds (few Cupanieae and most Nephelieae). The seeds or fruits are mainly dispersed by birds and mammals. The smaller fruits, like those with drupaceous fruits and the seeds of the Cupanieae (dehiscent capsules), are mainly eaten by birds, which are attracted by the contrasts in color between the fruit (yellow to mainly red), arillode (also mainly yellow to red, but different shade than fruit), and the seed (shiny dark brown to black). Several genera (Guioa, Mischocarpus, Sarcopteryx) add movement as an attractant: the arillode has an appendage (pseudofunicle) that attaches to the basal corner of the fruit and the seed remains dangling on this appendage after fruit dehiscence. The seeds of the larger fruits in this group are mainly preyed upon by parrots, parakeets, and presumably pigeons, especially the crown pigeons. The indehiscent and mainly larger fruits are eaten by mammals. The commercially interesting species are in this category. These fruits also have a red color as attractant. They are reported to be eaten and dispersed by humans, monkeys, bats, and pigs; parakeets and parrots may eat these fruits too. Winged and inflated fruits are reported to be dispersed by wind or (sea)water, just like the woody seeds of Pometia.

The seeds of Sapindaceae usually germinate readily, within a week, and passage through an animal’s gut is not a prerequisite. The seeds are short-lived and do not show dormancy. During germination the radicle and petioles swell and become terete. The testa usually opens at the place where the embryonic radicle is present. The radicle is quite often enclosed in a testal pouch. The testa subsequently opens along the pleurograms, after which the petioles, hypocotyl, and radicle elongate. The cotyledons, turning green, remain loosely enclosed in the testa. This type of germination is classified as Horsfieldia type and subtype and has been reported for several genera. The seedlings may differ considerably in macromorphology from

................. 16157$

CH31

02-08-07 10:35:44

PS

PAGE 460

Angiosperms: Sapindaceae of Papua / 461

the mature plants. The first leaves are usually opposite (alternate in adults) and the rachis is usually slightly winged (usually unwinged in adults). The margin of the juvenile leaflets is (crenate to) serrate (usually entire in adults). There are no papillae and domatia on the lower surface of the juvenile leaflets. In size and number of leaflets the leaves of seedlings may differ dramatically from those in mature plants. Plants still showing immature characters may flower and fruit, thus hampering species delimitation. At least several of the mountain species in Guioa seem to retain the juvenile syndromes when mature (van Welzen 1989).

Several genera, Alectryon, Guioa, Harpullia, and Sarcopteryx, show species associated with ants. The branchlets, which are hollow, are inhabited by black, foulsmelling ants. The branchlets are usually swollen below the nodes and the nest openings may be found in the swellings. The nature of the relationship is still unknown, but likely to be symbiotic, ant protection in exchange for accommodation. Something similar may occur with the leaflet domatia, present in many genera and species. There appears to be a relationship between the presence of domatia on leaves and the leaves’ occupation by predaceous or fungivorous mites (bug or fungus-eating bugs) (O’Dowd and Wilson 1989).

Literature Cited Adema, F., P.W. Leenhouts, and P.C. van Welzen. 1994. Sapindaceae. Flora Malesiana ser. I, 11 (3): 419–768. Rijksherbarium/Hortus Botanicus, Leiden. Appanah, S. 1982. Pollination of androdioecious Xerospermum intermedium Radlk. (Sapindaceae) in a rain forest. Biol. J. Linn. Soc. 18: 11–34. Judd, W.S., C.S. Campbell, E.A. Kellogg, and P.F. Stevens. 1999. Plant Systematics: A Phylogenetic Approach. Sinauer, Sunderland, Massachusetts. O’Dowd, D.J., and M.F. Wilson. 1989. Leaf domatia and mites on Australasian plants: ecological and evolutionary implications. Biol. J. Linn. Soc. 37: 191–236. van Welzen, P.C. 1989. Guioa Cav. (Sapindaceae): Taxonomy, phylogeny, and historical biogeography. Leiden Bot. Ser. 12: 1–315. van Welzen, P.C., A. Lamb, and W.W.W. Wong. 1988. Edible Sapindaceae in Sabah. Nature Malaysiana 13 (1): 10–25.

................. 16157$

CH31

02-08-07 10:35:44

PS

PAGE 461

Sapotaceae of Papua

Number of Genera and Species are a pantropical family. The latest estimates of the size of this family (Pennington 1991, 2004) indicate about 1,100 species in 53 genera. Of the tropical areas, Central and South America contain the highest number of species, whereas Africa and Asia are richer in genera. In Malesia 14 genera are found, but of these only nine are represented in New Guinea, including Magodendron, endemic to eastern Papua New Guinea. Although in the field the Sapotaceae often seem to be one of the more important families because of the number of individuals observed, on the species level New Guinea is rather poor. The nine genera represented in New Guinea have together a total of 784 species, but New Guinea as a whole accounts only for about 75 species and Papua for 48. Of the large genera, Pouteria (313 species, pantropical but mainly neotropical) accounts for 44, more than half of all sapotaceous species in New Guinea; Palaquium (119 species in Asia and the Pacific, with a center in Borneo) is second most speciose, with 12 species. The other large genera, Madhuca (116 species from India to New Guinea), Manilkara (82 species, pantropical), and Chrysophyllum (81 species, mainly neotropical) have only four, three, and two outliers, respectively, as New Guinea representatives. The remaining genera each account for fewer than ten species.

T

Endemism Of the genera, only Magodendron is endemic to New Guinea. On the species level the endemism is rather high: of the 75 species known from New Guinea, 49 are endemic. The differences in the floristic exploration of Papua and Papua New Guinea (PNG) are also reflected in the numbers of species endemic to these regions: of the 48 sapotaceous species known from Papua, six species (12%) are regional endemics, whereas of the 64 species known from PNG, 27 species (42%) are regional endemics. Pouteria is the best represented in New Guinea, by 44 species of which 33 are endemics, mostly found in both Papua and PNG.

Characterization of New Guinean Sapotaceae Habit shrubs to large trees. The leaves are simple and entire, spirally arranged but sometimes subopposite (in Pouteria; Sarcosperma) with stipules present or absent. The small actinomorphic flowers are arranged in fascicles; in some species of Pouteria these fascicles occur on leafless shoots, in Sarcosperma fascicles or solitary Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

462

................. 16157$

CH32

02-08-07 10:35:51

PS

PAGE 462

Angiosperms: Sapotaceae of Papua / 463

flowers are arranged on racemose to paniculate leafless shoots. The 4–8 sepals are free or basally united, in one or two series, persistent in fruit. The corolla is gamopetalous, with a short to long tube and 4–8 lobes, the latter in Manilkara and Mimusops usually with lateral appendages. The 4–25 stamens are inserted in the tube or on the throat of the corolla, epipetalous, and between the lobes present, absent, or replaced by staminodes. A small disk (mainly found in Pouteria) may surround the ovary, free or (partly) connate with it. The superior 2–12locular ovary has one style; the ovules are solitary and axile to basal. The fruit is usually a fleshy berry. The seeds almost always have a hard, dark, shining testa but the area of attachment of the seed to the fruit wall (the ‘‘scar’’) can be very large (e.g., in Burckella, Pouteria). When the cotyledons are foliaceous they are embedded in thick albumen, but when they are thick the albumen is (almost) absent. The indumentum consists of malpighiaceous hairs, whether or not mixed with simple hairs. An important field character is the presence of white, sticky latex in the bark and fruits, often also in other parts.

Subdivision of the Family The family has had a checkered taxonomic history because in this family sets of characters are not universal, and always have exceptions. This inconsistency gave rise to different opinions on the weighting of individual characters or of character sets. Baehni (1965) even introduced variability as a generic character. The taxonomic history has been well documented by several authors, including Baehni (1938, 1965) and Aubre´ville (1964), and, most recently, by Pennington (1991). Presently five tribes are recognized and all five are represented in New Guinea: Mimusopeae (Manilkara, Mimusops); Isonandreae (Burckella, Madhuca, Palaquium); Sideroxyleae (Sarcosperma); Chrysopylleae (Chrysopyllum, Pouteria); and Omphalocarpeae (Magodendron).

Quick Recognition of the Genera in New Guinea (sepals are persistent in fruit; deviations in numbers do occur): 1. Inflorescences cauliflorous ................................................................Magodendron 1. Inflorescences axillary to leaves or leaf scars, or on leafless shoots (ramiflorous in Pouteria keyensis) .............................................................................................. 2 2. Sepals in two pairs of 2 ....................................................................... Madhuca 2. Sepals in one row of 4 ......................................................................... Burckella 2. Sepals in two rows of 3 .................................................................................... 3 3. Staminodes 0; seed scar lateral ....................................................Palaquium 3. Staminodes 6; seed scar subbasal ................................................ Manilkara 2. Sepals in two rows of 4 ......................................................................Mimusops 2. Sepals in one row of 5 ...................................................................................... 4 4. Petiole at top with tiny stipellae .......................................... Sarcosperma

................. 16157$

CH32

02-08-07 10:35:52

PS

PAGE 463

464 /

4. Petiole without stipellae ......................................................................... 5 5. Flowers with alternipetalous staminodes ..............................Pouteria 5. Flowers without staminodes .......................................Chrysophyllum Revisions of the Malesian Sapotaceae have been made by H. J. Lam and colleagues (especially P. van Royen) at the National Herbarium at Leiden in the series ‘‘Revision of the Sapotaceae of the Malaysian area in a wider sense’’ I–XXIII, published in Blumea and Nova Guinea between 1952 and 1960 (listed in van Royen 1960:432). The large genus Planchonella, treated in that series, is now included in Pouteria. The accumulation of collections made in the second half of the twentieth century necessitates a new round of revisional work.

Natural History Although a shrubby habit does occur (e.g., Pouteria lanatifolia), the Sapotaceae are usually small or large trees, up to 50 m high with a bole up to 120 cm in diameter (e.g., Pouteria thyrsoidea). The bole is mostly cylindrical; in Pouteria keyensis it can be fluted over its entire length. Buttresses, if present, may be up to 5 m high (in Pouteria thyrsoidea). I have not seen stilt-roots mentioned for New Guinea Sapotaceae. Only Pouteria keyensis is ramiflorous; the two species of Magodendron are cauliflorous. Most common in the New Guinea Sapotaceae is a tree architecture according to the Model of Aubre´ville: long bare horizontal branches terminating in an upright short part with densely clustered leaves, elongating sympodially from the base of the upright portion (Terminalia-branching). Flowers and fruits can be located between the foliage or below the leaves on the bare part of the branches. Chris Versteegh reported a tree of Pouteria ledermannii (Figure 3.6.30) in lowland rainforest to be completely defoliated, but generally flushes of new leaves appear beyond mature leaves.

Most Sapotacae have bisexual flowers. By reduction of the anthers, some to all flowers (in an individual) can become female, including Pouteria lanatifolia, P. monticola (Vink 2002), P. linggensis, and P. obovata (van Royen 1957). Female flowers can be smaller than bisexual ones. Subba Reddi and Bai (1980) found Mimusops elengi to be trioecious: plants with male, female, or bisexual flowers only.

Usually the reproductive entities in a collection are in the same stage of development: all buds, all open flowers, all corollas falling, all young fruits of the same stage, all ripe fruits. Sometimes fruit ripening can take a long time: several times flowers of the next flowering stage were found on fruiting trees of Burckella obovata (large fruits) and Palaquium amboinense (medium-sized fruits).

................. 16157$

CH32

02-08-07 10:35:52

PS

PAGE 464

Angiosperms: Sapotaceae of Papua / 465

Figure 3.6.30. Pouteria ledermannii; fresh gathering from a low-montane ultrabasic environment. The species is distributed throughout New Guinea.

Very little is known about the pollinators of the Sapotaceae. A ‘‘nectary disk’’ is found in some species of Pouteria and in Magodendron, but nectar has never been observed. The only record of nectar, produced at the base of the ovary, is for Mimusops elengi, which, however, does not have a disk (Subba Reddi and Bai 1980). On Waigeo Island, van Royen observed bees as visitors on flowers of Manilkara fasciculata. According to Wiselius (1998) the flowers of Chrysophyllum roxburghii are pollinated by insects. Mimusops elengi has very fragrant nocturnal flowers, but there are strong indications that this species is wind-pollinated and that the visiting bees do not contribute to the pollination (Subba Reddi and Bai 1980). In herbarium specimens I never found thrips caught in the dried flowers. In the Aru Islands, M. M. J. van Balgooy (pers. comm.) saw hordes of parrots on profusely flowering ramiflorous Pouteria keyensis.

Many sapotaceous fruits are fleshy and edible for animals, but in New Guinea observations of animals eating the fruits are rare. Although data on fruit color are incomplete, some generalities have emerged. There is a tendency for large fruits, of about 10 cm, to have a dull light greenish to yellowish brown color (Burckella,

................. 16157$

CH32

02-08-07 10:36:15

PS

PAGE 465

466 /

Pouteria doonsaf, P. maclayana, P. ripicola), except for the fruits of the cauliflorous Magodendron, which become almost black. There are species with medium-sized fruits (3–7 cm, Palaquium amboinense, Pouteria thyrsoidea) and small-sized fruits (1–3 cm, Manilkara fasciculata, Pouteria lanatifolia, P. monticola) that remain (yellowish) green when ripe, but fruits in these size classes are in some cases red (e.g., Pouteria suboppositifolia) or, more often, red becoming purple to almost black (e.g., Pouteria myrsinodendron). Others have a wide range of ripe fruit color, such as Pouteria obovata, which may have brown, orange, red, purple, or black fruits! Majnep and Bulmer (1977) describe fruits of Pouteria macropoda as one of three favorite foods of the Mountain Pigeon (Gymnophaps albertisi). Sterly (1997) names the same species as a food for birds and marsupials. On Yapen Island I have seen fruits of Manilkara fasciculata being eaten by crown pigeons. Hamann and Curio (1999) observed fruit pigeons and starlings foraging on Palaquium fruits in the Philippines. Stocker and Irvine (1983) report germination of seeds of Niemeyera chartacea and Pouteria species collected from cassowary dung in northern Queensland. Bats are also among the dispersal agents of Sapotaceae. Evidence is from surrounding countries only. For Indonesia, van der Pijl (1957) lists Mimusops elengi and Pouteria duclitan ( Sideroxylon sundaicum) among the plants dispersed by bats. In the Philippines Palaquium lanceolatum is a ‘‘suspected’’ bat plant (Stier 1990). In New South Wales Pouteria ( Planchonella) australis is included in the diet of the Grey-headed Flying Fox (Pteropus poliocephalus) (Nature Conservation Council of NSW 2002). In the neotropics Sapotaceae belong to the families important in the diet of bats (Fleming 1986; van der Pijl 1957). The fruits of the New Guinean Pouteria thyrsoidea dangle from their leafless inflorescence axis at the end of the branches below the tufts of large leaves. This ‘‘penduliflory’’ (van der Pijl 1957) might indicate bat dispersal, also hypothesized by van der Pijl for some species of Palaquium and Madhuca. In about 1645, all plant species on the volcanic Long Island, PNG, were extirpated by an explosive eruption, and no plant remains have been found in the upper layers of ash deposits. Recently Pouteria maclayana was found on the crater rim (Harrison et al. 2001). This species has dull fruits up to 13 cm large and seeds up to 5.5 by 3 cm; its dispersal agent is as yet unknown. The cauliflorous Magodendron apparently presents its flowers and rather large fruits to animals living on the forest floor. The importance of the preservation of dispersal vectors is emphasized by Wiselius (1998): ‘‘Due to the fragmentation of the Sungei Menyala forest (Peninsular Malaysia) the mammals that once ate and dispersed the seed of Chrysophyllum roxburghii have disappeared and today piles of rotting fruits accumulate below the parent trees.’’

There are two types of seed. One type has foliaceous cotelydons embedded in endosperm, such as, in New Guinea, Chrysophyllym, Manilkara, Mimusops, Pouteria p.p. The seedling is epigeal and phanerocotylar (Macaranga type (de Vogel

................. 16157$

CH32

02-08-07 10:36:16

PS

PAGE 466

Angiosperms: Sapotaceae of Papua / 467

1980), e.g., Chrysophyllum roxburghii (pers. obs.)). The second type of seed has thick planoconvex cotelydons with storage function, (almost) lacking the endosperm, characteristic of, for example, Burckella, Madhuca, Magodendron, Palaquium, Pouteria p.p., and Sarcosperma in New Guinea. The seedling is hypogeal and more or less cryptocotylar (Sloanea type, Palaquium subtype, e.g., Palaquium amboinense; de Vogel 1980). Germination periods were found to be from 6–23 days in Pouteria obovata, to 24–82 days in Mimusops elengi (Ng 1992).

Of the native species, Burckella obovata is a fruit tree that is spared from felling when people make new gardens. The ripe mellow fruit smells very pleasant and is readily eaten (Peekel 1984). According to Kambuou (1995), it is a highly preferred fruit in the Bismarck Archipelago; in the Solomon Islands it is listed with eight other species of edible fruit of prime importance (Ministry of Agriculture and Fisheries 1996). Kirch (1989) lists Burckella obovata with the currently important tree crops in Melanesia and reports its presence among the tree crops in the Lapita culture site in the Mussau Islands, PNG, dating from 1,600–500 . . . Of the introduced American species, Manilkara zapota, Chrysophyllum oliviforme, and especially C. cainito, are fruit trees, but they are not widely used in New Guinea. Sapotaceous fruits must be fully ripe and soft to be eaten; otherwise the tannin content and the latex make them completely inedible (Ng 1992).

Apparently the Sapotaceae belong to the families that do not form mycorrhizas. Galls are very rare in Sapotaceae.

Although the Sapotaceae are often mentioned as one of the important families in the lowland rainforest, sapotaceous species almost never account for 1% of the trees 34 cm dbh and over. There are, however, several exceptions to this rule. The hydrological reserve ‘‘Tafelberg’’ (G. Meja) above Manokwari is at 150– 175 m altitude on a plateau of old sediments surrounded by coral limestone. Here, the dryland mixed forest is dominated by Pometia (21%), Intsia bijuga (7.5%), Neonauclea (4.5%), and Palaquium amboinense (3.5%). Other Sapotaceae are Pouteria (0.4%) and Chrysophyllum roxburgii (0.2%) (Zieck 1960). The Tami coastal plateau east of Jayapura is a terrace of coral limestone at 10–80 m altitude. According to Zieck (1959), 23% of the canopy trees in the closed forest are Pometia and 22% consist of species of Sapotaceae genera (Pouteria, Palaquium, Chrysophyllum, Manilkara). The most frequent in the very diverse remainder are Alstonia (5%), Moraceae (5%), and Burseraceae (2%). In the fluvial-deltaic plains of the Buna-Kokoda area, PNG, Taylor (1964) recognized a ‘‘Pometia pinnata–Dillenia quercifolia–Palaquium spec. alliance’’ that included the ‘‘P. pinnata–D. quercifolia–Palaquium spec. association’’ and the

................. 16157$

CH32

02-08-07 10:36:16

PS

PAGE 467

468 /

‘‘Palaquium spec. association,’’ the latter on more poorly drained sites. The three species mentioned usually make up over 50% of the trees in the dominant layer of any one stand in this alliance. Other common species in this forest are Vitex cofassus, Alstonia scholaris, Anisoptera thurifera subsp. polyandra, and Ficus species. There are extensive lowland rainforests on the Pacific side of the Vogelkop: the Warsamson Valley in the northwest and the Arfak Plain in the northeast. The demography of these forests shows relicts of historic widespread disturbance (Vink 1998). The Warsamson Valley is 32,600 ha and was inventoried at 0.4–0.5% of the trees of 34 cm dbh and over and 695 botanical collections were made; the Arfak Plain of 23,600 ha was inventoried at the 2% level and 1,412 collections were made. No sapotaceous species reach 1% of the tree number. In the Warsamson Valley nine species of Sapotaceae were found, including Burckella magusun and B. polymera, which did not appear in the more intensively studied Arfak Plain. Fifteen species of Sapotaceae in the genera Chrysophyllym, Madhuca, Palaquium, and Pouteria were collected on the Arfak Plain. Of these, Chrysophyllum roxburghii, Madhuca leucodermis, Pouteria keyensis, and P. ledermannii were apparently quite common. During his Guttapercha- und Kautschuk-Expedition (1907–1909) to the Torricelli Mts, the area south of Madang, and the Finisterre Mts in Papua New Guinea, Schlechter (1911) found P. supfianum and P. warburgianum in low densities, but widespread and with sufficient regrowth (0–6 large trees per ha with a mean of ca one or two large trees per ha). For three of his harvesting trials he could use between 34 and 82 trees of 30–100 cm diameter. According to Schlechter these species are never found in disturbed forest, but seedlings show increased growth after being exposed to more light. Some species are locally gregarious although otherwise they occur scattered in the forest; this might indicate that they react to some disturbance by massive regrowth. Examples are Manilkara fasciculata on the south coasts of the McCluer Gulf, Pouteria anteridifera (Figure 3.6.31) in the Trans Busu Timber Lease east of Lae, PNG, and Pouteria thyrsoidea on Biak Island, now probably all logged. Pouteria chartacea can occur in dense uniform stands on frequently inundated river banks or lake shores in seasonal areas.

A number of endemics are known from a few collections only and their rarity inhibits a circumscription of their preferences. Many more common species are found in a wide variety of habitats. Most versatile is probably Pouteria obovata. It grows on sandy or rocky beaches and is named as a constituent of the Barringtonia formation; it is even reported as colonizing pebble beach in the sea spray. It also occurs on the landward side of mangroves, in coastal swamps, on freshwater lake shores, and in riverine forest. Drier sites include small forest patches on limestone savannas, well-drained primary and secondary lowland forests, and oak forests. In New Guinea Pouteria obovata is found up to 400 m altitude, in the Lesser Sunda Islands up to 1,650 m altitude. Pouteria keyensis also grows in very different habi-

................. 16157$

CH32

02-08-07 10:36:17

PS

PAGE 468

Angiosperms: Sapotaceae of Papua / 469

Figure 3.6.31. Pouteria anteridifera (Lane-Poole) Baehni. a. habit (X 1). b. flower (X 8). c. stamen (X 6). d. gynoecium (X 12). e. longitudinal section of gynoecium (X 12). f. cross-section of ovary (X 12). g. fruit (X 1). h. seed. Drawing: Ruth van Crevel; courtesy Flora Malesiana Foundation.

................. 16157$

CH32

02-08-07 10:37:10

PS

PAGE 469

470 /

tats on a wide variety of soils: in flatland well-drained primary (and old secondary) forests and hill forests, but also along watercourses, on marshy terrain, or on terrain inundated in the wet season. Near Oriomo, PNG, it is associated with Acacia, but near Vanimo, PNG, it occurs in beach forest on the shoreline. Its altitudinal range is from sea level up to 250 (once at 1,980) m.

Beach Forest In addition to Pouteria obovata, Manilkara kauki has a preference for beach forests but it also grows inland on grass plains under seasonal conditions.

Low-Altitude Freshwater Swamp Forest Palaquium ridleyi is common in mixed swamp forest, but is sometimes found in dryland forests. Palaquium galactoxylon is found in permanent swamp or in periodically flooded lowland forest, but also in dryland hill forest and nonseasonal lowland forests up to Castanopsis forest at 450 m altitude.

Rainforests at Low Altitudes (up to 700 m) These are the habitat of most of the New Guinea Sapotaceae, such as Chrysophyllum roxburghii, Manilkara fasciculata, Palaquium amboinense, P. lobbianum, P. obtusifolium, P. supfianum, P. warburgianum, Pouteria maclayana, P. menait (sometimes in floodplain forest or swamp forest), P. rhopalocarpa, P. ripicola (riverine), P. thyrsoidea (rarely in seasonal and nonseasonal swamp), P. toricellensis.

Forests at Wide Altitudinal Ranges A further number of Pouteria species have the ability to occupy quite different habitats. It is not possible to specify the optimum in their habitat ranges. In rainforests from sea level up to about 1,000 m altitude are found Pouteria linggensis var. linggensis (also in coastal areas), P. luzoniensis var. papuana, P. myrsinodendron (up to 2,500 m), P. pullenii, P. sussu (up to 1,600 m). Pouteria ledermannii is rather common in lowland forest on alluvial clays, and is found scattered in Melaleuca forest, in Lithocarpus-Castanopsis forest, and in lower montane forest on ultrabasic soil; its altitudinal range is from sea level up to 1,500 m. The widespread but rare Sarcosperma paniculata occurs in primary and secondary rainforests, less often in forest edges or in (bamboo) thickets, from sea level up to 1,100 m altitude.

(Sub)montane Forests A small number of Sapotaceae are apparently restricted to higher altitudes. Pouteria kaernbachiana is a canopy or subcanopy tree 24–30 m high in forests at 800– 1,500 m altitude. Pouteria lamii, an endemic of the Hunstein Range, is a smaller tree found at 100 and 1,300 m altitude. Pouteria monticola occurs throughout New Guinea as a (sub)canopy tree or treelet at 450–2,200 m altitude, often associated with Castanopsis, Lithocarpus, or Nothofagus. At higher altitudes, 1,750–2,800 m, Pouteria macropoda var. macropoda grows in mossy forests. The endemic of the Anggi Lakes, Pouteria lanatifolia, is a 2–6 m high shrub growing in secondary myrtaceous-ericaceous shrubbery or in forest edges at 2,140–2,200 m.

................. 16157$

CH32

02-08-07 10:37:10

PS

PAGE 470

Angiosperms: Sapotaceae of Papua / 471

Literature Cited Aubre´ville, A. 1964. Sapotace´es. Adansonia, Me´moire 1: 1–157. Baehni, C. 1938. Me´moires sur les Sapotace´es. 1. Syste`me de classification. Candollea 7: 394–508. Baehni, C. 1965. Me´moire sur les Sapotace´es. 3. Inventaire des genres. Boissiera 11: 1–262. de Vogel, E.F. 1980. Seedlings of dicotyledons. Centre for Agric. Publ. and Doc., Wageningen. Fleming, T.H. 1986. Opportunism versus specialization: the evolution of feeding strategies in frugivorous bats. Pp. 105–118 in Estrada, A., and T.H. Fleming (eds.) Frugivores and Seed Dispersal. W. Junk, Dordrecht. Hamann, A., and E. Curio. 1999. Interactions among frugivores and fleshy fruit trees in a Philippine submontane rainforest. Conservation Biology 13: 766–773. Harrison, R.D., R. Banka, I.W.B. Thornton, and M. Shanahan. 2001. Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. II. The vascular flora. J. Biogeography 28: 1311–1337. Kambuou, R.N. 1996. Papua New Guinea: country report to the FAO International Technical Conference on Plant Genetic Resources, Leipzig: 26. Kirch, P.V. 1989. Second millennium . . arboriculture in Melanesia: archeological evidence from the Mussa Islands. Economic Botany 43: 225–240. Lam, H.J. 1948. Sarcospermaceae. Flora Malesiana I, 4: 32. Majnep, I.S., and R. Bulmer. 1977. Birds of My Kalam Country. Auckland University Press/Oxford University Press, Oxford. Ministry of Agriculture and Fisheries. 1996. Solomon Islands: country report to the FAO International Technical Conference on Plant Genetic Resources, Leipzig: 13. Nature Conservation Council of NSW. 2002. Flying Foxes Policy (as endorsed by the Annual Conference, October 2002), Appendix 2. Available at http://www.nccnsw. org.au/data/files/general/flying_foxes_policy2002.rtf (2004). Ng, F.S.P. 1992. Manual of forest fruits, seeds and seedlings. Malayan For. Rec. 34: 505–510. Peekel, P.G. 1984. Flora of the Bismarck Archipelago for Naturalists (trans. E.E. Henty). Division of Botany, Lae: 431. Pennington, T.D. 1991. The Genera of Sapotaceae. Royal Botanic Gardens, Kew, and New York Botanical Garden, Bronx. Pennington, T.D. 2004. Sapotaceae. Pp. 390–421 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants VI. Springer-Verlag, Berlin. Schlechter, R. 1911. Die Guttapercha- und Kautschuk-Expedition des KolonialWirtschaftlichen Komitees nach Kaiser Wilhelmsland 1907–1909. Kol. Wirtsch. Kom., Berlin. Stier, S.C. 1990. Dietary habits of two threatened co-roosting flying foxes (Megachiroptera), Subic Bay, Philippines. M.A. thesis, University of Montana, Montana. Sterly, J. 1997. Simbu Plant-Lore, vol. II. Reimer Verlag, Berlin. Stocker, G.C., and A.K. Irvine. 1983. Seed dispersal by Cassowaries (Casuarius casuarius) in Northern Queensland’s rainforests. Biotropica 15: 170–176. Subba Reddi, C., and A.J. Bai. 1980. Floral biology of Mimusops elengi Linn. J. Bombay Nat. Hist. Soc. 77: 471–475. Taylor, B.W. 1964. Vegetation of the Buna-Kokoda area. Pp. 89–98 in Haantjens, H.A. (ed.) General Report on Lands of the Buna-Kokoda Area, Territory of Papua and New Guinea. Land Research Series, 10.

................. 16157$

CH32

02-08-07 10:37:11

PS

PAGE 471

472 / van der Pijl, L. 1957. The dispersal of plants by bats (chiropterochory). Acta Bot. Neerl. 6: 291–315. van Royen, P. 1957. Revision of the Sapotaceae of the Malaysian area in a wider sense. VII. Planchonella Pierre. Blumea 8: 235–445. van Royen, P. 1960. Revision of the Sapotaceae of the Malaysian area in a wider sense. XXIII. Palaquium Blanco. Blumea 10: 432–606. Vink, W. 1998. Notes on some lowland rainforests of the Bird’s Head peninsula, Irian Jaya. Pp. 91–109 in Bartstra, G.-J. (ed.) Bird’s Head Approaches. Balkema, Rotterdam. Vink, W. 2002. Some malesian species of Pouteria (Sapotaceae). Blumea 47: 95–147. Wiselius, S.I. 1998. Chrysophyllum L. Pp. 162–164 in Sosef, M.S.M., L.T. Hong, and S. Prawirohatmodjo (eds.) Plant Resources of South-East Asia, 5 (3), Timber Trees: Lesser Known Timbers. Backhuys, Leiden. Zieck, J.F.U. 1959. Bosverkenning van het Tami-kustplateau en de Tami-Mossovlakte (O.Afd. Hollandia) 1955–1956. 2nd ed. Bosplanalogie, Manokwari, mimeo. Zieck, J.F.U. 1960. Hydrologische bosreserve ‘‘Tafelberg’’ te Manokwari. Bosplanalogie, Manokwari, mimeo.

................. 16157$

CH32

02-08-07 10:37:11

PS

PAGE 472

Zingiberaceae of Papua . Number of Genera and Species ca 52 genera and 1,300 species of Zingiberaceae worldwide (Larsen et al. 1998). In New Guinea there are six genera, Alpinia, Amomum, Etlingera, Hornstedtia, Pleuranthodium, and Riedelia, all of them found both in Papua and in Papua New Guinea. Curcuma, Globba, and Zingiber are probably not native but some species are cultivated or naturalized, especially ginger, Zingiber officinale Roscoe, and turmeric, Curcuma longa L. It is difficult to give an accurate estimate of the number of species. Newman et al. (2004) list ca 240 names, ca 140 in Papua and ca 130 in Papua New Guinea, but most of them are only provisionally accepted. With new discoveries, it looks likely that there will be more than 300 species found on the island.

T

Distribution and Habitat Forty-six of the genera and the vast majority of species are Asiatic, being found from Sri Lanka to the western Pacific. Aframomum, Aulotandra, and Siphonochilus occur only in Africa, including Madagascar in the case of Aulotandra, while Renealmia is the only genus found in two continents, namely tropical America and Africa. The northern limit of the family follows a line from the Himalayas through southern China to Japan where Alpinia japonica (Thunb.) Miq. occurs just north of Tokyo. To the south, Alpinia caerulea (R. Br.) Benth. reaches almost as far south as Sydney in northern New South Wales and to the east there are several species in Fiji. Most Zingiberaceae grow in the understory of tropical forests though all but two species of Siphonochilus are found in savannas in Africa (Poulsen and Lock 1999). Many continental Asiatic species are geophytes of seasonal forests which die down to the underground rhizome during the dry season but almost all species of the Malay Archipelago are evergreen. The highest altitudes are reached by certain species of Roscoea in the Himalayas and China which have been found at around 4,000 m, while Riedelia montana Valeton var. goliathensis Valeton has been collected at 3,200 m in New Guinea. The earliest fossils of Zingiberaceae date to the Upper Cretaceous of North America where leaf fossils in the genus Zingiberopsis have been found (Benton 1993). Fruit fossils dating to the Upper Eocene are also known from North America and Europe. However, there are no fossil records from areas in which the Zingiberaceae grow now. Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

473

................. 16157$

CH33

02-08-07 10:37:43

PS

PAGE 473

.

474 /

Family Classification The Zingiberaceae have long been recognized as a highly natural group. There have been changes in the rank at which certain groups in the Zingiberales have been classified but the groups themselves have not changed much in the last 150 years. Several classifications of the genera of Zingiberaceae exist, all of them placing great emphasis on the size of the lateral staminodes. Kress et al. (2003) have recently produced a classification that is supported by molecular and morphological evidence. This divides the family into four subfamilies, two of which contain a single genus; the two larger subfamilies are then divided into two tribes, as follows: subfamily Siphonochiloideae W.J. Kress (Siphonochilus only); subfamily Tamijioideae W.J. Kress (Tamijia only); subfamily Zingiberoideae Blume ex Hassk., including tribe Globbeae Meisn. (Gagnepainia, Globba, Hemiorchis) and tribe Zingibereae Meisn. (ca 26 genera); and subfamily Alpinioideae Link, including tribe Riedelieae W.J. Kress (Burbidgea, Pleuranthodium, Riedelia, Siamanthus), and tribe Alpinieae A. Rich. (ca 17 genera).

Features of the Family Habit: herbs, sometimes very robust, containing essential oils. Leaves composed of clasping sheaths, ligule and blade, the sheaths forming pseudostems; ptyxis supervolute, venation pinnate. Inflorescences terminal on leafy shoots or on a separate, leafless shoot, spicate, racemose or made up of cincinni borne in the axils of bracts. Flowers zygomorphic, bisexual or functionally male, calyx fused with three more or less distinct teeth; corolla fused with three lobes; labellum oblong to lobed; lateral staminodes present at base of labellum, small or large and petaloid; one fertile stamen opposite the labellum, anther with two thecae; gynoecium syncarpous, the ovary inferior with three locules, style passing between the anther thecae but not fused to them, stigma punctate; placentation axile, ovules few to many per locule, arillate, anatropous. Fruits fleshy or dry, indehiscent or (especially Pleuranthodium and Riedelia) dehiscent. Seeds several to many per locule; cotyledon single.

Gaps in Floristic Documentation or Knowledge It is often difficult to collect gingers for the herbarium, especially the large, ground-flowering species, so collectors tend to pass them by. Thus, the Zingiberaceae are undercollected, even by comparison with other plant families of New Guinea. Clear advice on how to make good herbarium specimens was published by Burtt and Smith (1976) but there have been few collectors in New Guinea since then. Little has been published on the taxonomy of gingers in New Guinea since Schumann’s monograph of the family (1904). At that time New Guinea was divided into three colonies, Dutch in the west, German in the northeast, and British

................. 16157$

CH33

02-08-07 10:37:44

PS

PAGE 474

Angiosperms: Zingiberaceae of Papua / 475

in the southeast. A particular problem relates to the former German colony. Collectors such as Lauterbach and Schlechter sent their material to Schumann in Berlin. Much of this was lost in a fire in 1943, though a few duplicates have been found at BO, WRSL (formerly Breslau), and other places. Without type material it is very difficult to determine new material and to judge species limits. A great deal of fieldwork will be required before neotypes can be chosen for these names. Not long after Schumann, Valeton, working at Herbarium Bogoriense, described a large number of new taxa (Valeton 1913, 1914). Since then, however, progress has been slow and intermittent. Some revision of the genera has occurred (Smith, 1986a,b, 1991) but a new revision of the species is still awaited.

Natural History Studies of reproductive biology of gingers in China (Gao et al. 2004; Zhang et al. 2003; Li et al. 2002) and Borneo (Sakai et al. 1999) have shown that most species are pollinated by bees and spiderhunter birds. No such studies have been carried out in New Guinea.

Literature Cited Benton, M.J. (ed.). 1993. The Fossil Record 2. Chapman & Hall, London. Burtt, B.L., and R.M. Smith. 1976. Notes on the collection of Zingiberaceae. Flora Males. Bull. 29: 2599–2601. Gao, J.-Y., L. Zhang, X.-B. Deng, P.-Y. Ren, J.-L. Kong, and Q.-J. Li. 2004. The floral biology of Curcumorpha longiflora (Zingiberaceae): a ginger with two-day flowers. Amer. J. Bot. 91: 289–293. Kress, W.J., L.M. Prince, W.J. Hahn, and E.A. Zimmer. 2001. Unravelling the evolutionary radiation of the families of the Zingiberales using morphological and molecular evidence. Syst. Biol. 50: 926–944. Kress, W.J., L.M. Prince, and K.J. Williams. 2003. The phylogeny and a new classification of the gingers (Zingiberaceae): evidence from molecular data. Amer. J. Bot. 89: 1682–1696. Larsen, K. 1998. Costaceae. Pp. 128–132 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants, vol. IV. Flowering Plants. Monocotyledons. Alismatanae and Commelinanae (except Gramineae). Springer-Verlag, Berlin. Larsen, K., J.M. Lock, H. Maas, and P.J.M. Maas. 1998. Zingiberaceae. Pp. 474–495 in Kubitzki, K. (ed.) The Families and Genera of Vascular Plants, vol. IV. Flowering Plants. Monocotyledons. Alismatanae and Commelinanae (except Gramineae). SpringerVerlag, Berlin Li, Q.-J., W.J. Kress, Z.-F. Xu, Y.-M. Xia, L. Zhang, X.-B. Deng, and J.-Y. Gao. 2002. Mating system and stigmatic behaviour during flowering of Alpinia kwangsiensis (Zingiberaceae). Plant Syst. Evol. 232: 123–132. Newman, M.F., A. Lhuillier, and A.D. Poulsen. 2004. Checklist of the Zingiberaceae of Malesia. Blumea, Suppl. 16. Poulsen, A.D., and J.M. Lock. 1999. A review of African forest Zingiberaceae. Pp. 51–64 in Timberlake, J., and S. Kativu (eds.) African Plants. Biodiversity, Taxonomy, and Uses.

................. 16157$

CH33

02-08-07 10:37:44

PS

PAGE 475

.

476 /

Proceedings of the 1997 AETFAT Congress, Harare, Zimbabwe. Royal Botanic Gardens, Kew. Sakai, S., M. Kato, and T. Inoue. 1999. Three pollination guilds and variation in floral characteristics of Bornean gingers (Zingiberaceae and Costaceae). Amer. J. Bot. 86: 646–658. Schumann, K.M. 1904. Zingiberaceae. Pp. 1–458 in Engler, A. (ed.) Das Pflanzenreich IV. 46 (Heft 20), Leipzig. Smith, R.M. 1986a. Etlingera: the inclusive name for Achasma, Geanthus and Nicolaia (Zingiberaceae). Notes RBG Edinburgh 43: 235–241. Smith, R.M. 1986b. New combinations in Etlingera Giseke (Zingiberaceae). Notes RBG Edinburgh 43: 243–254. Smith, R.M. 1991. Pleuranthodium replaces the illegitimate Psychanthus (Zingiberaceae). Edinburgh J. Bot. 48: 63–68. Valeton, T. 1913. Zingiberaceae. Nova Guinea 8: 923–988. Valeton, T. 1914. Die Zingiberaceen Deutsch-Neu-Guineas. Bot. Jahrb. Syst. 52: 40–100. Zhang, L., Q.-J. Li, X.-B. Deng, P.-Y. Ren, and J.-Y. Gao. 2003. Reproductive biology of Alpinia blepharocalyx (Zingiberaceae): another example of flexistyly. Plant Syst. Evol. 241: 67–76.

................. 16157$

CH33

02-08-07 10:37:45

PS

PAGE 476

section four 

The Fauna

................. 16157$

PRT4

04-13-07 13:50:09

PS

PAGE 477

................. 16157$

PRT4

02-08-07 10:21:55

PS

PAGE 478

4.1. Introduction to the Fauna of Papua a rich and diverse fauna that includes at least 3,764 vertebrates and probably more than 200,000 invertebrates. To put this fauna into a wider perspective, I first compare it, and its various taxonomic components, to the faunas of the whole island of New Guinea and that of the entire world. I then review the status of our knowledge of the fauna of the New Guinea region, examine geographic patterns of endemism within Papua, and conclude with a brief discussion of the need for improved understanding and documentation of the fauna. In this treatment Papua includes the western half of New Guinea and satellite islands, including the Raja Ampat group, the islands in Cenderawasih Bay and the Aru Islands. Although politically part of Maluku, the Aru Islands are biogeographically most similar to New Guinea and are usually included in treatments of that biota (e.g., Mittermeier et al. 2002). I treat New Guinea as the entire island together with satellite islands, including the Aru Islands but excluding the Bismarck and Admiralty archipelagos. The 3,764 vertebrate species known from Papua constitute 81% of the vertebrates known from the island of New Guinea (Table 4.1.1). Marine fishes make up 62% of the Papuan fauna (Figure 4.1.1). Freshwater and brackish fish species comprise nearly 8%. Birds make up nearly 15% of Papuan vertebrates, followed by amphibians and reptiles at nearly 10%. Mammals comprise 5% of the total. A comparison of the 4,665 vertebrates currently known from the island of New Guinea shows a comparable breakdown in percentages by class. If we focus on land and freshwater vertebrates (i.e., exclude marine and brackish water fishes as well as other exclusively marine vertebrates such as sea turtles and sea snakes), from Papua there are 1,240 species known but only 250 of these (20%) are endemic (Table 4.1.2). In comparison, from New Guinea as a whole there are 1,647 land and freshwater vertebrates known; 1,130 (69%) of these are endemic. The level of endemism in New Guinea ranges from a low of 49% for snakes, which tend to have large geographic ranges, to a high of 92% for frogs, which have limited dispersal capacity and are represented by a high percentage of restricted range species (Stuart et al. 2004). Most native species that are not endemic to Papua are endemic to the island of New Guinea (i.e., shared with Papua New Guinea); most remaining nonendemic species are shared with Australia and to a lesser extent with Southeast Asia. New Guinea vertebrates comprise nearly 8% of currently recognized world vertebrates and range from about 4% of the world total for snakes, lizards, and mammals, to a high of nearly 11% for fishes (Table 4.1.1). If we restrict the comparison

P

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

479

................. 16157$

CH34

02-08-07 10:37:52

PS

PAGE 479

480 /

Table 4.1.1. Vertebrate fauna of Papua, New Guinea, and the world

Taxon

No. of species in Papua

No. of species in New Guinea

No. of species in the world

Fishes

2,650

3,200

30,000

Frogs

130

282

Turtles

15

17

Crocodiles

Papua as % of New Guinea

Papua as % of world

New Guinea as % of world

82.8

8.8

10.7

5,067

46.1

2.6

5.6

307

88.2

4.6

5.2

2

2

23

100.0

8.7

8.7

Lizards

141

193

4,765

73.6

3.0

4.1

Snakes

83

109

2,978

76.1

2.8

3.7

552

578

9,702

95.5

5.7

6.0

Birds Mammals Total

191

284

5,338

67.3

3.6

5.3

3,764

4,665

58,180

80.7

6.5

8.0

Note: Mammal totals do not include cetaceans. Birds figure includes breeding land birds only. Source: Data for fishes based on Allen (1991, Ch. 4.8), Allen and Swainson (1992), and Eschmeyer (1998). Data for amphibians and reptiles from a database maintained at Bishop Museum and are current as of September 2005. World totals for amphibians and reptiles based on Frost (2005) and Uetz (2005), respectively. Data for birds from Dumbacher and Mack (Ch. 4.9), and Beehler and Finch (1985). Mammal data from Helgen (Ch. 4.10), the Hotspots database maintained by Conservation International, and Wilson and Reeder (2005).

to land and freshwater vertebrates, the New Guinea total drops to about 4% of the world total if we include fishes and 6.5% if fishes are excluded. The overall number of invertebrates in the world and in New Guinea is not known to a level of accuracy that would allow for a meaningful comparison, but details are available for a few groups (Table 4.1.3). The proportion of the world’s species found in New Guinea ranges from 3.3% for Diptera (flies) to 9.4% for Odonata (dragonflies and damselflies). The two best known groups of New Guinea insects, butterflies and aquatic Heteroptera, comprise, respectively, 5.5% and 7.0% of the world totals for these groups. The biogeographic affinities of New Guinea vertebrates are complex and are discussed in detail in the chapters that follow. Repeated land connections during the late Cenozoic facilitated faunal exchange between New Guinea and Australia (Aplin 1993; Walker 1972), resulting in many similarities of the vertebrate faunas of these two areas, particularly at the level of genus and above. This is particularly true for mammals, for which marsupials and monotremes are such important and distinctive components of the fauna, but is also true for many groups of birds and other vertebrates. For example, of the 119 genera of amphibians and reptiles that occur in New Guinea, 56 (47%) also occur in Australia. New Guinea also has a strong affinity with Southeast Asia, but oceanic barriers have always separated New Guinea from that region, as demarcated by Wallace’s Line (Wallace 1859; Schulte 2003). These oceanic barriers have prevented a wide array of vertebrate

................. 16157$

CH34

02-08-07 10:37:53

PS

PAGE 480

Introduction to the Fauna of Papua / 481

Figure 4.1.1. Taxonomic composition of the vertebrate fauna of Papua. Crocodiles are represented by the thin wedge between lizards and turtles. lineages from reaching New Guinea. They seem to have been less of a barrier to plants and invertebrate groups (de Boer and Duffels 1996; Vane-Wright 1991) but the evolutionary history of these groups is also closely intertwined with the geological history of the region. With a land area of 790,000 km2 and mountains exceeding 5,000 m in elevation, New Guinea is the world’s largest and highest tropical island. Its biota is the result of evolutionary diversification in one of the most geologically complex regions of the world. To put this into perspective, it is informative to compare New Guinea to the island of Borneo. That island, with a land area of 743,330 km2, is only slightly smaller than New Guinea, has high mountains ( 4,000 m), and many other similar features. Both islands formed through a complex process of accretion of island arcs, oceanic crustal material, and other tectonic fragments onto a Paleozoic continental core (Pigram and Davies 1987; Hall 2002; Moss and Wilson 1998). Borneo began forming during the Mesozoic and was connected by land to mainland Southeast Asia until at least the Eocene (Hall 2002); it has subsequently been reconnected to that region for varying amounts of time coinciding with periods of lowered sea level during the Tertiary and Quaternary (Voris 2000). New Guinea began forming somewhat later, during the mid-Cenozoic, at the leading edge of the northward-moving Australian plate and is composed of at least 32

................. 16157$

CH34

02-08-07 10:38:22

PS

PAGE 481

482 /

Table 4.1.2. Endemicity of land and freshwater vertebrates of Papua and New Guinea (including satellite islands) Taxon

Papua Province No. of Total endemic no. of % species species endemic

New Guinea No. of Total endemic no. of % species species endemic

Freshwater fishes

64

151

42.4

179

213

84.0

Frogs

69

130

53.1

260

282

92.2

Turtles

1

9

12.5

8

11

72.7

Crocodiles

0

2

0.0

1

2

50.0

Lizards

30

141

21.3

124

193

64.2

Snakes

8

64

12.5

41

84

48.8

Birds

38

552

6.9

324

578

56.1

Mammals

40

191

20.9

195

284

68.7

250

1,240

20.2

1,132

1,647

68.7

Total

Note: Mammal totals do not include cetaceans. Birds figure includes breeding land birds only. Source: Figures for fishes from G. Allen, based on updated totals from Allen (1998). Amphibian and reptile figures from database maintained at Bishop Museum; current as of September 2005. Figures for birds from Dumbacher and Mack (Ch. 4.9), Beehler and Finch (1985), and Wheately (1998). Figures for mammals based on Helgen (Ch. 4.10) and the Hotspots database maintained by Conservation International.

Table 4.1.3. Number of species of selected insect taxa of New Guinea and the world Insect group

Total Number of Species New Guinea World

New Guinea as % of world

Diptera

5,000

3.3

150,000

Odanata

579

6,189

9.4

Aquatic Heteroptera

330

4,746

7.0

Papilionoidea/Hesperoidea

955

17,500

5.5

6,864

178,435

3.8

Total

Source: Figures for Diptera from Evenhuis (1989) and Irwin, Schlinger, and Thompson (2003); New Guinea total updated to include species described since 1989 (N. Evenhuis, pers. comm.). Figures for Odonata from J. van Tol (pers. comm.); include species and non-nominal subspecies. Figures for aquatic Heteroptera from D. Polhemus (pers. comm.). Figures for Papilionoidea/Hesperoidea from d’Abrera (1977) and Parsons (1991, 1998), updated in October 2005 from the Zoological Record.

................. 16157$

CH34

02-08-07 10:38:22

PS

PAGE 482

Introduction to the Fauna of Papua / 483

geological terranes. Many of these involve isolated mountain ranges or lowland basins that have produced pockets of endemism, particularly at the northern margins of the Australian craton (Heads 2001; Pigram and Davies 1987; van Welzen 1997). Borneo, in contrast, is topographically less complex, with wide expanses of lowlands surrounding the central mountains. The occurrence of past land connections between Borneo and continental Asia, the similarity of the climate throughout the region during the Tertiary (BrandonJones 1996), and the absence of much topographic relief, has resulted in the fauna of Borneo being rather similar to that of other parts of Southeast Asia. New Guinea, by contrast, with its more complex geological history, its relative isolation from continental areas with similar climates, its greater topographic diversity compared to Borneo, and the impact of these factors on the evolutionary diversification of species (Dow 1977; Hall 2002; Hamilton 1979; Kroenke 1984, 1996; Chapter 4.6) has developed a richer and more endemic biota than has Borneo. However, Borneo is generally richer in higher taxa (e.g., families and orders). There are 11 orders and at least 219 species of mammals from Borneo; 46 (21%) species are endemic (Payne 1985; updated). New Guinea has six orders of mammals (monotremes, three orders of marsupials [Dasyuromorphia, Peramelemorphia, and Diprotodontia], rodents, and bats) with 284 species, of which 195 (69%) are endemic. A total of 434 species of breeding birds are known from Borneo but only 39 of these are endemic (Smythies 2000). New Guinea in contrast has 578 species of breeding birds, of which 324 species are endemic (Beehler and Finch 1985; Beehler, Pratt, and Zimmerman 1986; Wheatley 1998). Borneo has a higher number of families and genera of amphibians and reptiles than does New Guinea and, in addition, has caecilians, an amphibian order that is absent from New Guinea. At the species level, Borneo also has a higher number of snakes than does New Guinea, reflecting the importance of this group in Southeast Asia, but Borneo has far fewer frog and lizard species and a much lower overall total number of amphibian and reptile species than does New Guinea (Table 4.6.2). Borneo and New Guinea have a similar number of freshwater fishes, about 400 species (Kottelat et al. 1993; Chapter 4.8) but about a quarter of the freshwater ichthyofaunas of both areas are composed of fishes with a marine larval stage (Chapter 4.8). Borneo is dominated by primary division fishes (species belonging to families that have always been intolerant of salt water); about 40% of these species are endemic. In contrast, New Guinea essentially lacks primary division fishes and most species of its freshwater fishes are thought to have evolved from marine ancestors; approximately 180 (84%) of currently recognized New Guinea freshwater species lacking a marine larval stage are endemic. In general, species richness in Borneo tends to be higher within a drainage basin than in New Guinea. For example, the large Kapuas River system in western Kalimantan has 320 species (Roberts 1989; Kottelat and Whitten 1996). By contrast the Fly River drainage in New Guinea has fewer than 110 species (Roberts 1978). The New Guinea region is estimated to harbor around 2,600–3,000 species of

................. 16157$

CH34

02-08-07 10:38:23

PS

PAGE 483

484 /

marine fishes (Allen and Swainson 1992), including 30% of the world’s reef fishes (Myers 1999; Pyle 1995); Borneo likely has a comparable number of marine species. Many of the marine species are widespread in the Indo-Pacific region and are found in both Borneo and New Guinea. Species richness of the marine biota of the Indo-Australian region is among the highest is the world (Roberts et al. 2002).

Status of the Fauna It is likely that most species of resident birds are known for New Guinea; the last currently recognized species was described in 1959 and a graph of the cumulative number of species treated by Beehler and Finch (1985) has reached an asymptote (Figure 4.1.2). Our knowledge of other groups, however, is far from complete. The mammals, particularly the bats and some groups of rodents, are incompletely known and about ten new species were described during the past decade. Ongoing systematic studies by K. Helgen, T. Flannery, G. Musser, T. Leary, and others is likely to significantly increase the number of species based on material currently in collections. A cumulative graph of species (Figure 4.10.1) demonstrates that only about half the currently known species had been described by 1905. During the past five years three species have been described from Papua and descriptions of at least seven more Papuan species are in preparation (K. Helgen, pers. comm.; T. Leary, pers. comm.). The frogs are probably the most poorly known group of New Guinea vertebrates and the current New Guinea total of 282 species will almost certainly double

Figure 4.1.2. Cumulative number of currently recognized New Guinea bird species. All breeding birds and regular migrants are included. Source: Beehler and Finch (1985); Beehler, Pratt, and Zimmerman (1986).

................. 16157$

CH34

02-08-07 10:38:41

PS

PAGE 484

Introduction to the Fauna of Papua / 485

and could even triple when all species have been scientifically named. Seventyseven new species of frogs have been described during the past decade (see Figure 4.1.3) and at least 150 additional species in collections await description. Each major herpetofaunal survey during the past five years has turned up five to ten and frequently more new species in single drainage basins. Approximately 28 (60%) of the frogs collected during a survey of the Wapoga River drainage in northwest Papua proved to be undescribed species (Richards, Iskandar, and Allison 2000). On a broader scale, during a series of surveys of Milne Bay Province, one of the better known areas of Papua New Guinea, Kraus and Allison (2004a,b,c, 2005, in prep) have discovered at least 38 new species of frogs. As for the reptiles, the lizards are somewhat better known than frogs and only 12 new species have been described during the past decade (Figure 4.1.3). However, descriptions of at least eight new species are currently in press and ongoing studies of the large genus Sphenomorphus by G. Shea is likely to result in the recognition and description of many additional species. Only four new species of snakes have been described during the past decade (Figure 4.1.3) but at least eight new species in collections await description and it is likely that the total number, which is currently 109, will ultimately increase to about 140. The number of invertebrates known from New Guinea is not known with any accuracy and there are no comprehensive checklists of most groups. Miller (Chapter 4.3) estimates that there are slightly more than 300,000 terrestrial species. This is very likely a conservative estimate and the actual number could be far higher, perhaps 500,000 or more. In the best known group, butterflies (d’Abrera 1977; Parsons 1991, 1998), the descriptions of the new species appears to be reaching an

Figure 4.1.3. Cumulative number of currently recognized New Guinea frog, lizard, and snake species. Source: Database maintained at Bishop Museum; includes all species described to September 2005.

................. 16157$

CH34

02-08-07 10:38:59

PS

PAGE 485

486 /

asymptote (Figure 4.1.4) but it is likely that fewer than 50% of species have been described in most insect orders. Faunal survey work in Papua has greatly lagged that in Papua New Guinea. As a rough indication of this, the number of museum specimens from poorly known groups from Papua is only a quarter, or less, of the number available from Papua New Guinea. This is apparent in Brown’s (1991) monograph of the lizard genus Emoia, which is represented in New Guinea by 36 species. Brown borrowed material from 34 museums throughout the world and endeavored to obtain specimens for as many different species and localities as possible. He ultimately examined 1,605 lots (8,648 specimens) from Papua New Guinea but was able to obtain only 316 lots (1,215 specimens) from Papua. For the combined Papua/Papua New Guinea area only 21% of the lots and 12% of the specimens were from Papua. A quick search of catalogs of museums in the United States that have herpetology collections with more than 10,000 New Guinea specimens showed that the ratio of Papua to Papua New Guinea specimens (amphibians and reptiles) was on the order of 1 to 40. This, of course, does not include museums such as the National Museum of Natural History at Leiden, Zoological Museum Amsterdam, or the Museum Zoologense Bogoriense which have significant collections from Papua. If we restrict analysis to the 18 museums from which Brown (1991) obtained material from Papua for his monograph on Emoia, we obtain a ratio (for Emoia specimens) of around one specimen from Papua for every seven specimens from Papua New Guinea. It is probably safe to conclude that at a world level, the number of museum herpetological specimens available from Papua New Guinea

Figure 4.1.4. Cumulative number of currently recognized New Guinea butterfly species. Source: Parsons (1998); updated from the literature to September 2005.

................. 16157$

CH34

02-08-07 10:39:21

PS

PAGE 486

Introduction to the Fauna of Papua / 487

is probably at least seven to ten times greater than the amount of material available from Papua. The situation appears to be similar for mammal specimens. For example, Taylor, Calaby, and van Deusen (1982) obtained specimens from about 275 sites on New Guinea (including the Aru Islands and satellite islands) but only 73 (27%) of these were from Papua. Similarly, the number of specimens they examined was strongly skewed to Papua New Guinea. There are no compelling reasons to expect that species richness in Papua is much different from species richness in the eastern half of New Guinea. Both halves of New Guinea have accreted terranes along the north coast and have complex composite terranes such as the Vogelkop (Bird’s Head) in Papua and the southeastern peninsula of Papua New Guinea (East Papua Composite Terrane). Both areas have isolated mountain ranges and extensive topographic relief. And both areas are also fringed by satellite islands, some of which are themselves centers of endemism. Indeed, if we look at the best-known group, birds, both halves of New Guinea have similar numbers of species (Coates 1985; Chapter 4.9). However, for the more poorly known groups (e.g., amphibians and reptiles) it is clear that the lower number of species known from Papua and the paucity of specimens from there, compared to Papua New Guinea, both indicate a crucial need for fieldwork and associated taxonomic studies. It is especially important to note that there are large geographic gaps in Papua where there has been little or no fauna survey work (Allison, Kraus, and McShane 2004). For example, there are essentially no specimens of amphibians or reptiles from the Foja or Kumawa mountains, both areas that, based on geological history and geographic isolation, are likely to be rich in endemic species.

Patterns of Endemism Identifying areas of endemism and high species richness in Papua helps us to understand overall faunal distribution patterns and to associate these patterns with past geological and other events. This, in turn, helps us to understand the evolution of the Papuan fauna and to identify potential areas that require formal conservation protection. This information is also crucial for determining priorities for field surveys. The 1997 Conservation Priority-setting Workshop (CPSW; Conservation International 1999) identified areas of endemism for various taxonomic groups. For amphibians and reptiles (Allison 1998; Conservation International 1999), a total of 19 areas were identified that included islands, isolated mountain ranges/terranes, drainage basins, and woodland savanna as follows: islands: Batanta, Waigeo, Numfoor, Biak/Supiori, and Yapen; isolated mountain ranges/terranes: Charles Louis Mts, Weyland Mts, Kumawa Mts, Fakfak Mts, Tamrau and Arfak Mts, Wandammen Peninsula, Van Rees Mts, Foja Mts, Jayawijaya Mts, Cyclops Mts; drainage basins: Digul River, Lorentz National Park, Mamberamo River; seasonal dry forest: southern savannas.

................. 16157$

CH34

02-08-07 10:39:21

PS

PAGE 487

488 /

These CPSW areas, with the addition of the Aru Islands (Figure 4.6.1), are inhabited by at least 85% and probably up to 90% of the 107 amphibian and reptile species that are endemic to the Aru Islands-Papua region and for which the distribution is known (Chapter 4.6). The scheme proposed at the CPSW for mammals is very similar to that proposed for amphibians and reptiles, except that it includes the western versant of the Jayawijaya Mts (Sudirman Mts) and a large area of the southern Vogelkop region (the ‘‘Bird’s Neck’’). The scheme proposed for birds is mostly similar with respect to islands, except that Yapen, which lacks endemic birds, is excluded (Chapter 2.4). It differs somewhat from other schemes in that entire mountain ranges are included, not just parts of ranges as for amphibians and reptiles and mammals. It also differs from the plan for amphibians and reptiles by including large lowland areas around Bintuni and Arguni bays. The CPSW scheme for mammals is similar to that for birds in that it includes an area on the southern parts of the Bird’s Neck (Arguni, Etna, and Triton bays). It differs from the scheme for birds in not including an area around Bintuni Bay (Chapter 4.10). Polhemus and Allen (Chapter 2.5) analyzed the distribution of freshwater fauna and identified 23 areas in Papua, including five that are shared with Papua New Guinea, as centers of species richness and endemism. There is considerable agreement between this scheme and the CPSW schemes. There are essentially no differences in the selection of satellite islands except that Polhemus and Allen include Misool within the group of important islands. There are at least 25 species of amphibians and reptiles known from Misool but only one of these, the recently described Varanus reisingeri (Eidenmu¨ller and Wicker 2005), is endemic. There are relatively few differences between the two schemes for mountains/ terranes, except that Polhemus and Allen incorporate the Wandammen Peninsula and adjacent mountains into a fairly large Vogelkop anticlines region. This region is particularly rich in freshwater fish (Chapter 4.8) but has not been well surveyed for amphibians and reptiles. There is, however, some evidence (e.g., Zug and Allison 2006) to suggest that the greater Bird’s Neck area may be important for amphibians and reptiles. The southern areas of the Bird’s Neck are included in the mammal and bird schemes. The Mamberamo foreland region designated by Polhemus and Allen largely on the basis of collections from the Third Archbold Expedition in 1939 is certainly an important area for amphibians and reptiles, largely for the same reason. This area would be included in the Jayawijaya region designated by the CPSW. The Arafura Foreland region designated by Polhemus and Allen essentially coincides with the Lorentz National Park region designated by the CPSW. Similarly the Trans-Fly Foreland would correspond to the Digul River drainage designated by the CPSW. Several northern or western lowland areas not recognized as important for amphibians and reptiles were identified as important for aquatic fauna. This includes an area near Jayapura that Polhemus and Allen call the northeast Papuan lowlands. Because of its proximity to Jayapura, this area has been relatively well-

................. 16157$

CH34

02-08-07 10:39:22

PS

PAGE 488

Introduction to the Fauna of Papua / 489

surveyed for aquatic fauna compared to surrounding areas and, as Polhemus and Allen point out, its high level of endemicity may be an artifact that simply reflects a higher level of collecting in this area compared to surrounding regions. Although the Jayapura area has been relatively well collected for amphibians and reptiles, many adjacent lowland areas remain poorly known, although at least one species of frog, Litoria mystax, is thought to occur in this region. The Vogelkop lowlands area designated by Polhemus and Allen was not designated as an endemic area by the CPSW. However, at least two species of lizards are endemic to this area: a gecko, Gehyra leopoldi, and a skink, Lipinia venemai. The southern savanna region designated by the CPSW (for amphibians and reptiles) includes two areas designated by Polhemus and Allen: Arafura coastal lowlands and Trans-Fly coastal lowlands. Both these areas have a complex mosaic of forest and savanna vegetation but the Arafura coastal lowlands tends to be dominated by closed canopy forest and the Trans-Fly coastal lowlands have a greater dominance of woodland and savanna. The distribution of amphibians and reptiles in the overall region is too poorly documented to determine whether these vegetation differences are sufficient to warrant the designation of two different areas. Allen (1991) recognized seven zoogeographic regions in New Guinea that reflect distribution patterns of freshwater fish. Five of these of these occur in Papua: 1. Western Islands (Raja Ampat group); 2. Vogelkop Peninsula (mountains and lowlands); 3. Great Northern (includes north coast accreted terranes and river systems and extends across the much of northern Papua New Guinea); 4. Great Southern or Trans-Fly (includes most of southern New Guinea); and 5. the Aru Islands. In his treatment of the fishes of Papua (Chapter 4.8), Allen combines the Western Islands with the Vogelkop regions and includes the Aru Islands within the Great Southern region. The scheme proposed by Polhemus and Allen is essentially based on these same areas but differentiates within regions at a finer scale. Birdlife International has taken a somewhat similar approach to Allen (1991) and has recognized eight endemic bird areas in Papua (Stattersfield et al. 1998). These include for the Vogelkop Peninsula two areas, the west Papuan lowlands and west Papuan highlands, which cover all of western New Guinea and include the Raja Ampat Islands. The islands of Numfoor, Supiori, Biak, and Meos Num are included within the Geelvink (Cenderawasih) Islands area; Yapen is treated as a secondary area to this. The Foja and Cyclops Mts (together with the Bewani, Torricelli, and Prince Alexander mountains in Papua New Guinea) are included in the north Papuan mountains area. The lowlands of northern Papua drained by the Mamberamo River together with the Sepik, Ramu, and Markham drainage systems in Papua New Guinea are included in the north Papuan lowlands area. The central mountains, from the Weyland and Charles Louis mountains in the west to the Owen Stanley Mts in eastern Papua New Guinea are included in the large central Papuan mountains area. Southern watersheds from the Mimika River east to the upper Fly and entire Kikori and Purari drainages are included in the south Papuan lowlands areas. The area to the south of this, which has a strongly

................. 16157$

CH34

02-08-07 10:39:23

PS

PAGE 489

490 /

seasonal climate and is dominated by woodland savanna vegetation, is included in the Trans-Fly area, with the Aru Islands included as a secondary area to this. These eight areas include essentially all the land area in Papua and represent a classification system that is similar to the others in recognizing the importance to endemism of lowland basins, mountains, and islands, many of which represent separate terranes. The only significant differences between the endemic areas schemes proposed by the CPSW for amphibians and reptiles, birds, and mammals and the schemes proposed by Polhemus and Allen (Chapter 2.5), Allen (1991), and Birdlife International (Stattersfield et al. 1998), is the extent to which the various schemes subdivide areas of importance. These schemes all agree on the importance of the various major regions and recognize important differences between the north and south coasts and the Vogelkop in the west. They differ slightly on the boundaries of various areas. These differences probably reflect real differences in the biological characteristics of different taxa (e.g., birds generally have larger geographic ranges than do amphibians and reptiles) and also different levels of knowledge (e.g., bird species distributions are far better known than the ranges of other vertebrate taxa).

Future Considerations A persistent belief among biologists, conservation professionals, and others who work outside of the New Guinea region is that the biota of this region, while interesting, is comparatively well known and not particularly rich compared to other tropical regions (Pearson 1977; Clarke 2000). As this brief review and other recent work have established, neither assertion is true. New Guinea is clearly one of the megadiverse regions of the world (Mittermeier et al. 2002). It is also one of the most poorly known. As we have gained an understanding of New Guinea’s diversity, we have also come to understand the true dimensions of this diversity and how much additional work is required to discover and scientifically name the biota. At the same time we are mindful that mature forest is being lost at an accelerating rate (Chapter 7.1) due to timber harvesting, land conversion to agriculture, mining, urban development, and other factors. The 1997 Conservation Priority-setting Workshop helped to focus attention on areas of endemism in Papua Province. There is a compelling, indeed crucial, need for a comprehensive biological survey of Papua to better document the biota and help guide and inform the designation of protected areas.

Acknowledgments I thank C. Kishinami, L. Eldredge, K. Helgen, D. Polhemus, G. Allen, S. Miller, N. Evenhuis, J. van Tol, and T. Leary for information. I also thank C. Kishinami, L. Eldredge, and T. Leary for their helpful comments on drafts of the manuscript. Brad Evans assisted with the figures. I would also like to acknowledge financial support from the U.S. National Science Foundation.

................. 16157$

CH34

02-08-07 10:39:24

PS

PAGE 490

Introduction to the Fauna of Papua / 491

Literature Cited Allen, G.R. 1991. Field Guide to the Freshwater Fishes of New Guinea. Christensen Research Institute, Madang. Allen, G.R., and R. Swainston. 1992. Reef Fishes of New Guinea: A Field Guide for Divers, Anglers and Naturalists. Christensen Research Institute, Madang. Allison, A. 1998. Taxa group summary: reptiles and amphibians. Pp. 47–48 in Burnett, J.B., Y. de Fretes, and P. Kramadibrata (eds.) The Irian Jaya Biodiversity Conservation Priority-Setting Workshop: Final Report. Conservation International, Washington, DC. Allison, A., F. Kraus, and M. McShane. 2004. Patterns of species richness in the Papuan region: a preliminary assessment using amphibians and reptiles. Report prepared for The Nature Conservancy. Bishop Museum, Honolulu. Aplin, K.P., P.R. Baverstock, and S.C. Donnellan. 1993. Albumin immunological evidence for the time and mode or origin of the New Guinea terrestrial mammal fauna. Science in New Guinea 19: 131–145. Beehler, B.M., and B.W. Finch. 1985. Species-checklist of the birds of New Guinea. Australasian Ornithological Monographs 1: 1–127. Beehler, B.M., T.K. Pratt, and D.A. Zimmerman. 1986. Birds of New Guinea. Princeton University Press, Princeton, New Jersey. Brandon-Jones, D. 1996. The Asian Colobinae (Mammalia: Cercopithecidae) as indicators of Quaternary climatic change. Biological Journal of the Linnean Society 59 (3): 327–350. Brown, W.C. 1991. Lizards of the genus Emoia (Scincidae) with observations on their ecology and biogeography. Memoirs of the California Academy of Sciences 15: 1–94. Clarke, B. 2000. Review of Patterns of Distribution of Amphibians: A Global Perspective. Duellman, W.E. (ed.) [1999.] Johns Hopkins University Press, Baltimore. Coates, B.J. 1985. The Birds of Papua New Guinea, Including the Bismarck Archipelago and Bougainville. Vol. 1. Dove Publications, Aderley, Queensland. Conservation International. 1999. Peta Lokakarya Prioritax Konservasi di Irian Jaya. Okakarya Penentuan Daerah Prioritas Konservasi Keanekaragaman Hayati Irian Jaya. The Irian Jaya Biodiversity Conservation Priority-Setting Workshop, 7–12 January, Biak, Irian Jaya. Conservation International, Washington, D.C. d’Abrera, B. 1977. Butterflies of the Australian Region. Vol. 1. Butterflies of the World. 2nd ed. Lansdowne Press, Melbourne. deBoer, A.J., and J.P. Duffels. 1996. Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific: a geotectonic explanation. Palaeogeography Palaeoclimatology Palaeoecology 124 (1–2): 153–177. Dow, D.B. 1977. A geological synthesis of Papua New Guinea. Bureau of Mineral Resources, Geology and Geophysics Bulletin, Australian Government Publishing Service, Canberra 201: i–vii, 1–41. Eidenmu¨ller, B., and R. Wicker. 2005. Eine weitere neue Waranart aus dem Varanaus prasinus-Komplex von der Insel Misol, Indonesien. Sauria 27 (1): 3–8. Eschmeyer, W.N. (ed.). 1998. The Catalog of Fishes. 3 volumes. Center for Biodiversity Research and Information. Special Publication 1. California Academy of Sciences, San Francisco, California. Evenhuis, N.E. (ed.). 1989. Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Museum Press and E.J. Brill, Honolulu and Leiden. Frost, D.R. 2005. Amphibian Species of the World: An Online Reference. Version 3.0 (22 August 2004). Electronic database accessible at http://research.amnh.org/herpetology/ amphibia/index.html. American Museum of Natural History, New York.

................. 16157$

CH34

02-08-07 10:39:24

PS

PAGE 491

492 / Hall, R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions model and animations. Journal of Asian Earth Sciences 20: 353–431. Hamilton, W. 1979. Tectonics of the Indonesian region. U.S. Geological Survey Professional Paper 1078: 1–345. Heads, M. 2001. Regional patterns of biodiversity in New Guinea plants. Botanical Journal of the Linnean Society 136 (1): 67–73. Irwin, M.E., E.I. Schlinger, and F.C. Thompson. 2003. Diptera, true flies. Pp. 692–702 in Goodman, S.M., and J.P. Benstead (eds.) The Natural History of Madagascar. University of Chicago Press, Chicago, Illinois. Junk, W.J., and M.G.M. Soares. 2001. Freshwater fish habitats in Amazonia: state of knowledge, management and protection. Aquatic Ecosystems Health and Management 4 (4): 437–451. Kottelat, M., and T. Whitten. 1996. Freshwater Biodiversity in Asia: With Special Reference to Fish. World Bank, Washington, DC. Kottelat, M., T. Whitten, S.N. Kartikasari, and S. Wirjoatmodjo. 1993. Freshwater Fishes of Western Indonesia and Sulawesi. Periplus Editions, Hong Kong. Kraus, F., and A. Allison. 2004a. New records of amphibians and reptiles from Milne Bay Province, Papua New Guinea. Herpetological Review 35 (4): 413–418. Kraus, F., and A. Allison. 2004b. A new species of Litoria (Anura: Hylidae) from southeastern New Guinea. Herpetologica 60 (1): 97–103. Kraus, F., and A. Allison. 2004c. Two new treefrogs from Normanby Island, Papua New Guinea. Journal of Herpetology 38 (2): 197–207. Kraus, F., and A. Allison. 2005. A colorful new species of Albericus (Anura: Microhylidae) from southeastern Papua New Guinea. Pacific Science 59 (1): 43–53. Kroenke, L.W. 1984. Cenozoic tectonic development of the Southwest Pacific. United Nations Economic and Social Commission, Committee for Co-ordination of Joint Prospecting for Mineral Resources in South Pacific Offshore Areas (CCOP/SOPAC), Technical Bulletin 6: 1–122. Kroenke, L.W. 1996. Plate tectonic development of the western and southwestern Pacific Mesozoic to the present. Pp. 19–34 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Mittermeier, R.A., C.G. Mittermeier, P. Robles Gill, J. Pilgrim, G.A.B. Da Fonseca, T. Brooks, and W.R. Konstant (eds.). 2002. Wilderness: Earth’s Last Wild Places. CEMEX, Agrupacio´n Serra Madre, S.C., Mexico. Moss, S.J., and M.E.J. Wilson. 1998. Biogeographic implications of the Tertiary paleogeogramhic evolution of Sulawesi and Borneo. Pp. 133–163 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Myers, R.F. 1999. Micronesian Reef Fishes: A Comprehensive Guide to the Coral Reef Fishes of Micronesia. Coral Graphics, Barrigada, Guam. Parsons, M. 1991. Butterflies of the Bulolo-Wau Valley. Bishop Museum Press, Honolulu. Parsons, M. 1998. The Butterflies of Papua New Guinea: Their Systematics and Biology. Academic Press, San Diego. Payne, J., C.M. Francis, and K. Phillipps. 1985. A Field Guide to the Mammals of Borneo. Sabah Society and World Wildlife Fund Malaysia, Kota Kinabalu, Sabah and Kuala Lumpur, Malaysia. Pearson, D.L. 1977. A pantropical comparison of bird community structure on six lowland forest sites. Condor 79: 232–244.

................. 16157$

CH34

02-08-07 10:39:24

PS

PAGE 492

Introduction to the Fauna of Papua / 493 Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. BMR Journal of Australian Geology 10 (3): 193–211. Pyle, R.L. 1995. Pacific reef and shore fishes. Pp. 205–238 in Maragos, J.E., M.N.A. Peterson, L.G. Eldredge, J.E. Bardach, and H.F. Takeuchi (eds.) Marine and Coastal Biodiversity in the Tropical Pacific Region. Vol. 1. Species Systematics and Information Management Priorities. Program on Environment, East-West Center, Honolulu. Richards, S., D.T. Iskandar, and A. Allison. 2000. Amphibians and reptiles of the Wapoga River area, Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14: 54–57, 113–120. Robbins, R.K. 1982. How many butterfly species? News of the Lepidopterists’ Society 1982: 40–41. Roberts, C.M., C.J. McClean, J.E.N. Veron, J.P. Hawkins, G.R. Allen, D.E. McAllister, C.G. Mittermeier, F.W. Schueler, M. Spalding, F. Wells, C. Vynne, and T.B. Werner. 2002. Marine biodiversity hotspots and conservation priorities for tropical reefs. Science 295 (5558): 1280–1284. Roberts, T.R. 1978. An ichthyological survey of the Fly River in Papua New Guinea with descriptions of new species. Smithsonian Contributions to Zoology 281: 1–70. Roberts, T.R. 1989. The freshwater fishes of western Borneo (Kalimantan Barat, Indonesia). Memoirs of the California Academy of Sciences 14: 1–210. Schulte, J.A., J. Melville, and A. Larson. 2003. Molecular phylogenetic evidence for ancient divergence of lizard taxa on either side of Wallace’s Line. Proceedings of the Royal Society of London Series B-Biological Sciences 270 (1515): 597–603. Smythies, B.E. 2000. The Birds of Borneo [Revised by G.W.H. Davison]. Natural History Publications (Borneo), Kota Kinabalu, Sabah. Stattersfield, A.J., M.J. Crosby, A.J. Long, and D.C. Wege. 1998. Endemic Bird Areas of the World: Priorities for Biodiversity Conservation. BirdLife Conservation Series No. 7. BirdLife International, Cambridge. Stuart, S.N., J.S. Chanson, N.A. Cox, B.E. Young, A.S.L. Rodrigues, D.L. Fischman, and R.W. Waller. 2004. Status and trends of amphibian declines and extinctions worldwide. Science 306: 1783–1786. Taylor, J.M., J.H. Calaby, and H.M. van Deusen. 1982. A revision of the genus Rattus (Rodentia, Muridae) in the New Guinea region. Bulletin of the American Museum of Natural History 173 (3): 177–336. Uetz, P. 2005. The EMBL Reptile Database: http://www.embl-heidelberg.de/ uetz/ LivingReptiles.html (June 2004). Vane-Wright, R.I. 1991. Transcending the Wallace line: do the western edges of the Australian region and the Australian Plate coincide? Australian Journal of Botany 4: 183–197. van Welzen, P.C. 1997. Increased speciation in New Guinea: tectonic causes. Pp. 363–387 in Dransfield, J., M.J.E. Coode, and D.A. Simpson (eds.) Plant Diversity in Malesia III: Proceedings of the Third International Flora Malesiana Symposium 1995. Royal Botanic Gardens, Kew. Voris, H.K. 2000. Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. Journal of Biogeography 27 (5): 1153–1167. Walker, D., and Australian National University, Research School of Pacific Studies. 1972. Bridge and Barrier: The Natural and Cultural History of Torres Strait. Australian National University, Canberra. Wallace, A.R. 1859. Letter from Mr. Wallace concerning the geographical distribution of birds. Ibis 1: 449–454.

................. 16157$

CH34

02-08-07 10:39:25

PS

PAGE 493

494 / Wheatley, N. 1998. Where to Watch Birds in Australasia and Oceania. Princeton University Press, Princeton, New Jersey. Wilson, D.E., and D.M. Reeder. 2005. Mammal Species of the World: A Taxonomic and Geographic Reference. Johns Hopkins University Press, Baltimore. Zug, G.R., and A. Allison. 2006. New Carlia fusca complex lizards (Squamata: Scincidae) from New Guinea, Papua-Indonesia. Zootaxa 1237: 27–44.

................. 16157$

CH34

02-08-07 10:39:25

PS

PAGE 494

4.2. Marine Invertebrates of Papua . of the earth’s surface from the east coast of Africa to Hawaii, the Indo-West Pacific Ocean is well known for its incredible diversity of species. The biota inhabiting coral reefs systems in the Indo-West Pacific is so diverse that it exceeds that of the next three richest areas combined (eastern Pacific, western Atlantic, and eastern Atlantic; Briggs 1999). Within the Indo-West Pacific the ‘‘Coral Triangle,’’ extending roughly from the Philippines through Malaysia to Papua New Guinea, is the center of the megadiversity (Figure 5.2.4). Diversity declines in all directions with increasing distance from the center. To the north and south, diversity decreases with increasing latitude. In tropical areas within the Indo-Pacific, diversity falls off more sharply to the east in the Pacific Ocean than it does to the west in the Indian Ocean (Chapter 5.2). Lying on the western half of the island of New Guinea, Papua is well within the center of coral triangle megadiversity. Papua has considerable diversity across a large number of invertebrate phyla. There is a wealth of information about the broad distribution patterns of some groups. For example, Veron (2000a) provides distribution maps for almost all of the known coral species. Among mollusks, publications on families favored by shell collectors provide similar distribution maps; for example, cones were surveyed by Ro¨ckel et al. (1995) and cowries by Burgess (1985). However, these are generalized maps, which are based on relatively few museum records. They assume that if a species occurs to the east and west of an area such as Papua, the species also occurs in the intervening area, in this case Papua. While this may be true as a generalization, it does risk missing genuine distribution gaps. There have been very few detailed surveys of invertebrates at localities anywhere in the coral triangle, and the total diversity of most groups is unknown. The best data are for coral reefs, but there is no compilation of all species inhabiting a single reef. Even estimates for the total number of species on reefs vary widely. Reaka-Kudla (1997) estimated that 93,000 species of plants and animals have been described from coral reefs, but suggested this may represent only 1–15% of the total number of species present. If the fauna of reef systems is poorly known, the invertebrates of mangroves, the other key habitat in the region, are even less well known. The present paper deals with the benthic invertebrates of coral reefs and mangroves in Papua, the key shallow-water habitats in the area. Sandy shores have a depauperate biota, and rocky shores often have corals from the lower intertidal and subtidal. Deep-water habitats are not included for two reasons: firstly, there is very little information on invertebrates in these areas and, secondly, conservation issues are largely concentrated in shallow-water habitats.

C

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

495

................. 16157$

CH35

02-08-07 10:39:07

PS

PAGE 495

.

496 /

Historical Background The first collections of marine invertebrates in New Guinea appear to have been made by the naturalists Quoy and Gaimard aboard the French vessel L’Uranie in 1818 and 1819. A variety of species was collected from the Raja Ampat Islands, and these were described in the expedition results published in 1824. Later exploratory trips to the region also resulted in the incidental collection of material. The Raja Ampats were visited by other French vessels, including La Coquille (1823) and L’Astrolabe (1826). In addition to France, Papua was visited by vessels from Great Britain, Holland, Germany, Australia, and the United States. Papers on individual phyla were presented by a variety of authors in the expeditionary reports. The first major publication on mollusks in the region appears to have been Oostingh (1925), who reported in detail on the marine mollusks of the nearby Obi Major and Halmahera Islands, recording 298 species. In addition, he provided a summary of collecting that had been done in the area until 1925.

Origin and Composition One of the key questions about the Coral Triangle is: Why is the area so rich in species? While there is no single definitive answer, there is a wealth of evidence to show how this came about (see also Chapter 5.2). Shallow waters in the tropics have long been considered to be reasonably stable habitats that have existed for millions of years. While they may have contracted and expanded over time with phenomena such as the ice ages, the tropics have always existed. The stable tropical environment is thought to have allowed species to occupy smaller niches than they do in environments subject to greater fluctuations. However, there has been an increasing awareness in recent decades that the tropics is not as benign as it seems to a casual observer. Cyclones and typhoons can be extremely destructive over short periods of time. The Indian Ocean is subject to the annual monsoonal deluges that can severely affect coastal waters. Even the rainy season in areas such a Papua can result in considerable deposition of silt into the coastal marine environment. During the Pleistocene there were a number of periods of alternating high and low sea levels. During periods of low sea levels, the Indian and Pacific Oceans tended to be separate basins, with speciation occurring independently in each. As sea levels rose, the Indonesian archipelago became an area where the two biotas overlapped. The high diversity of corals in Indonesia is explained by the overlapping ranges of many widespread species; only 31 species are endemic to the region (Veron 2000a). Veron (2000b) places Papua in a biogeographical grouping to the east of Wallace’s Line that includes all of the island of New Guinea and the Australian Great Barrier Reef. Wallace et al. (2002) consider the Togian Islands in the Bay of Tomini in central Sulawesi to be the ultimate center of diversity in the Indo-Pacific. The

................. 16157$

CH35

02-08-07 10:39:08

PS

PAGE 496

Marine Invertebrates of Papua / 497

area is to the west of Wallace’s Line. In fact, Wallace et al. (2002) surveyed several faunal groups. Bivalves, gonodactyloid stomatopods, and terebellid polychaetes were all very diverse in the Togian Islands, but labrid and scarid fishes, fungiid corals and their associates, all had lower diversity than has been found in other parts of the coral triangle. In fact, for acroporid corals, the greatest similarly with the Togian Islands is with Kimbe Bay in northern PNG. They also emphasize the importance of habitat complexity in maintaining the considerable species diversity in the region. Wallace et al. (2002) propose that the Togian Islands area is a region which operated as a lagoonal refuge during Pleistocene low water levels. With the subsequent sea level rise, the refuges received additional species from the Pacific, carried into the area by the throughflow which moves water from the Pacific Ocean into the eastern Indian Ocean as part of the El Nin˜o Southern Oscillation phenomenon.

Major Invertebrate Groups There are literally dozens of invertebrate phyla that have benthic species living in the Coral Triangle. While they are often lumped together for convenience, invertebrates are not a distinct group. At the phylum level they range from very simple animals such as jellyfish to exceedingly complex organisms such as squid and octopus. The only unifying feature among invertebrates is that they lack the backbone of the vertebrates (birds, fish, reptiles, amphibians, and mammals). All of the vertebrates are classified together in the phylum Chordata, but the primitive chordates are invertebrates as they do not have a backbone. Many of these groups are extremely poorly known and have never been properly surveyed anywhere in the world, much less in an isolated area such as Papua. As an example of how poorly known marine invertebrates are, the results have been published on six marine biological workshops which have been held in the western half of Australia in the last two decades. Each workshop had approximately 17 days of fieldwork (Hanley et al. 1997; Walker and Wells 1999; Wells 1997; Wells et al. 1990, 1993, 2003, 2005), a relatively short period of fieldwork in which to document a taxonomic group in the area. The workshop proceedings and related papers have described a total of 2 new families, 23 genera, and 330 new species across a wide range of invertebrate groups, including worms (oligochaetes, polychaetes, nemerteans, and flatworms), marine mites, mollusks (bivalves, gastropods, and an aplacophoran), crustaceans (barnacles, hermit crabs, and a mystococarid), cnidarians, and sponges. Dr Christer Erse´us of the Swedish Museum of Natural History documented the presence for the first time in Australia of 90 species of marine oligochaetes, 64 of which were new to science. Dr Ilse Bartsch of the Senckenberg Institute in Germany described 82 new species of marine mites. If studies conducted over a limited time frame in Australia can uncover such an incredible number of new species, one can only imagine the undiscovered biodiversity in Papua! Because of the large number (over 50) of invertebrate phyla, and

................. 16157$

CH35

02-08-07 10:39:08

PS

PAGE 497

.

498 /

the poor state of our knowledge of most of the phyla, the present report concentrates on mollusks and corals. Corals are diverse, and provide the foundations of the reefs on which they live. Because of their symbiosis with unicellular zooxanthellae, they are also important as primary producers. Mollusks are by far the most diverse of the marine invertebrate phyla. In terms of diversity, mollusks are the insects of the sea. Gosliner et al. (1996) estimated that mollusks comprise 60% of invertebrate diversity in the marine environment. Many species of mollusks are ecologically important, with some species also being commercial fishery species. Knowledge of mollusks on reefs in the coral triangle has been considerably increased as they are the two invertebrate groups being examined by the Conservation International Marine Rapid Assessment Program (RAP).

Compared to the diversity of groups such as mollusks, crustaceans, and echinoderms, corals are not particularly diverse. However, they are the key building block of coral reefs. The calcified exoskeleton provides a solid attachment base for other organisms and a three-dimensional habitat for other species to colonize. Veron (2000a) has published broad distribution patterns for 793 coral species worldwide. These species are distributed over 18 families and 111 genera. Veron (2002) reported that 581 corals are known from Indonesia alone, meaning that the island nation has a majority (73%) of the world’s total number of species. He also recorded 465 species on the Conservation International Marine RAP of the Raja Ampat Islands, the highest number recorded on any of the five surveys in the Coral Triangle (Table 4.2.3). Wallace (2002) examined distribution patterns in the speciose genus Acropora, which has 113 species. The greatest diversity in the genus is in Indonesia (91 species) and Papua New Guinea (75 species).

The total number of shallow water mollusk species in the Indo-West Pacific Ocean, or any portion of it, is not known. Briggs (1995) estimated that there are 6,000 species of shallow water mollusks, but this is certainly an underestimate. Gosliner (2002) reported that there are 3,400 species of opisthobranchs, a relatively small component of total molluscan diversity. Diversity of opisthobranchs is greatest in the Coral Triangle, with 563 species recorded from the Philippines and 646 from Papua New Guinea. Data are not available for Indonesia, including Papua, but unpublished surveys by Gosliner and others suggest opisthobranch diversity may be higher in Indonesia. Using the proportions of cowries and volutes collected in the survey discussed below, Bouchet et al. (2002) estimated there are between 11,904 shallow water species of mollusks (based on the snail family of cowries) and an incredible 99,963 (based on another snail family, the volutes) in the Indo-Pacific. The most intensive survey of mollusk diversity has been undertaken by Bouchet et al. (2002), who examined species present in a 295 km2 site on the west coast of New Caledonia. The site is essentially a coral reef system with an outer barrier reef

................. 16157$

CH35

02-08-07 10:39:09

PS

PAGE 498

Marine Invertebrates of Papua / 499

STOMATOPOD CRUSTACEANS OF NORTHERN PAPUA (VOGELKOP SEASCAPE) Stomatopods, also known as mantis shrimp, are benthic marine crustaceans that occur predominantly in coral reef and seagrass habitats in the tropical oceans of the world, though some are found in temperate waters and a few species are deepsea specialists that occur in depths of up to 1,500 meters. Stomatopods are a diverse group, with at least 450 species representing over 100 genera in 17 families. Though related to the more commonly-known decapods such as crabs, shrimp, and lobsters, stomatopods are quite different from these groups in having two enlarged raptorial appendages (much like praying mantis insects) instead of pincers as their primary defensive and predatory appendages. The morphology of these raptorial appendages naturally divides the stomatopods into two functional groups: the spearers and the smashers. The raptorial appendages of the spearers are lined with long, sharp teeth that are used to impale soft-bodied prey such as fish and prawns. Spearers typically live in burrows they excavate in soft-bottom habitats, where they frequently occur in monogamous pairs. The largest spearers reach up to 40 cm in length and can live for 20 years or more. Smashers are generally much smaller, with the largest reaching 17 cm, but most species are less than 6 cm. They have a hardened, calcified heel on their raptorial appendages that they use to smash apart hard-bodied prey such as crabs, snails, and other gastropods. This habit has earned them the nickname ‘‘thumbnail splitters’’ and ‘‘devil shrimp’’ in some languages, and they have even been known to smash aquarium glass with their strike, which is slightly faster than a .22 caliber bullet. The smashers live mostly on coral reefs, where they inhabit cavities in hard substrates including coral rubble and live coral. Unlike the monogamous spearers, smashers display generally promiscuous reproductive behavior. The author has been sampling stomatopod crustaceans from around Indonesia and the Coral Triangle region for the past 14 years, focusing predominantly on the superfamily Gonodactyloidea, which includes all of the smashers. Of all the areas studied, the Vogelkop region of northern Papua, particularly the Raja Ampat Archipelago, is the most diverse by far. In 15 days of sampling from 36 sites (32 in Raja Ampat, 4 in the Biak/Padaido Islands), I collected a total of 37 gonodactyloid species (see Table 4.2.1), including five previously undescribed species, one

Table 4.2.1. Gonodactyloid stomatopods collected from Vogelkop region Family

Species

Gonodactylidae Geisbrecht 1883

Gonodactylaceus falcatus (Forska˚l 1775) Gonodactylaceus glabrous (Brooks 1886) Gonodactylellus affinis (de Man 1902) Gonodactylellus annularis Erdmann and Manning 1998

................. 16157$

CH35

02-08-07 10:39:10

PS

PAGE 499

.

500 /

Gonodactylellus erdmanni Ahyong 2001 Gonodactylellus espinosus (Borradaile 1898) Gonodactylellus kandi Ahyong and Erdmann 2006 Gonodactylellus micronesicus (Manning 1971) Gonodactylellus rubriguttatus Erdmann and Manning 1998 Gonodactylellus snidsvongi (Naiyanetr 1987) Gonodactylellus viridis (Sere`ne 1954) Gonodactylellus sp. A (Ahyong and Erdmann, in prep) Gonodactylellus sp. B (Erdmann, in prep) Gonodactylopsis sp. A (Erdmann, in prep) Gonodactylus childi Manning 1971 Gonodactylus chiragra (Fabricius 1781) Gonodactylus platysoma (Wood-Mason 1895) Gonodactylus smithii Pocock 1893 Hoplosquilla said Erdmann and Manning 1998 Odontodactylidae Manning 1980

Odontodactylus cultrifer (White 1851) Odontodactylus latirostris Borradaile 1907 Odontodactylus scyllarus (Linnaeus 1758) Odontodactylus sp. A (Erdman, in prep)

Protosquillidae Manning 1980

Chorisquilla brooksii (de Man 1888) Chorisquilla gyrosa (Odhner 1923) Chorisquilla hystrix (Nobili 1899) Chorisquilla mehtae Erdmann and Manning 1998 Chorisquilla pococki (Manning 1975) Chorisquilla spinosissima (Pfeffer 1888) Chorisquilla sp. A (Erdmann, in prep) Echinosquilla guerini (White 1861) Haptosquilla glyptocercus (Wood-Mason 1875) Haptosquilla pulchella (Meiers 1880) Haptosquilla pulchra (Hansen 1926) Haptosquilla trispinosa (Dana 1852) Haptosquilla tuberosa (Pocock 1893) Haptosquilla sp. A (Erdmann, in prep) Haptosquilla sp. B (Erdmann in prep) Haptosquilla sp. C (Erdmann, in prep) Siamosquilla laevicaudata (Sun and Yang 1998) Siamosquilla sp. A (Erdmann, in prep)

Pseudosquillidae Manning 1977

Pseudosquilla ciliata (Fabricius 1787) Pseudosquilla megalophthalma (Bigelow 1893) Raoulserenea ornata (Miers 1880) Raoulserenea oxyrhyncha (Borradaile 1898)

Takuidae Manning 1995

Taku spinosocarinatus (Fukuda 1909)

................. 16157$

CH35

02-08-07 10:39:10

PS

PAGE 500

Marine Invertebrates of Papua / 501 of which appears endemic to the Raja Ampat Archipelago. The 37 species collected represent 14 of the 24 genera and 5 of the 7 families of gonodactyloids known worldwide. This is the highest known species diversity of reef-associated stomatopods for a seascape of this size anywhere in the world. By comparison, Australia’s Queensland coast has 26 known gonodactyloids (Ahyong 2001), Vietnam has 24 (Manning 1995), and the entire Caribbean area has 17 known gonodactyloids (Manning 1969, 1970; Schotte and Manning 1993). Within the Coral Triangle, the next most diverse seascapes are northern Sulawesi (32 gonodactyloids) and the Wakatobi area of southeast Sulawesi (31 gonodactyloids), both of which have been examined with more than triple the sampling effort of the Vogelkop. Table 4.2.2 shows the known zoogeographic distributions of the stomatopod species collected from northern Papua (following the classification of Allen, 2002). While roughly 65% of the collected species have a relatively wide distribution (i.e., found at least as far as the central Pacific or Indian Ocean), another 35% show a much more restricted range (e.g., found only in the Western Pacific or within the Indo-Australian Archipelago). Fully 13% of these species are found only in eastern Indonesia, including one species apparently endemic to Raja Ampat’s reefs.

Table 4.2.2. Zoogeographic distribution of stomatopod species collected from the Vogelkop region Zoogeographic distribution

Number of species

Percentage of world total

Circumtropical

1

2.10%

Indo-west and central Pacific

7

14.90%

Indo-west Pacific

7

14.90%

East Indian and west and central Pacific

4

8.50%

East Indian and west Pacific

10

21.30%

West and central Pacific

1

2.10%

West Pacific (incl. South China Sea)

3

6.40%

Indo-Australian Archipelago

2

4.30%

Indo-Philippines

2

4.30%

Eastern Indonesia

4

8.50%

Endemic to Vogelkop

6

12.70%

The fantastic diversity of stomatopods in the Vogelkop seascape is likely a direct result of the equally impressive diversity of reef habitats represented in the area, with a sharp gradient of exposure, sedimentation, and current regimes. Among the varied habitats sampled are protected bay fringing reefs, fringing reefs subject to freshwater influx, steep walls and platform reefs with strong currents, clear water

................. 16157$

CH35

02-08-07 10:39:11

PS

PAGE 501

.

502 /

slopes and lagoons, rocky reefs, and wave-pounded reef flats. The sheer diversity of stomatopod species alone would argue for the prioritization of this area for conservation efforts, with the strong representation of restricted range species further emphasizing the importance of protecting this unique area. Ongoing population genetic studies of stomatopods from the region should help elucidate patterns of genetic connectivity of the Vogelkop seascape with other reefs in the Coral Triangle and thereby give a clear indication of the importance of including these diverse reefs in Indonesia’s developing marine protected area (MPA) network. Mark V. Erdmann

Acknowledgments The author wishes to thank the Indonesian Institute of Sciences, and especially Dr. M. Kasim Moosa, for support and guidance in stomatopod research over the past decade. Literature Cited Ahyong, S.T. 2001. Revision of the Australian stomatopod Crustacea. Records of the Australian Museum, Supplement 26: 1–326. Allen, G.R. 2002. Reef fishes of the Raja Ampat Islands, Papua Province, Indonesia. Pp. 46–57 in McKenna, S.A., G.R. Allen, and S. Suryadi (eds.) A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. Conservation International, Washington, D.C. Manning, R.B. 1969. Stomatopod Crustacea of the Western Atlantic. Studies in Tropical Oceanography, Miami 8: 1–380. Manning, R.B. 1970. Nine new American stomatopod crustaceans. Proceedings of Biological Society of Washington 83: 99–114. Manning, R.B. 1995. Stomatopod Crustacea of Vietnam: the legacy of Raoul Sere`ne. Crustacean Research, Special No. 4: 1–339. Schotte, S.R., and R.B. Manning. 1993. Stomatopod Crustacea from Tobago, West Indies. Proceedings of Biological Society of Washington 106: 566–581.

9–12 km from shore, a lagoon, and a shoreline of sedimentary bays, mangroves, and rocky shores. Four hundred person-days were spent in the field collecting and sorting specimens from the intertidal to depths of 120 m. A total of 2,738 species of mollusks was collected, several times the maximum diversity published by previous expeditions to other areas. Nearly a third of all species collected were found at a single station, including the 20% of species represented by a single individual. Even with the 2,738 species found, Bouchet et al. (2002) concluded that softbodied species were underrepresented. Because of the specialized sampling required, cephalopods were not sampled at all. Bouchet et al. (2002) predicted the total number of mollusk species at the site could be as high as 3,971 species. Wells (2002c) divided the tropical Indo-West Pacific into ten regional areas and surveyed the distribution of 1,268 species of mollusks in each area. The greatest diversity was in the Coral Triangle, in which 745 of the species occurred. Forty-

................. 16157$

CH35

02-08-07 10:39:12

PS

PAGE 502

19

Milne Bay, Papua New Guinea, Expedition 1

................. 16157$

CH35

12

02-08-07 10:39:13

465

15

Note: Collected in five Marine RAP surveys in the Coral Triangle conducted by Conservation International.

15

11

Raja Ampat Islands, Indonesia

314

17

Togian and Banggai Islands, Indonesia

19

16

16

11

19

Calamianes Islands, Philippines

305

362

Collecting days

30 19

15

Number of species

Total for Milne Bay

Milne Bay, Papua New Guinea, Expedition 2

Collecting days

Location

Corals Number of families

Table 4.2.3. Biodiversity of corals and mollusks

94

103

96

110

109

Mollusks Number of families

699

541

651

945

643

637

Number of species

Veron 2002; Wells 2002b

Fenner 2002; Wells 2002a

Veron 2000; Wells 2000

Wells in press

Veron in press; Wells in press

Veron 1998; Wells 1998

Reference Marine Invertebrates of Papua / 503

PS

PAGE 503

.

504 /

nine of these species were endemic to the Coral Triangle, but distributional information is not sufficient to know if any of the 49 is restricted to Papua. Wells (2002b) recorded 699 species of mollusks on the Raja Ampats expedition. As with corals, the Raja Ampats had the greatest number of species of mollusks for any of the five Marine RAPs in the coral triangle (Table 4.2.3). The distributions of 258 species of mollusks collected on the Raja Ampats expedition for which there is good distributional information have been examined. Two hundred and three species (79% of the total) are widespread Indo-West Pacific species and 54 species have ranges which are restricted to the western Pacific, central and western Pacific, or western Pacific and eastern Indian Ocean (Table 4.2.4). Only a single species, Terebra caddeyi, is currently thought to be restricted to the north coast of New Guinea. This is a relatively recently described species, and further investigations are likely to show its distribution to be more widespread than is presently known. There are few marine laboratories on the coast of New Guinea. During the 1980s the Christensen Research Institute was developed in Madang, on the north coast of PNG. An incredible diversity of species has been recorded from the Madang area. Kohn (2001) considers the north coast of Papua New Guinea to be a Biodiversity Hotspot, with many invertebrate groups reaching their highest known species diversity, including 536 species of opisthobranch mollusks in more than 50 families at Madang (Ghiselin 1993; Gosliner 1993). With over 500 extant species, the gastropod genus Conus is the most diverse genus in the marine invertebrates. Kohn (2001) found 63 species on a single platform fringing Laing Island and 32 species on four small reefs near Madang. Given the absence of similar studies on the north coast of Papua, it would not be surprising to find a similarly high diversity there.

Invertebrate Habitats Invertebrates are the most diverse groups on the coral reefs of Papua. Veron (2002) estimated that 565 species of coral have been recorded or are likely to occur

Table 4.2.4. Geographical distribution of selected species of mollusks Geographic area

Number of species

Indo-west Pacific

Percentage of total species

203

79

Western Pacific

34

13

Central and western Pacific

12

5

Western Pacific and eastern Indian Ocean

8

3

Endemic to Papua New Guinea and the Coral Sea

1

0

258

100

Total

Note: Collected during the Raja Ampat Islands Survey (Wells 2002b).

................. 16157$

CH35

02-08-07 10:39:13

PS

PAGE 504

Marine Invertebrates of Papua / 505

on the reefs of the region. A total of 465 species were recorded on a single survey of the Raja Ampat Islands. Not only do the corals themselves provide a significant diversity, but they considerably increase habitat structure in the environment. Living corals provide habitats for other invertebrates to live in, on, below, or just above the live coral. Additional niches are found in the skeletons of dead coral and among coral rubble. The survey by Conservation International provided considerable detail on the distribution of corals in the Raja Ampat Islands in the western portion of Papua. It is the only area to have been surveyed in detail, but further expeditions to the Raja Ampats would undoubtedly provide additional records in the groups surveyed. Allen (Chapter 4.8) reports that there are other significant reefs in Papua at Cenderawasih Bay and around Biak on the north coast, near Jayapura, and along the southwestern coast off the Fakfak Peninsula and between Kaimana and Etna bays. Coral reefs come in a huge variety of shapes and sizes, but can be divided into the two categories of offshore and inshore. Offshore reefs, such as atolls, are in environments where there is little or no terrestrial runoff. The water is clear, and there is often open sea wave action. Turbidity is often much higher on the coastal inshore reefs as a result of runoff from adjacent terrestrial areas. Inshore reefs may be located in protected bays where wave action is slight. There can be considerable differences in the fauna inhabiting reefs in the two areas. In the early 1980s, the Western Australian Museum examined a series of offshore atolls, and found that across a range of faunal groups, 20–25% of the species did not occur on inshore continental reefs (Berry 1986, 1983). Coral studies undertaken during the Raja Ampat expedition resulted in the finding of nine potentially new species (Veron 2002) in the silty inshore waters. In the Raja Ampat survey, Wells (2002b) recorded 699 species of mollusks in 15 collecting days, the largest number on any of the Marine RAP surveys conducted by Conservation International. Because of time limitations, these surveys concentrated on macromollusks 1 cm or larger. The work of Bouchet et al. (2002) demonstrated that the actual number of species present in areas such as the Raja Ampats would be several times greater if a detailed survey was undertaken. However, such a survey, with 400 days in the field and considerably increased time in the laboratory cannot be done in a limited time frame. Forty-four species of cowries (Cypraea) were collected during the expedition. Hugh Morrison, a very experienced shell collector, later visited the same area for seven days. He raised the number of cowries known from the area to 63. Wells (2002b) found that most species tended to be found at one or a few stations, often only as dead shells. In contrast, a small number of species was found at 30 or more of the 44 stations investigated. These included the ark shell Arca avellana, the scallops Gloripallium radula and Pedum spondyloidaeum, the giant clams Tridacna squamosa and T. crocea, the venerids Antigona restriculata and Venus toreuma, the boring mussels Lithophaga spp., and the coral dwelling gastropod Coralliophila neritoidea. Many of these records were based on dead shells. The numerically dominant living species were burrowing ark shells, Lithophaga spp., P. spondyloidaeum, and C. neritoidea.

................. 16157$

CH35

02-08-07 10:39:14

PS

PAGE 505

.

506 /

Estuaries are the transitional zones between freshwater rivers flowing off the land and the marine waters of the open sea. There tends to be a gradation in salinity ranging from the very low salinity of freshwater (often near 0 parts per thousand) in the upper part of the estuary to open seawater salinities of 36 ppt near the mouth. Species diversity in the estuary varies with the salinity. In the upper areas freshwater species capable of tolerating a salinity of a few parts per thousand encroach into the estuary. There are relatively few invertebrates capable of surviving entirely in the estuary itself, where salinities range from a few to perhaps 15 ppt. Diversity increases in the lower estuary as one approaches the mouth. As salinities become higher, progressively more essentially marine species are able to tolerate the salinity levels. These animals may be present in the estuary throughout the year or may move into it seasonally when conditions are right for their survival. In tropical areas, most estuaries are fringed with mangroves, one of the underappreciated habitats of tropical seas, both among scientists and the general public. The mangroves of Papua, particularly on the south coast, are among the largest in the world (Chapter 5.4). In western countries mangroves are considered to be fetid swamps which breed mosquitoes, midges, and other unpleasant insects. As such they are thought to ‘‘stand in the way of progress’’ and are often cleared to provide space for the development of harbors or housing estates. In developing countries, such as Papua, mangroves are cut down for their wood or to provide space for shrimp farming. The continuing destruction of mangroves is an ecological disaster, as mangroves are one of the key habitats in coastal ecosystems in the tropics. In contrast to their poor popular image, mangroves are essential systems which provide considerable primary and secondary production, and are important nursery areas for commercially valuable species of fish and shellfish, including shrimp (Robertson 1988). A mangrove species is simply a plant that is adapted to live in a coastal environment where it is in periodic contact with seawater (see Chapter 5.4 for further details). The term mangrove (which can be used to describe either the habitat type or the plants that live therein) has no taxonomic meaning: there are approximately 80 species of plants which are considered to be mangroves, but they belong to approximately 30 genera spread over 20 plant families. Mangroves are both full grown trees and smaller shrubs. Mangroves occur in the middle to upper intertidal areas of the shoreline, where at least the lower surfaces of the plants are typically covered during high tide on a daily basis. Some of the mangroves on the upper part of the shoreline may be reached by the sea only during spring tides, which occur every two weeks, or possibly on a monthly basis. The fauna that inhabits mangroves is complex, and can be divided into a number of different categories. Perhaps the most useful is to consider species that are of marine, freshwater, and terrestrial origins. Freshwater species are limited to the landward margins of mangroves, and are typically only a minor component of the fauna. During monsoons in the Indo-Pacific regions or during flooding, additional freshwater species can be washed into the mangroves.

................. 16157$

CH35

02-08-07 10:39:14

PS

PAGE 506

Marine Invertebrates of Papua / 507

Marine animals living in the mangroves can be readily divided into two groups: permanent residents and transients. Permanent inhabitants may have planktonic larvae that settle into the mangroves at metamorphosis, but once the benthic stage is reached, the animals live in the mangroves throughout their juvenile and adult stages. Oysters and shipworms are examples of species that remain at one site once they settle. There is an unending tidal movement of transient species. As the tide rises, a wide variety of animals enters the mangroves to feed, reproduce, and undertake their other biological functions. Among these are a wide variety of fish, crustaceans, sea snakes, turtles, and the estuarine crocodile, Crocodylus porosus. These animals leave the mangroves when the tide falls and are replaced by immigrants from the land, which include birds, mammals, reptiles, and insects. Animals that are permanent residents of mangroves can be further divided into species that characteristically occur only in mangroves and those which are common on adjacent rocky, sandy, or muddy shores and use the mangrove as only part of the habitat they occupy. The number of species characteristic of mangroves is relatively small; most species also occur in adjacent habitats. For example, several hundred species of mollusks have been recorded in mangroves in the IndoWest Pacific, but fewer than 50 of these are characteristic of mangroves. Many are uncommon or cryptic, but others such as three species of the mudwhelk genus Terebralia are conspicuous inhabitants of mangroves. Four major benthic marine habitats can be found in the mangroves: infaunal, epifaunal, epibiotic, and species that live within the trees. Infaunal species, those that live within the substrate, are rare in the mangroves. Working in the Kimberley region of Western Australia, Wells and Slack-Smith (1981) examined the distribution of mollusks in six zones of the mangroves. Not a single living mollusk was found in any of the infaunal samples from the tree zones. This work was later extended to cover all invertebrate groups in a study undertaken at North West Cape in Western Australia (Wells 1983, 1984). Again, infaunal species were only a minor component of the fauna in the tree zone, but were abundant in the mudflat seaward of the trees. Very few infaunal mollusks are found in the mangroves. An exception is the bivalve Geloina, which is abundant on the surface of the mangrove floor of many Papuan mangroves. It can form an important part of the diet of local people. Burrowing crabs are uncommon among the trees, but are often abundant along the margins of tidal creeks. Fiddler crabs (Uca) and sesarmids are readily observed in this habitat, and the mangrove crab Scylla is also common. Epifaunal species living on the mud surface are probably the dominant faunal group in mangroves. The diversity, density, and biomass of invertebrates, including epifaunal species, varies considerably in different tree zones. Wells (1983, 1984) found these characters to be highest on the mudflat seaward of the mangroves, lower in monospecific areas of Avicennia, and lower still in Rhizophora. The decrease in abundance of mollusks in going from the seaward mudflat into Avicennia occurred at the extreme limit of the pneumatophores, not the trunks of the trees (Wells 1986). Despite the generally low abundance of epifaunal species in the tree zones, some species can be spectacularly abundant. In particular, mudwhelks of

................. 16157$

CH35

02-08-07 10:39:15

PS

PAGE 507

.

508 /

the genus Terebralia paulstris can reach a shell length of 12 cm and a density of over 100 per m2 (Wells 1980). It is also often eaten by local people. A number of mollusk groups are abundant in mangroves. The snail genus Littoraria has speciated into a variety of mangrove habitats, and some species are abundant among the foliage of the upper portions of the trees. Reid (1986) examined the species critically and recognized 20 species in the Indo-West Pacific, whereas previous studies had concluded there was only a single species. Despite their name, shipworms are bivalve mollusks of the family Teredinidae. There are approximately 66 species worldwide, many of which occur in mangroves. Turner (1966) reported 17 species from the coral triangle region, including Papua. While little is known of the biology of the group, some species occur in living mangroves and others colonize dead trees or logs. Their presence can be determined by the small pair of openings which allow the siphons of the animals to contact surrounding seawater (Turner 1966). These bivalves are frequently eaten by people in Papua. Potamidid snails, which include the genera Terebralia, Telescopium, and Cerithidea, are often abundant in mangroves. The other diverse group of mangrove gastropods is the air breathing family Ellobiidae, which live in dead logs, under leaves, or on the mud surface. A single species of Nerita is also abundant on the mangrove trees. Crustaceans are a key group in mangroves, and are important in converting the plant biomass of fallen leaves into animal biomass available to higher trophic levels (Lee 1998). Important crustaceans include the crab genera Sesarma and Macropthalmus, the mud lobster Thalassina anomala, and fiddler crabs (Uca). There is a single species of sipunculan, Phascolosoma arcuatum, which is often abundant in the sediments of mangroves. Zonation patterns of invertebrates in the mangroves can be related to those of the trees. Wells and Slack-Smith (1981) divided the mangroves at Port Warrender in the Kimberley region of Australia into five tree zones. Zonation patterns of mollusks closely matched those of the trees, with individual species having very different zonation patterns. As indicated above, many terrestrial animals enter the mangroves at low tide to forage on the forest floor or on the lower trunks of the trees. The lower parts of the trees, or even entire trees near the seaward margins of the mangroves, are submerged during high tides. However, much of the upper foliage of the mangrove is well above the upper limits of spring tides and is exposed on even the highest tides. Terrestrial species of insects, spiders, birds, possums, snakes, and other animals can colonize this portion of the mangroves. As with the marine fauna lower down on the trees, the terrestrial species are a combination of species which occur in adjacent habitats on land and those which are restricted to mangroves.

Biology of Marine Invertebrates It is impossible to summarize in a few paragraphs, or even pages, the biology of the many thousands of marine invertebrates that are found in a tropical area the

................. 16157$

CH35

02-08-07 10:39:15

PS

PAGE 508

Marine Invertebrates of Papua / 509

size of Papua. Among benthic shallow water mollusks, for example, there is a full range of species which varies from minute forms which reach only one or a few millimeters in length to the largest of the giant clams, Tridacna gigas, which can be up to 137 cm long and weigh 250 kg. These species have a variety of life styles. Almost all bivalves are filter feeders, trapping minute particles present in the water flowing over their gills. Less common are deposit feeders that obtain material from the sediment surface. Rarer still are carnivorous species or those that depend on sulfur-oxidizing bacteria. Scaphopods are surface deposit feeders, and cephalopods are well known as carnivorous predators. Chitons tend to be surface raspers, meaning they scrape tiny bits of algae from rock surfaces with their teeth. There is a wealth of feeding mechanisms among gastropods. Many are surface raspers; others are herbivorous, carnivorous, omnivorous, filter feeders, or even parasitic. Reproductive strategies are similarly diverse. Many species of mollusks have external fertilization after which the young develop into a planktonic veliger stage (a larval type found only in mollusks), which may be preceded by a trochophore larval stage (another larval type which also occurs in some other phyla). Other species have internal fertilization but retain the planktonic veliger stage after the egg has been fertilized. The veliger stage lasts only a few days in many groups. Species of Conus are competent to settle after periods ranging from 8 to 30 days (Kohn 2001). Species of other groups, such as ranellids, may be have planktonic lifespans of over a year, allowing individual larvae to travel widely across open ocean systems. Benthic egg capsules are found in many gastropods from which the young hatch as crawling juveniles. In other species developing embryos are retained within the body or shell of the female and are released as crawling juveniles. Despite the incredible variety of life styles among marine invertebrates, two biological features are worth mentioning here: symbiotic zooxanthellae and planktonic larval stages. Despite their sessile habitat as adults, corals are animals. They are cnidarians, the group to which sea anemones and jellyfish belong. Corals have stinging cells, or pneumatocysts, in their tentacles that they use to trap zooplankton as it is washed over the reef by tides and currents. Thus they are carnivorous. However, in addition to receiving nutrition as carnivores, corals have within their tissues millions of single celled algae called zooxanthellae, with which they have a symbiotic relationship (see Chapter 5.2 for additional information). Corals provide the zooxanthellae with protection, nutrients, and carbon dioxide. In return the corals are nourished by the zooxanthellae and also receive oxygen. This relationship is one of the important mechanisms that coral reefs employ to develop high population densities and biomass of tissue in open ocean waters that are oligotrophic, or nutrient poor. Zooxanthellae are best known and most ecologically important in corals, but also occur in other groups of animals, such as giant clams and jellyfish. The second key attribute of marine invertebrates is that across the entire range of marine invertebrate phyla, the great majority of species have planktonic larvae. We have already mentioned that in some snails the larval stage lasts long enough for the larvae to cross oceans. While some species remain in the plankton for over

................. 16157$

CH35

02-08-07 10:39:16

PS

PAGE 509

.

510 /

a year, feeding and growing, most remain in the water column for only a few days and do not feed. However, even a few days is enough for the larvae to move from one reef to another over a wide range. This allows the various populations of a species to remain in genetic contact over a wide geographic range. Maintaining a wide distribution is much more difficult for species that do not have a planktonic larval stage. However, natural processes, such as being carried about by drifting logs, do provide some means of genetic interchange. In general, species lacking a planktonic stage have smaller distributions. Within gastropods, for example, the volutes hatch as crawling juveniles from benthic egg masses. Groups such as this tend to speciate because populations are too far apart to maintain contact with each other. Compared to those of mollusks, reproductive mechanisms in corals are relatively simple. Approximately three-fourths of the reef-building corals are hermaphroditic; the remainder have separate sexes. Three-quarters have external fertilization in which sperm and egg are fertilized in the water column, then develop into a planula larva. The remainder brood young to the planula stage. Some species use different modes of reproduction in different geographic areas. Synchronous spawning, in which all individuals of a population on a reef, and indeed most species, release their reproductive materials simultaneously is now well known. It occurs in October/November on the Great Barrier Reef, in March in Western Australia, and in July in southern Japan. Synchronous spawning enhances the chances of successful fertilization. At the same time, predators are swamped with so much food they cannot consume it all. Planulae can remain in the water column for variable periods, even months, which allows them to attain widespread distributions. They are aided in their planktonic existence by the zooxanthellae which provide nutrition to the planulae (Veron 2000a).

Conservation Unless they have a particular human focus, such as being actively fished, the best way to conserve marine invertebrate species is to conserve the habitats in which they live. For example, a coral reef is best protected by managing the habitat, including adjacent terrestrial areas, rather than trying to protect individual species. If, for example, logging denudes an adjacent catchment area, the reef will be under threat from increased terrestrial runoff, sedimentation, and a host of other mechanisms that degrade the reef system. Efforts to protect individual species will be useless if the habitat is destroyed. Once the general area is protected, mechanisms can be developed that are aimed directly at protecting the reef. This can include the establishment of marine parks, which can be zoned to allow particular activities to occur in areas where they will not degrade the reef. While such parks can be established in areas such as Papua, there are often not sufficient political will and resources to manage the parks effectively. The general consensus has been that most invertebrate species have very widespread distributions. The money cowry (Cypraea moneta), for example, extends

................. 16157$

CH35

02-08-07 10:39:17

PS

PAGE 510

Marine Invertebrates of Papua / 511

from the east coast of Africa through the entire tropical expanse of the Indian and Pacific Oceans, including Papua, to Hawaii. Along with a few other species, it has been recorded along the west coast of the Americas. There would be little risk of species with such vast distributions becoming extinct through human activities. This is in stark contrast to freshwater and terrestrial species which may be restricted to one drainage system or an isolated hilltop. There have been many documented extinctions of both freshwater and terrestrial invertebrates, but documented extinctions of invertebrates are rare in the marine environment. While human activities in local areas are unlikely to result in extinctions, there has been a recent accumulation of evidence that suggests that there are significant levels of endemism in marine invertebrates. Even in widespread species, there is now an understanding that there may be local populations with significant levels of genetic differentiation. In many cases the differentiation has progressed far enough to create closely related, but separate, species. In others differentiation has not progressed that far, but there are still recognizably different populations which should be conserved. Many scientists now consider retention of the pool of genetic diversity, not simply the retention of species, to warrant significant conservation activity. In a recent paper, Roberts et al. (2002) analyzed the distributions of 3,235 species of coral reef fish, mollusks, corals, and rock lobsters. Between 7.2% and 53.6% of each group had restricted ranges which made them more susceptible to extinction through global warming and other effects of human activities which are acting on a global scale. The definition of restricted ranges was a species that occurred in ten or fewer cells with a side of 2 of latitude or longitude. If a species occurred throughout ten cells, it would still inhabit a considerable geographic range. However, coral reef species do not occur uniformly throughout the cells. The effective range of a species is much smaller because it occurs only on the limited coral reef habitats within the larger area. Eighteen Hotspots of coral diversity and restricted range species were documented worldwide. Despite the high diversity of these groups in Papua, the province was not included on the list of Hotspots because of the low level of endemic species in the area. Ponder (2003) extended consideration of endemism in the ocean to smaller scales. Many species lack a planktonic larval stage, thus potentially severely restricting their distributional capability. Others with a short planktonic stage may have mechanisms such as negative phototaxis which are useful in retaining larvae near where they were spawned. Such a mechanism has positive survival value in areas such as a seamount where the wide distribution of larvae would severely decrease survival. However, it leads to the development of small ranges that a species may not be able to expand. If a species lives in a restricted habitat, such as an isolated rock outcrop in the middle of a sandy beach, there is little opportunity for it to be distributed beyond the beach. Ponder (2003) points out the small number of marine species that are known to have recently become extinct, but balances this against our lack of knowledge and the difficulty of convincingly documenting extinction in the marine environment.

................. 16157$

CH35

02-08-07 10:39:17

PS

PAGE 511

.

512 /

Many invertebrates are widely collected for food or other human uses. This includes many mollusks such as squid, octopus, abalone, giant clams, conches, and bivalves. Crustaceans are widely eaten, including lobsters, crabs, and shrimp. Beˆche-de-mer is a delicacy in the Asian market. Other animals, such as mollusks and corals are collected for their shells or use in aquaria. Special conservation measures are required for these groups which are directly targeted by people.

Literature Cited Barber, P.H., S.R. Palumbi, M.V. Erdmann, and M. Kasim Moosa. 2000. A marine Wallace’s line? Nature 406: 692–693. Berry, P.F. (ed.). 1986. Faunal Survey of the Rowley Shoals and Scott Reef, Western Australia. Records of the Western Australian Museum Supplement 25: 41–58. Berry, P.F. (ed.). 1993. Marine faunal surveys of Ashmore Reef and Cartier Island, Northwestern Australia. Records of the Western Australian Museum Supplement 44. Bouchet, P., P. Lozouet, P. Maestrati, and V. Heros. 2002. Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of mollusks at a New Caledonia site. Biological Journal of the Linnean Society 75: 421–436. Briggs, J.C. 1995. Global Biogeography. Elsevier, Amsterdam. Briggs, J.C. 1999. Coincident biogeographic patterns: Indo-West Pacific Ocean. Evolution 53: 326–335. Burgess, C.M. 1985. Cowries of the World. Gordon Verhoef, Seacomber Publications, South Africa. Fenner, D. 2002. Reef corals of the Togean and Banggai Islands, Sulawesi, Indonesia. Pp. 27–37, 64–71 in Allen, G.R., and S.A. McKenna (eds.) A Marine Rapid Assessment of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment 20. Conservation International, Washington, D.C. Ghiselin, M. 1993. How well known is the opisthobranch gastropod fauna of Madang, Papua New Guinea? Proceedings of the Seventh International Coral Reef Symposium 2: 697–701. Gosliner, T.M. 1993. Biodiversity of tropical opisthobranch gastropod faunas. Proceedings of the Seventh International Coral Reef Symposium 2: 702–709. Gosliner, T.M. 2002. Biodiversity, endemism and evolution of opisthobranch gastropods on Indo-Pacific coral reefs. Pp. 937–940 in Kasim Moosa, M., S. Soemodihardjo, A. Soegiarto, K. Romimohtarto, A. Nontji, Soekarno, and Suharsono (eds.) Proceedings of the 9th International Coral Reef Symposium, Bali, Indonesia, 23–27 October 2000. Ministry of Environment, Indonesian Institute of Sciences and International Society for Reef Studies, Jakarta. Gosliner, T.M., D.W. Behrens, and G.C. Williams. 1996. Coral Reef Animals of the IndoPacific. Sea Challengers, Monterey, California. Hanley, J.R., G. Caswell, D. Megerian, and H.K. Larson (eds.). 1997. The Marine Flora and Fauna of Darwin Harbour, Northern Australia. Northern Territory Museum, Darwin, and the Australian Marine Sciences Association. Kohn, A.J. 2001. Maximal species richness in Conus: diversity, diet and habitat on reefs of northeast Papua New Guinea. Coral Reefs 20: 25–38. Lee, S.Y. 1998. Ecological role of grapsid crabs in mangrove ecosystems: a review. Marine and Freshwater Research 49 (4): 335–344. Ponder, W.F. 2003. Narrow range endemism in the sea and its implications for conser-

................. 16157$

CH35

02-08-07 10:39:18

PS

PAGE 512

Marine Invertebrates of Papua / 513 vation. Pp. 89–102 in Hutchings, P.A., and D. Lunney (eds.) Conserving Marine Environments? Out of Sight Out of Mind. Royal Zoological Society of New South Wales, Mosman, New South Wales. Reaka-Kudla, M.L. 1997. An estimate of known and unknown biodiversity and potential for extinction on coral reefs. Reef Encounter 17: 8–12. Reid, D.G. 1986. The Littorinid Mollusks of Mangrove Forests in the Indo-Pacific Region the Genus Littoraria. British Museum (Natural History), London. Roberts, C.M., C.J. McClean, J.E.N Veron, J.P. Hawkins, G.R. Allen, D.E. McAllister, C.G. Mittermeier, F.W. Schueler, M. Spalding, F. Wells, C. Wynne, and T.B. Werner. 2002. Marine biodiversity hotspots and conservation priorities for tropical reefs. Science 295: 1280–1284. Robertson, A.I. 1988. Abundance, diet and predators of juvenile banana prawns, Penaeus merguiensis, in a tropical mangrove estuary. Australian Journal of Marine and Freshwater Research 38: 467. Robertson, A.I., and D.M. Alongi (eds.). 1992. Tropical Mangrove Ecosystems. Coastal and Estuarine Studies 41. American Geophysical Union, Washington, D.C. Ro¨ckel, D., W. Korn, and A.J. Kohn. 1995. Manual of the Living Conidae, Vol. 1: IndoPacific Region. Verlag Christa Hemmen, Wiesbaden. Turner, R.D. 1966. A Survey and Illustrated Catalogue of Teredinidae. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. Veron, J.E.N. 1998. Corals of the Milne Bay region of Papua New Guinea. Pp. 26–34 in Werner, T., and G.R. Allen (eds.) A Rapid Biodiversity Assessment of the Coral Reefs of Milne Bay Province, Papua New Guinea. RAP Working Papers 11. Conservation International, Washington, D.C. Veron, J.E.N. 2000a. Corals of the World. Vols. 1–3. Australian Institute of Marine Science, Townsville, Australia. Veron, J.E.N. 2000b. Corals (Zooxanthellata Scleractinia) of the Calamianes Islands, Palawan Province, Philippines. Pp. 24–26, 66–74 in Werner, T. B., and G.R. Allen (eds.) A Rapid Marine Biodiversity Assessment of the Calamianes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Conservation International, Washington, D.C. Veron, J.E.N. 2002. Reef corals of the Raja Ampat Islands, Papua Province, Indonesia. Part 1. Overview of Scleractinia. Pp. 26–28, 90–103 in McKenna, S.A., G.R. Allen, and S. Suryadi (eds.) A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, D.C. Walker, D.I., and F.E. Wells (eds.). 1999. Proceedings of the Ninth International Marine Biological Workshop: The Seagrass Flora and Fauna of Rottnest Island, Western Australia. Western Australian Museum, Perth. Wallace, C.C. 1997. The Indo-Pacific centre of coral diversity re-examined at species level. Proceedings of the Eighth International Coral Reef Symposium 1: 365–370. Wallace, C.C., G. Paulay, B.W. Hoeksema, D.R. Bellwood, P.A. Hutchings, P.H. Barber, M. Erdmann, and J. Wolstenholme. 2002. Nature and origins of unique high diversity reef faunas in the Bay of Tomini, central Sulawesi: the ultimate centre of diversity? Pp. 185–192 in Kasim Moosa, M., S. Soemodihardjo, A. Soegiarto, K. Romimohtarto, A. Nontji, Soekrano, and Suharsono (eds.) Proceedings of the 9th International Coral Reef Symposium, Bali, Indonesia, 23–27 October 2000. Ministry of Environment, Indonesian Institute of Sciences and International Society for Reef Studies, Jakarta. Wells, F.E. 1980. A comparative study of distribution of the mudwhelks Terebralia sulcata

................. 16157$

CH35

02-08-07 10:39:19

PS

PAGE 513

.

514 /

and T. palustris in a mangrove swamp in northwestern Australia. Malacological Review 13: 1–5. Wells, F.E. 1983. An analysis of marine invertebrate distributions in a mangrove swamp in northwestern Australia. Bulletin of Marine Science 33: 736–744. Wells, F.E. 1984. Comparative distribution of macromolluscs and macrocrustaceans in a North-western Australian mangrove system. Australian Journal of Marine and Freshwater Research 35: 591–596. Wells, F.E. 1986. Distribution of mollusks across a pneumatophore boundary in a small bay in Northwestern Australia. Journal of Molluscan Studies 52: 83–90. Wells, F.E. (ed). 1997. Proceedings of the Seventh International Marine Biological Workshop: The Marine Flora and Fauna of the Houtman Abrolhos Islands, Western Australia. Western Australian Museum, Perth. Wells, F.E. 1998. Part 3. Mollusks of Milne Bay Province, Papua New Guinea. In Werner, T.B., and G.R. Allen (eds.) A Rapid Biodiversity Assessment of the Coral Reefs of Milne Bay Province, Papua New Guinea. RAP Working Papers 11. Conservation International, Washington, D.C. Wells, F.E. 2000. Mollusks of the Calamianes Islands, Palawan Province, Philippines. Pp. 27–30, 81–94 in Werner, T.B., and G.R. Allen (eds.) A Marine Rapid Biodiversity Assessment of the Calamianes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Conservation International, Washington, D.C. Wells, F.E. 2002a. Mollusks of the Gulf of Tomini, Sulawesi, Indonesia. Pp. 38–43, 81–97 in Allen, G.R., and S.A. McKenna (eds.) A Marine Rapid Assessment of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment 20. Conservation International, Washington, D.C. Wells, F.E. 2002b. Mollusks of the Raja Ampat Islands, Papua Province, Indonesia. Pp. 37–45, 113–131 in McKenna, S.A., G.R. Allen, and S. Suryadi (eds.) A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, D.C. Wells, F.E. 2002c. Centres of species richness and endemism of shallow water marine mollusks in the tropical Indo-West Pacific. Pp. 941–945 in Kasim Moosa, M., S. Soemodihardjo, A. Soegiarto, K. Romimohtarto, A. Nontji, Soekrano, and Suharsono (eds.) Proceedings of the 9th International Coral Reef Symposium, Bali, Indonesia, 23–27 October 2000. Ministry of Environment, Indonesian Institute of Sciences and International Society for Reef Studies, Jakarta, Indonesia. Wells, F.E., and S.M. Slack-Smith. 1981. Zonation of mollusks in a mangrove swamp in northwestern Australia. Pp. 265–274 in Biological Survey of Mitchell Plateau and Admiralty Gulf, Kimberley, Western Australia. Western Australian Museum, Perth. Wells, F.E., D.I. Walker, and D.S. Jones. 2003. Proceedings of the Eleventh International Marine Biological Workshop: The Marine Flora and Fauna of Dampier, Western Australia. Western Australian Museum, Perth. Wells, F.E., D.I. Walker, and G. Kendrick. 2005. Proceedings of the Twelfth International Marine Biological Workshop: The Marine Flora and Fauna of Esperance, Western Australia. Western Australian Museum, Perth. Wells, F.E., D.I. Walker, H. Kirkman, and R. Lethbridge (eds.). 1990/1991. Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia. 2 vols. Western Australian Museum, Perth. Wells, F.E., D.I. Walker, H. Kirkman, and R. Lethbridge, R. (eds.). 1993. Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and Fauna of Rottnest Island, Western Australia. Western Australian Museum, Perth.

................. 16157$

CH35

02-08-07 10:39:19

PS

PAGE 514

4.3. Insects of Papua . terrestrial invertebrates are tremendously diverse in Papua, and play vital roles in providing ecosystem services, but are poorly known. Insects are interesting for conservation studies for the following reasons: (1) generality of distribution: insects are found in almost every conceivable habitat and niche; (2) within this overall versatility, there is much specialization; (3) many taxa show rapid responses to environmental perturbation; (4) some taxa are readily identifiable without specialized training; (5) many taxa are good indicators of areas of endemism; and (6) many taxa are readily sampled with quantitative methods, providing high quality data for statistical analyses (see Brown 1991; Holloway and Stork 1991; Kremen et al. 1993; Miller and Rogo 2002; Sutton and Collins 1991).

I

History Frodin and Gressitt (1982; Chapter 2.1) have presented excellent descriptions of the history of biological exploration in New Guinea and only the major elements are repeated here (see also van Steenis-Kruseman 1950). The Papuan fauna cannot be understood in isolation because much of the fauna is shared with Papua New Guinea, the Solomon Islands, and Australia. Most groups of insects have been more extensively sampled in Papua New Guinea than Papua, so it is often necessary to extrapolate from knowledge of Papua New Guinea to understand the Papuan fauna. The entomological literature for the New Guinea region was compiled by Gressitt and Szent-Ivany (1968), and is available online in an updated version at http://entomology.si.edu. The first comprehensive insect collecting in Papua was undertaken by European collectors starting in the 1870s and continuing into the 1930s. These collectors included E. Cheesman, W. Doherty, H. A. Lorentz, E. Mayr, A. S. Meek, W. G. Meek, A. E. Pratt, H. Pratt, and A. F. R. Wollaston. The Third Archbold Expedition (1938–1939) was the only one of the Archbold Expedition series that visited Papua and it provided an unsurpassed sample of the fauna of the Mamberamo Basin and headwaters (Archbold et al. 1942). During World War II, many United States and Australian entomologists collected in coastal Papua. In the 1950s, J. L. Gressitt of the Bishop Museum organized surveys of Papuan insects. Little research was done on insects in Papua from the 1960s through the 1980s. Terrestrial insects have been included in conservation surveys starting in the 1990s, but only in a small way (e.g., Polhemus and Polhemus 2000; Rosariyanto et al. 2002; van Mastright and Rosariyanto 2002; Oppel 2006). Major collections of Papua’s terrestrial invertebrates are housed at Bishop MuMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

515

................. 16157$

CH36

02-08-07 10:39:31

PS

PAGE 515

.

516 /

seum, Honolulu (BPBM); Museum Zoologense Bogeriense, Bogor (MZB); Natural History Museum (formerly British Museum of Natural History), London (NHM); National Museum of Natural History (formerly Rijksmuseum van Natuurlijke Historie), Leiden (RNH); American Museum of Natural History, New York (AMNH); Smithsonian Institution, Washington DC (USNM); and the University of Amsterdam. A small collection, which should be upgraded, exists at Cenderawasih University, Manokwari campus, Papua.

Status of General Knowledge Knowledge of insects and other invertebrates of New Guinea varies tremendously among taxonomic groups and localities, according to the history of sampling and study, and the biological complexity of the group. For example, butterflies are well known in some areas (e.g., Gotts and Pangemanan 2001; van Mastrigt and Rosariyanto 2005), but little is known about soil arthropods at most sites in New Guinea (Hammen 1983). The components involved in understanding insects include the following: 1. Sampling of basic diversity: collecting and preservation for study. Basic sampling has taken place for many groups, although the smaller and more difficult to study taxa are almost always undersampled. Geographic coverage of sampling is poor in Papua, even for butterflies. 2. Systematic study of these samples: diagnosis of species and other taxonomic units. The quality and quantity of systematic study varies with historical interest from group to group. Some taxa are relatively well known, but information transfer is a problem for all groups. The recent text on Australian insects (CSIRO 1991) is very helpful for understanding the New Guinea fauna. 3. Field studies on the biology, ecology, and geographic distribution of species. Except for species of economic interest to agriculture (e.g., Simon Thomas 1962; Ubaidillah 1991), forestry (Nair 2000), or medicine (Bangs and Subianto 1999; Owen 2005), little is known about the ecology of Papua’s insects. In addition, little is known about the distribution of insects in ecological space, including issues such as host specificity, and the distribution of individuals and biomass per species within ecosystems (Novotny et al. 2002a, 2006).

However, as noted above, the great diversity of insects is a positive feature, allowing researchers to choose specific groups that can answer specific questions in a statistically meaningful manner. Modern information management techniques provide at least a partial solution to the information transfer problem.

Status of Taxonomic Knowledge As noted above, taxonomic knowledge of New Guinea insects varies greatly from group to group. There is no comprehensive review for any major group except the flies, and the bibliography by Gressitt and Szent-Ivany (1968) is the only broad

................. 16157$

CH36

02-08-07 10:39:31

PS

PAGE 516

Insects of Papua / 517

attempt to synthesize knowledge of New Guinea insects. The following is a brief overview of the state of taxonomic knowledge of New Guinea insects, including major references. Major taxonomic units follow the recent text on Australian insects (CSIRO 1991) which is an important general review. Simon Thomas (1962) reviewed agricultural entomology in Papua, and Hammen (1983) includes a review of the meager knowledge of soil arthropods. ‘‘Very poorly known’’ in the list below indicates that only very basic surveys have been done, little is known about biogeography and distribution, and no ecological studies have been done (except sometimes for selected pest organisms). There is no comprehensive list of insects recorded from Papua (or New Guinea), and any estimate must be based on assumptions and extrapolation (e.g., Novotny et al. 2002c). Miller (1996) estimated the total insect fauna of New Guinea might be 300,000 species (Table 4.3.1). The diversity of insects in Australia has recently been reviewed by Austin et al. (2004) and Yeates et al. (2003), who reached the conclusion that the insect fauna of Australia is something over 200,000 species. In light of the new Australian estimates, and the lower host specificity estimates in Novotny et al. (2002c), the insect fauna of New Guinea is likely less than 300,000 species, of which many remain to be described. Larger insects in the lowland areas are relatively well known, at least in the sense of having species names (Novotny et al. 2005; S. Miller, unpublished data). However, it is often impossible to associate the species name with a specimen without reference to historic collections in Europe because the species have never been adequately characterized in the literature. But the fauna of larger insects at higher elevations (above about 1,000 m) is poorly known (Novotny et al. 2005; Riedel 2001; S. Miller, unpublished data), and the fauna of small insects (as well as mites and other tiny arthropods) is poorly known throughout Papua. There is very little overlap in species between the lowlands and higher elevations, although the nature of the transition in faunas with elevation is not yet adequately explored (Allison et al. 1993; Novotny et al. 2005).

Table 4.3.1. Numbers of invertebrate species known from New Guinea Taxon Terrestrial flatworms Freshwater rotifers Nematodes of plants Terrestrial mollusks Leeches (part)

Number of species 4

Segers and de Meester 1994

63

Bridge and Page 1984

1,000

Cowie 1993

5 42

Onychophora

7

Insects

de Beauchamp 1972

135

Earthworms

van der Lande 1993 Nakamura 1992 van der Lande 1993

300,000

................. 16157$

Source

Miller 1996

CH36

02-08-07 10:39:32

PS

PAGE 517

.

518 /

Order Collembola (Springtails): Very poorly known in Papua (Yoshii and Suhardjno 1992). Order Protura: Very poorly known for Papua (Tuxen 1964). Order Diplura: Very poorly known for Papua (Paclt 1985). Order Archaeognatha ( Microcoryphia) (Bristletails): Very poorly known for Papua (Strum 1999, Strum and Machida 2001). Order Thysanura (Silverfish): Very poorly known for Papua (Paclt 1982). Order Ephemeroptera (Mayflies): Very poorly known for New Guinea (Edmunds and Polhemus 1990). Order Odonata (Dragonflies and damselflies): Relatively well known from extensive work of Lieftinck (1949). Order Plecoptera (Stoneflies): The only record in Papua is an unidentified species recently collected by D. A. Polhemus (pers. comm.). Order Blattodea ( Blattaria) (Cockroaches): Relatively poorly known. There is only a scattered literature for Papua, partially reviewed by Roth (2003). Order Isoptera (Termites): Relatively poorly known for Papua. Cataloged by Snyder (1949). Nasute termites of Papua New Guinea revised by Roisin and Pasteels (1996). Order Mantodea (Praying mantids): Relatively well known for New Guinea (Beier 1965; Rentz 1996). Order Dermaptera (Earwigs): Relatively poorly known for Papua; see world checklist by Steinmann (1989). Order Orthoptera (Grasshoppers, locusts, katydids, crickets): Larger taxa relatively well known in Papua, but the literature for New Guinea is scattered. See Rentz (1996) and Willemse (2001). Order Phasmatodea (Stick insects): Relatively well known in New Guinea (van Herwaarden 1998; Nakata 1961; Rentz 1996). Order Embioptera ( Embiidina) (Web-spinners): Relatively poorly known in Papua (Ross 1948). Order Zoraptera: Not yet known to occur in New Guinea (Smithers, in CSIRO 1991). Order Psocoptera (Psocids, booklice): Generally poorly known in New Guinea, although some groups recently revised (Lienhard and Smithers 2002; Smithers and Thornton 1981). Order Phthiraptera (including Mallophaga and Anoplura) (Lice): Generally poorly known in New Guinea (Durden and Musser 1994; Ferris 1951; Price 2003). Order Hemiptera (including Heteroptera and Homoptera) (Bugs, leafhoppers, cicadas, aphids, scale insects, etc.): Knowledge very uneven for New Guinea. Some groups have modern revisions (especially some Cicadidae, Coccoidea, Miridae, Flatidae, Cicadellidae, aquatic Heteroptera, etc.), but others do not. See Stonedahl and Dolling (1991) for literature on Heteroptera. Order Thysanoptera (Thrips): Generally poorly known for New Guinea but partially treated in recent revisions by L. A. Mound and others. Order Megaloptera (Alderflies, dobsonflies): Not yet known from New Guinea. Order Raphidioptera (Snakeflies): Not yet known from New Guinea.

................. 16157$

CH36

02-08-07 10:39:32

PS

PAGE 518

Insects of Papua / 519 Order Neuroptera (Lacewings, etc.): Relatively well known in New Guinea because of publications by T. R. New and colleagues, reviewed by New (2003). Order Coleoptera (Beetles): A very diverse group in New Guinea, ranging from relatively well known (e.g., Carabidae, Chyrsomelidae) to very poorly known (e.g., Scarabaeidae, Curculionidae). General review by Gressitt and Hornabrook (1977). Several large families reviewed by Bigger and Schofield (1983). Order Strepsiptera: A small parasitic group reviewed by Kifune and Hirashima (1989) and Kathirithamby (1989). Order Mecoptera (Scorpionflies): Not known from New Guinea (Penny and Byers 1979). Order Siphonaptera (Fleas): Relatively well known in New Guinea; monographed by Holland (1969). Order Diptera (Flies): A very diverse group in New Guinea, ranging from relatively well known (e.g., Culicidae, Simuliidae; e.g., Takaoka 2003) to very poorly known (e.g., Chironomidae). Reviewed by Evenhuis (1989) and Oosterbroek (1998). Order Trichoptera (Caddisflies): Reviewed by Neboiss (1986, 1989), but many taxa remain to be sampled and described. Order Lepidoptera (Moths and butterflies): A very diverse group in New Guinea, ranging from relatively well known (e.g., butterflies) to very poorly known (e.g., Microlepidoptera). General review and identification guide by Holloway et al. (2001). Butterflies partially reviewed by Parsons (1998) and van Mastrigt and Rosariyanto (2005); Papilionidae are reviewed by Ubaidillah et al. (1994). Microlepidoptera reviewed by Diakonoff (1952–1955), but many taxa remain to be described. Order Hymenoptera (Wasps, bees, ants, sawflies): A very diverse group in New Guinea, ranging from relatively well known (e.g., ants) to very poorly known (e.g., many parasitic wasps).

Terrestrial invertebrates other than insects have generally been very poorly sampled and studied in New Guinea. It is worth noting that terrestrial and freshwater mollusks are better known in Papua than in Papua New Guinea, although they remain poorly understood (Cowie 1993). For further discussion of invertebrate diversity see Ewers (1973), Miller (1996), and Sekhran and Miller (1996).

Biogeography New Guinea and surrounding islands provide major challenges for understanding the complex interrelationships of ‘‘mobile organisms in a geologically complex area’’ (Holloway 1982). The island of New Guinea has evolved as a major land area only over the past ten or so million years, through fusion and compression of inner and outer Melanesian island arcs between the northward moving Australian continent on the Indian Ocean tectonic plate and the westward moving margin of the Pacific tectonic plate. Only in the Pliocene and Pleistocene were its mountain ranges uplifted to the snow line, to be glaciated extensively during the latter pe-

................. 16157$

CH36

02-08-07 10:39:33

PS

PAGE 519

.

520 /

riod. Despite this geological youth, the insect fauna has attained a diversity that appears to be equivalent to that of the much older tropical lands of Southeast Asia (Gressitt 1982b). The majority of this fauna has its closest relationships with that of Asia, with the genera tending to be fewer in number but more species-rich. For instance, the majority of moth genera endemic to Melanesia are restricted to, or have their species richness strongly centered in, New Guinea (Holloway 1984b). Rapid development of high diversity in a geologically unstable environment with high relief is counter to some hypotheses of tropical diversity, particularly those based on environmental stability over long periods of geological time. Therefore studies of the biogeographic origins of the New Guinea fauna and speciation patterns within it are likely to throw considerable light on the processes leading to increases in species richness (Holloway 1991). Although many authors have addressed the biogeographic relationships of the New Guinea fauna to those of adjacent regions, particularly in regard to the importance of Australian versus Asian elements, few authors have attempted analyses of biogeography within New Guinea (especially patterns of regional endemism). The paucity of data for regionalization of New Guinea became very clear in an extensive literature survey for the Papua New Guinea Conservation Needs Assessment, an attempt to identify the areas within the country of greatest importance to biodiversity conservation. Even among the relatively well known ground beetles (Coleoptera: Carabidae), patterns based only on literature records proved misleading. For example, of 11 carabids that were known only from the Northern Province lowlands (putative regional endemics), four were represented in the Bishop Museum by specimens from outside of the Northern Province lowlands (Miller 1993). Thus, previously unpublished data from museum collections will be important to a comprehensive view of invertebrate biogeography in New Guinea. The best analyses of regional patterns are those for cicadas (de Boer 1995; de Boer and Duffels 1996a,b; Duffels and de Boer 1990; and Chapter 4.4) and aquatic insects (Chapter 2.5; Polhemus 1996, 1998). There are relatively few modern revisions (i.e., those applying modern species concepts to adequate samples) of speciose groups in New Guinea. Among the taxa for which such data exist are some groups of mosquitoes, black flies, sarcophagid flies, aquatic bugs, ground beetles, chrysomelid beetles, pyraloid moths, butterflies, and cicadas. Based on intensive studies of cicada distributions, Papua can be subdivided into four main areas of endemism, each with its own distinct biota (see Chapter 4.4): the Vogelkop peninsula, the northern mountain ranges, the central mountain ranges, and south New Guinea. The latter three continue into Papua New Guinea. For most groups of invertebrates, all of Papua needs intensive collecting using modern sampling techniques. Even where samples do exist, they are often limited to material from early expeditions and usually to larger species (e.g., butterflies, cicadas). But a huge area of Papua has received no serious sampling for insects. Some of the areas that are especially poorly known are: (1) The Vogelkop area, other than the Arfak Mountains; areas around Sorong, Fakfak, and Manokwari, and the Wasior Peninsula, are almost unknown; (2) Papua north of the Central

................. 16157$

CH36

02-08-07 10:39:33

PS

PAGE 520

Insects of Papua / 521

Range is largely unknown except for the coastal areas (especially around Nabire and Jayapura) and the Cyclops Mts; (3) The Central Range from the Weyland Mts to the Baliem Valley, and the Star Mts; and (4) the southeastern lowlands from the Lorentz National Park to the Wasur area (including Yos Sudarso (Kimaam) Island) remain unknown (Miller 1998).

Ecology and Conservation So little is known about the ecology of most New Guinea insects that it is difficult to identify ecologically critical areas. In general, the highest diversity is found in primary forest, but little study has been devoted to the ecology of savanna and wetlands insects in New Guinea and it is possible that important patterns of diversity are being overlooked (e.g., Pimm and Gittleman 1992). Because of the need for calcium for shell formation, the highest diversity of land snails is expected in limestone areas with high soil pH (Andrews and Little 1982; Cowie 1993). Preservation of isolated limestone outcrops will be especially important in the conservation of maximum diversity of land snails. The impact of habitat disturbance on insects depends upon, among other things, the life history strategies of the insect species involved. Although this has received little study in New Guinea (Bowman et al. 1990; Novotny et al. 2004), most native insects respond poorly to severe habitat degradation. Loss of diversity from disturbance of forests affects some taxonomic groups more seriously than others (see Miller and Holloway 1992 and Hill et al. 1995 for examples from other Indonesian islands). Insects with narrow dietary requirements or other habitat specialization tend to be most readily affected by habitat degradation. However, some insect populations recover relatively well after disturbance. For example, in Seram, moth diversity in shifting cultivation areas that were abandoned several decades ago, which are now well forested, is comparable with that of undisturbed forest at similar altitudes (Miller and Holloway 1992). It is therefore possible that a sustainable cycle of forest exploitation or farming that permits a full regeneration cycle will only temporarily depress biodiversity, although this may be dependent on the proportion of climax forest present throughout the progression of such cycles. However, conversion of lowland forest to field or cash crop plantations will depress moth diversity to a much lower level, as seen in Sulawesi (Miller and Holloway 1992).

Economically Important Species The most widely recognized economically important species of insects are pests of agriculture and forestry, and vectors of diseases. Although a great many species have some negative economic importance in Papua (Simon Thomas 1962), they constitute only a very minor fraction of the total fauna. However, as primary forest resources are depleted in the future, and more plantations (particularly of

................. 16157$

CH36

02-08-07 10:39:34

PS

PAGE 521

.

522 /

leguminous trees) are established, these economic activities and developments will be affected by insect attack, particularly from defoliating insects. In Papua, most of the agricultural pest insects are native, and new ‘‘pests’’ frequently emerge from the indigenous fauna in response to natural or humaninduced changes (Simon Thomas 1962, Kumar 2001). Szent-Ivany (1961) showed that while 98% of the then insect pests of Hawai’i were introduced by humans, less than 3% (1 of 34) of the then major agricultural insect pests in New Guinea were introduced. Furthermore, many of the economic crops (e.g., sugar) are native to the region. Thus a high ‘‘pest’’ potential, especially in crop monocultures, exists in this region. The current emphasis in forestry to plant legumes and Eucalyptus in monoculture after natural forest is cleared, will almost inevitably lead to major pest problems because these crops are in the preferred host families for many moths. Thus, for natural resource management, it will be important to have basic data on the insect fauna of Papua. Pest exclusion and detection programs, such as inspections and quarantines of incoming commodities, regular surveys for target pests, and local diagnostic services supported by international laboratories are important to prevent future problems. Large-scale conversion of primary forest to other uses can result in outbreaks of a variety of pests, especially disease vectors. This should be factored into costbenefit analyses of programs. Ross River alphavirus and Murray River encephalitis present likely prospects for intensified disease prevalence wherever lowland forests are disturbed and people aggregated (Jenkins 1992). Clearing of native forest barriers has allowed pest locusts to move into new areas in Africa and could happen in New Guinea (Miller 1993). Many of the large, colorful insects of Papua, especially butterflies, have a commercial market value. Butterfly farming activities in New Guinea are generally considered successful (e.g., studies in Papua New Guinea by Clark and Landford 1991; Collins and Morris 1985:29; Orsak 1993; Parsons 1998; Sekhran and Miller 1996; and in Papua by Mercer 1997; Morris 1986; and Parsons 1982). Nevertheless, a detailed analysis of the economic and conservation impact of butterfly farming would be useful in planning future expansion of the program (e.g., Goldstein 1991). It is clear that butterfly farming has raised the awareness, both locally and internationally, of conservation issues and of the potential value of New Guinean native fauna. Butterfly farming could clearly benefit from more research on effective methods of rearing rare species in captivity. Insects are also important food resources for many local people (Mercer 1997; Ponzetta and Paoletti 1997). Perennial malaria, vectored by mosquitoes in the Anopheles punctulatus complex, is a major public health problem for most coastal, lowland, and foothill populations in Papua. Malaria at higher elevations (above 1,500 m) is considered intermittent and highly unstable, providing a constant threat of epidemics (Bangs and Subianto 1999). Relatively few species of zoonotic parasites have been recorded in humans in Papua New Guinea (Owen 2005). Among them are flies (Chrysomya bezziana), bed bugs (Cimex sp.), fleas (Ctenocephalides spp.), and mites (Leptotrombidium spp., Sarcoptes scabiei, and Demodex sp.). The paucity of

................. 16157$

CH36

02-08-07 10:39:35

PS

PAGE 522

Insects of Papua / 523

zoonotic parasite species can be attributed to long historical isolation of the island of New Guinea and its people, and the absence until recent times of large placental mammals other than pigs and dogs (Owen 2005).

Threats The largest important threat to most insects is habitat disturbance. In most cases, this means destruction of the integrity of primary forests and other natural habitats. Some species might be threatened by alien species (e.g., by predation, parasitism, or competition). Alien species are probably a minor threat to native insects in most of mainland Papua, but can be significant in areas of heavy human disturbance (e.g., Snelling 1998) and islands that host many endemic species (e.g., Howarth and Ramsay 1991; Nishida and Evenhuis 2000; Williams 1994). However, alien species can be difficult to detect before they become widespread. The arboreal termite originally known as the New Guinea endemic Nasutitermes polygynus Roisin and Pasteels was recently found to be an accidental introduction of Nasutitermes corniger (Motschulsky) from the New World tropics (Scheffrahn et al. 2005). Biological control introductions should be carefully screened to prevent excessive impact on nontarget organisms (Howarth 1991). Overcollecting is a threat only to large species with very restricted populations, such as birdwing and Delias butterflies. Many butterflies are already protected by Indonesian law (Simbolon and Iswari 1990), although it is not clear how effective this protection has been. The effects of pollution on invertebrates in Papua have received almost no study, but may prove to be important. Chemical pollutants from mining and increased sedimentation from deforestation are likely to heavily impact aquatic insects, crustacea, and mollusks. Chemical pesticides used for agriculture, forestry, and vector control can impact both terrestrial and aquatic invertebrates. Note: Since this chapter was written, the Papua Insects Foundation has started a web site at www.papua-insects.nl that includes diverse information on Papua insects.

Acknowledgments A. Allison, M. Amir, Y. Basset, R. Cowie, A. de Boer, K. Darrow, H. Duffels, N. L. Evenhuis, J. D. Holloway, K. Kami, H. van Mastrigt, E. G. Munroe, T. R. New, Y. Novotny, L. J. Orsak, M. Parsons, D. A. Polhemus, F. C. Thompson, G. Weiblen, A. Yaku, and R. I. Vane-Wright contributed information or ideas for this paper over many years of study, which have been supported by the Bishop Museum, National Science Foundation, and Smithsonian Institution.

Literature Cited Allison, A. 1992. The role of museums and zoos in conserving biological diversity in Papua New Guinea. Pp. 59–63, 74 in Pearl, M., et al. (eds.) Conservation and

................. 16157$

CH36

02-08-07 10:39:35

PS

PAGE 523

.

524 /

Environment in Papua New Guinea: Establishing Research Priorities. Embassy of Papua New Guinea, Washington, D.C., and Wildlife Conservation International, Bronx, New York. Allison, A., G.A. Samuelson, and S.E. Miller. 1993. Patterns of beetle species diversity in New Guinea rain forest as revealed by canopy fogging: preliminary findings. Selbyana 14: 16–20. Andrews, E.B., and C. Little. 1982. Renal structure and function in relation to habitat in some cyclophorid land snails from Papua New Guinea. Journal of Molluscan Studies 48: 124–143. Archbold, R., A.L. Rand, and L.J. Brass. 1942. Results of the Archbold Expeditions, no. 41. summary of 1938–39 New Guinea expedition. Bulletin of the American Museum of Natural History 79: 197–288. Ashe, P. 1990. Ecology, zoogeography and diversity of Chironomidae (Diptera) in Sulawesi with some observations relevant to other aquatic insects. Pp. 261–268 in Knight, W.J., and J.D. Holloway (eds.) Insects and the Rain Forests of South East Asia (Wallacea). Royal Entomological Society, London. Austin, A.D., D.K. Yeates, G. Cassis, M.J. Fletcher, J. LaSalle, J.F. Lawrence, P.B. McQuillan, L.A. Mound, D.J. Bickel, P.J. Gullan, D.F. Hales, and G.S. Taylor. 2004. Insects ‘‘Down Under’’—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43: 216–234. Bangs, M.J., and D.B. Subianto. 1999. El Nin˜o and associated outbreaks of severe malaria in highland populations in Irian Jaya, Indonesia: a review and epidemiological perspective. Southeast Asian Journal of Tropical Medicine and Public Health 30: 608–619. Beier, M. 1965. Die Mantodeen Neu-Guineas. Pacific Insects 7: 473–502. Bigger, M., and P. Schofield. 1983. Checklist of Cerambycidae, Curculionidae, Attelabidae, Scolytidae and Platypodidae of Melanesia. Overseas Development Administration (London) Miscellanous Report 60: 1–62. Bowman, D.M.J.S., J.C.Z. Woinarski, D.P.A. Sands, A. Wells, and V.J. McShane. 1990. Slash-and-burn agriculture in the wet coastal lowlands of Papua New Guinea: response of birds, butterflies and reptiles. Journal of Biogeography 17: 227–239. Bridge, J., and S.L.J. Page. 1984. Plant nematode pests of crops in Papua New Guinea. Journal of Plant Protection in the Tropics 1: 99–109. Brown, K.S., Jr. 1991. Conservation of neotropical environments: Insects as indicators. Pp. 349–404 in Collins, N.M., and J.A. Thomas (eds.) The Conservation of Insects and Their Habitats. Academic Press, London. Clark, P.B., and A.D. Landford. 1991. Farming insects in Papua New Guinea. International Zoo Yearbook 30: 127–131. Collins, N.M., and M.G. Morris. 1985. Threatened Swallowtail Butterflies of the World: The IUCN Red Data Book. IUCN, Cambridge. Cowie, R.H. 1993. Non-Marine Mollusca of Papua New Guinea. Pp. 311–320 in Beehler, B.M. (ed.) Papua New Guinea Conservation Needs Assessment, vol. 2. Papua New Guinea Department of Environment and Conservation, Boroko. Cranston, P.S. 1990. Biomonitoring and invertebrate taxonomy. Environmental Monitoring and Assessment 14: 265–273. CSIRO. 1991. The Insects of Australia: A Textbook for Students and Research Workers. 2nd ed. 2 vols. Melbourne University Press, Melbourne. de Beauchamp, P. 1972. Planaires terrestres de Nouvelle-Guine´e. Cahiers du Pacifique 16: 181–192. de Boer, A.J. 1989. The taxonomy and biogeography of the bloetei group of the genus Baeturia Sta˚l, 1866 (Homoptera, Tibicinidae). Beaufortia 39: 1–43.

................. 16157$

CH36

02-08-07 10:39:36

PS

PAGE 524

Insects of Papua / 525 de Boer, A.J. 1995. Islands and cicadas adrift in the west-Pacific. Biogeographic patterns related to plate tectonics. Tijdschrift voor Entomologie 138: 169–244. de Boer, A.J., and J.P. Duffels. 1996a. Biogeography of Indo-Pacific cicadas east of Wallace’s Line. Pp. 297–330 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. de Boer, A.J., and J.P. Duffels. 1996b. Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific: a geotectonic explanation. Palaeogeography, Palaeoclimatology, Palaeoecology 124: 153–177. Diakonoff, A. 1952–1955. Microlepidoptera of New Guinea. Results of the Third Archbold Expedition (1938–39). Parts I–V. Verh. Akad. Wet Amst. (Natuurk.), ser. 2, 49 (2): 1–167; 49 (3): 1–166; 49 (4): 1–164; 50 (1): 1–191; 50 (3): 1–210. Duffels, J.P. 1977. A revision of the genus Diceropyga Sta˚l, 1870 (Homoptera, Cicadidae). Mono. Ned. Ent. Vereen. 8: 1–227. Duffels, J.P. 1983a. Distribution patterns of Oriental Cicadoidea (Homoptera) east of Wallace’s Line and plate tectonics. GeoJournal 7: 491–498. Duffels, J.P. 1983b. Taxonomy, phylogeny and biogeography of the genus Cosmopsaltria, with remarks on the historic biogeography of the subtribe Cosmopsaltriaria (Homoptera: Cicadidae). Pacific Insects Monograph 39: 1–127. Duffels, J.P. 1986. Biogeography of Indo-Pacific Cicadoidea: a tentative recognition of areas of endemism. Cladistics 2: 318–336. Duffels, J.P., and A.J. de Boer. 1990. Areas of endemism and composite areas in East Malesia. Pp. 249–272 in Baas, P., K. Kalkman, and R. Geesink (eds.) The Plant Diversity of Malesia. Kluwer Academic Publishers, Dordrecht. Durden, L.A., and G.G. Musser. 1994. The sucking lice (Insecta, Anoplura) of the world: a taxonomic checklist with records of mammalian hosts and geographical distributions. Bulletin of the American Museum of Natural History 218: 1–90. Edmunds, G.F., Jr., and D.A. Polhemus. 1990. Zoogeographical patterns among mayflies (Ephemeroptera) in the Malay Archipelago, with special reference to Celebes. Pp. 49–56 in Knight, W.J., and J.D. Holloway (eds.) Insects and the Rain Forests of South East Asia (Wallacea). Royal Ent. Soc., London. Erwin, T.L. 1975. Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini), Part III: systematics, phylogeny, and zoogeography of the genus Tachyta Kirby. Smithsonian Contributions to Zoology 208: 1–68. Evenhuis, N.L. (ed.). 1989. Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Museum Press and E.J. Brill, Honolulu and Leiden. Ewers, W.H. 1973. Bibliography of invertebrate animals from New Guinea. Science in New Guinea 1: 41–46. Ferris, G.F. 1951. The sucking lice. Memoirs of the Pacific Coast Entomological Society 1: 1–321. Frodin, D.G., and J.L. Gressitt. 1982. Biological exploration of New Guinea. Pp. 87–130 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Goldstein, J.H. 1991. The prospects for using market incentives to conserve biological diversity. Environmental Law 21: 985–1014. Gotts, R.I.C., and N. Pangemanan. 2001. Mimika butterflies: a guide to the butterflies of the Mimika subdistrict of Papua. PT Freeport Indonesia Biodiversity Centre, Timika, Indonesia. Gressitt, J.L. 1982a. Pacific-Asian biogeography with examples from the Coleoptera. Entomologica Generalis 8: 1–11.

................. 16157$

CH36

02-08-07 10:39:37

PS

PAGE 525

.

526 /

Gressitt, J.L. 1982b. Zoogeographical summary. Pp. 897–918 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Gressitt, J.L., and R.W. Hornabrook. 1977. Handbook of Common New Guinea Beetles. Handbook 2. Wau Ecology Institute, Wau. Gressitt, J.L., and J.J.H. Szent-Ivany. 1968. Bibliography of New Guinea entomology. Pacific Insects Monograph 18: 1–674. Hammen, L. van der. 1983. Contribution to the knowledge of the soil-fauna of New Guinea. Zoologische Verhandelingen 206:1–36, pl. 1–19. Haugum, J., and A.M. Low. 1978–1985. A Monograph of the Birdwing Butterflies. 2 vols. Scandinavian Science Press, Klampenborg, Denmark. Hebert, P.D.N. 1980. Moth communities in montane Papua New Guinea. Journal of Animal Ecology 49: 593–602. Hill, J.K., K.C. Hamer, L.A. Lace, and W.M.T. Banham. 1995. Effects of selective logging on tropical forest butterflies on Buru, Indonesia. Journal of Applied Ecology 32: 752–760. Holland, G.P. 1969. Contribution towards a monograph of the fleas of New Guinea. Memoirs of the Entomological Society of Canada 61: 1–77. Holloway, J.D. 1977. The Lepidoptera of Norfolk Island: Their Biogeography and Ecology. W. Junk, The Hague. Holloway, J.D. 1979. A Survey of the Lepidoptera, Biogeography and Ecology of New Caledonia. W. Junk, The Hague. Holloway, J.D. 1983. Insect surveys—an approach to environmental monitoring. Pp. 239–261 in Atti XII Cong. Naz. Ital. Ent. (Rome, 1980). Holloway, J.D. 1984a. Moths as indicator organisms for categorizing rain-forest and monitoring changes and regeneration processes. Pp. 235–242 in Chadwick, A.C., and S.L. Sutton (eds.) Tropical Rain-Forest: The Leeds Symposium. Leeds Philosophical and Literary Society, Leeds. Holloway, J.D. 1984b. The larger moths of the Gunung Mulu National Park: a preliminary assessment of their distribution, ecology, and potential as environmental indicators. Sarawak Museum Journal 30: 149–190. Holloway, J.D. 1984c. Lepidoptera and the Melanesian Arcs. Pp. 129–169 in Radovsky, F.J., et al. (eds.) Biogeography of the Tropical Pacific. Bishop Museum Special Publication 72. Bishop Museum, Honolulu. Holloway, J.D. 1987. Bracken-feeding Geometridae in the genus Idiodes Guenee, 1857, and allied taxa in the tribe Lithinini (Lepidoptera). Tinea 12 (supplement): 242–248. Holloway, J.D. 1989. Moths. Pp. 437–453 in Lieth, H. (ed.) Tropical Rain Forest Ecosystems. Ecosystems of the World, vol. 14B. Elsevier, Amsterdam. Holloway, J.D. 1991. Patterns of moth speciation in the IndoAustralian archipelago. Pp. 340–372 in Dudley, E.C. (ed.) The Unity of Evolutionary Biology: Proceedings of the Fourth International Congress of Systematic and Evolutionary Biology. Dioscorides Press, Portland, Oregon. Holloway, J.D. 1993. Aspects of the biogeography and ecology of the Seram moth fauna. Pp. 91–114 in Edwards, I.D., A.A. Macdonald, and J. Proctor (eds.) The Natural History of Seram, Maluku, Indonesia. Intercept, Andover. Holloway, J.D., and H.S. Barlow. 1992. Potential for loss of biodiversity in Malaysia, illustrated by the moth fauna. Pp. 293–311 in Kadir, A.A.S.A., and H.S. Barlow (eds.) Pest Management and the Environment in 2000. CAB International, Wallingford, U.K. Holloway, J.D., G.S. Robinson, and K.R. Tuck. 1990. Zonation in the Lepidoptera of northern Sulawesi. Pp. 153–166 in Knight, W.J., and J.D. Holloway (eds.) Insects and the Rain Forests of South East Asia (Wallacea). Royal Entomological Society, London.

................. 16157$

CH36

02-08-07 10:39:37

PS

PAGE 526

Insects of Papua / 527 Holloway, J.D., and N.E. Stork. 1991. The dimensions of biodiversity: the use of invertebrates as indicators of man’s impact. In Hawksworth, D.L. (ed.) The Ecological Foundations of Sustainable Agriculture. CAB International, Wallingford, U.K. Howarth, F.G. 1991. Environmental impacts of classical biological control. Ann. Rev. Ent. 36: 485–509. Howarth, F.G., and G.W. Ramsay. 1991. The conservation of island insects and their habitats. Pp. 71–107 in Collins, N.M., and J.A. Thomas (eds.) The Conservation of Insects and Their Habitats. Academic Press, London. Hurlbert, S.H., and A. Villalobos-Figueroa (eds.). 1982. Aquatic Biota of Mexico, Central America and the West Indies. San Diego State University, San Diego, California. Janzen, D.H. 1984. Two ways to be a tropical big moth: Santa Rosa saturniids and sphingids. Oxford Surveys in Evolutionary Biology 1: 85–140. Jenkins, C. 1992. Environmental change and human health in Papua New Guinea. Pp. 75–82 in Pearl, M., et al. (eds.) Conservation and Environment in Papua New Guinea: Establishing Research Priorities. Embassy of Papua New Guinea, Washington, D.C., and Wildlife Conservation International, Bronx, New York. Kathirithamby, J. 1989. Review of the order Strepsiptera. Systematic Entomology 14: 41–92. Kifune, T., and Y. Hirashima. 1989. Taxonomic studies on the Strepsiptera in the collection of Bishop Museum (Notulae Strepsipterologicae–XX). Esakia 28: 11–48. Kremen, C., R.K. Colwell, T.L. Erwin, D.D. Murphy, R.F. Noss, and M.A. Sanjayan. 1993. Terrestrial arthropod assemblages: their use in conservation planning. Conservation Biology 7: 796–808. Kumar, R. 2001. Insect Pests of Agriculture in Papua New Guinea. Part 1. Principles and Practice, Pests of Tree Crops and Stored Products. Science in New Guinea, Port Moresby, Papua New Guinea. Lieftinck, M.A. 1949. The dragonflies (Odonata) of New Guinea and neighbouring islands. Part VII. Nova Guinea, n.s. 5: 1–271. Lienhard, C., and C. N. Smithers. 2002. Pscoptera (Insecta): World Catalogue and Bibliography. Muse´um d’Histoire Naturelle, Geneva. Lugo, A.E. 1988. Diversity of tropical species: questions that elude answers. Biology International Special Issue 19: 1–37. Mercer, C.W.L. 1997. Sustainable production of insects for food and income by New Guinea villagers. Ecology of Food and Nutrition 36: 151–157. Miller, S.E. 1993. Biodiversity and conservation of the nonmarine invertebrate of Papua New Guinea. Pp. 227–325 in Beehler, B.M. (ed.) Papua New Guinea Conservation Needs Assessment, volume 2. Papua New Guinea Department of Environment and Conservation, Boroko. Miller, S.E. 1996. Biogeography of Pacific insects and other terrestrial invertebrates: a status report. Pp. 463–475 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Miller, S.E. 1998. Taxa group summary: insects/terrestrial invertebrates. Pp. 46–47 and map in Burnett, J.B. (ed.) The Irian Jaya Biodiversity Conservation Priority-Setting Workshop: Final Report. Conservation International, Washington, D.C. Miller, S.E., and J.D. Holloway. 1992. Priorities for conservation research in Papua New Guinea: non-marine invertebrates. Pp. 44–58 in Pearl, M., et al. (eds.) Conservation and Environment in Papua New Guinea: Establishing Research Priorities. Embassy of Papua New Guinea, Washington, DC, and Wildlife Conservation International, Bronx, New York.

................. 16157$

CH36

02-08-07 10:39:38

PS

PAGE 527

.

528 /

Miller, S.E., and L.M. Rogo. 2001. Challenges and opportunities in understanding and utilisation of African insect diversity. Cimbebasia 17:197–218. Morris, M.G. 1986. A butterfly farming and trading agency in Irian Jaya. Unpublished report on consultancy in Indonesia for World Wildlife Fund and USAID. Nair, K.S.S. (ed.). 2000. Insect Pests and Diseases in Indonesian Forests: An Assessment of the Major Threats, Research and Efforts. CIFOR, Bogor. Nakamura, M. 1992. An ecological study of earthworms in Papua New Guinea. Journal of Chuo University 13: 19–33. Nakata, S. 1961. Some notes on the occurrence of Phasmatodea in Oceania. Pacific Insects Monograph 2: 107–121. Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific-Australian Region. W. Junk, Dordrecht. Neboiss, A. 1989. Additions and corrections to the Atlas of Trichoptera of the SW PacificAustralian region. Occasional Papers from the Museum of Victoria 4: 63–67. New, T.R. 2003. The Neuroptera of Malesia. Brill, Leiden. New, T.R., and N.M. Collins. 1991. Swallowtail Butterflies: An Action Plan for Their Conservation. IUCN Gland, Switzerland. Nishida, G.M., and N.L. Evenhuis. 2000. Arthropod pests of conservation significance in the Pacific: a preliminary assessment of selected groups. Pp. 115–142 in Sherley, G. (ed.) Invasive Species in the Pacific: A Technical Review and Draft Regional Strategy. SPREP, Apia, Samoa. Novotny, V., Y. Basset, S.E. Miller, P. Drozd, and L. Cizek. 2002a. Host specialization of leaf-chewing insects in a New Guinea rain forest. Journal of Animal Ecology 71: 400–412. Novotny, V., Y. Basset, S.E. Miller, G.D. Weiblen, B. Bremer, L. Cizek, and P. Drozd. 2002c. Low host specificity of herbivorous insects in a tropical forest. Nature 416: 841–844. Novotny, V., P. Drozd, S.E. Miller, M. Kulfan, M. Janda, Y. Basset, and G.D. Weiblen. 2006. Why are there so many species of herbivorous insects in tropical rainforests? Science 313: 1115–1118. Novotny, V., S.E. Miller, Y. Basset, L. Cizek, K. Darrow, B. Kaupa, J. Kua, and G.D. Weiblen. 2005. An altitudinal comparison of caterpillar (Lepidoptera) assemblages on Ficus trees in Papua New Guinea. Journal of Biogeography 32: 1303–1314. Novotny, V., S.E. Miller, Y. Basset, L. Cizek, P. Drozd, K. Darrow, and J. Leps. 2002b. Predictably simple: communities of caterpillars (Lepidoptera) feeding on rainforest trees in Papua New Guinea. Proceedings of the Royal Society of London B 269: 2337–2344. Novotny, V., S.E. Miller, J. Leps, Y. Basset, D. Bito, M. Janda, J. Hulcr, K. Damas, and G.D. Weiblen. 2004. No tree an island: the plant-caterpillar food web of a secondary rainforest in New Guinea. Ecology Letters 7: 1090–1100. Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill, Leiden. Oppel, S. 2006. Comparison of two Odonata communities from a natural and modified rainforest in Papua New Guinea. International Journal of Odonatology 9 (1): 89–102. Orsak, L.J. 1993. Killing butterflies to save butterflies: a tool for tropical forest conservation in Papua New Guinea. News of the Lepidopterist’s Society 1993 (3): 71–80. Owen, I.L. 2005. Parasitic zoonoses in Papua New Guinea. Journal of Helminthology 79: 1–14. Paclt, J. 1982. On some Solomon Islands, Papua New Guinea and Sarawak Thysanura. Annotationes Zoologicae et Botanicae 151: 1–10.

................. 16157$

CH36

02-08-07 10:39:39

PS

PAGE 528

Insects of Papua / 529 Paclt, J. 1985. Diplura from the Bismarck Archipelago and the Solomon Islands. International Journal of Entomology 27: 208–211. Parsons, M.J. 1992. The butterfly farming and trading industry in the Indo-Australian region and its role in tropical forest conservation. Tropical Lepidoptera 3 (Supplement 1): 1–31. Parsons, M.J. 1998. The Butterflies of Papua New Guinea: Their Systematics and Biology. Academic Press, San Diego. Penny, N.D., and G.W. Byers. 1979. A check-list of the Mecoptera of the world. Acta Amazonica 9: 365–388. Pimm, S.L., and J.L. Gittleman. 1992. Biological diversity: where is it? Science 255: 940. Polhemus, D.A. 1996. Island arcs, and their influence on Indo-Pacific biogeography. Pp. 51–66 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Polhemus, D.A., and J.T. Polhemus. 1998. Assembling New Guinea: 40 million years of island arc accretion as indicated by the distributions of aquatic Heteroptera (Insecta). Pp. 327–340 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Polhemus, D.A., and J.T. Polhemus. 2000. A biodiversity survey of aquatic insects in the Ajkwa River Basin and adjacent areas, Irian Jaya, Indonesia. Tropical Biodiversity 5: 197–216. Ponzetta, M.T., and M.G. Paoletti. 1997. Insects as food of the Irian Jaya populations. Ecology of Food and Nutrition 36: 321–346. Price, R.D. 2003. The Chewing Lice: World Checklist and Biological Overview. Illinois Natural History Survey, Champaign. Raven, P.H. (ed.). 1980. Research Priorities in Tropical Biology. National Academy of Sciences, Washington, D.C. Rentz, D.C.F. 1996. Grasshopper Country: The Abundant Orthopteroid Insects of Australia. University of New South Wales Press, Sydney. Ricklefs, R.E., and K. O’Rourke. 1975. Aspect diversity in moths: a temperate-tropical comparison. Evolution 29: 313–324. Riedel, A. 2001. Revision of the simulans-group of Euops Schoenherr (Coleoptera, Curculionoidea, Attelabidae) from the Papuan region. Deutsche Entomologische Zeitschrift 48: 139–221. Robinson, M.H. 1969. Defenses against visually hunting predators. Evolutionary Biology 3: 225–259. Roisin, Y., and J.M. Pasteels. 1996. The nasute termites (Isoptera: Nasutitermitinae) of Papua New Guinea. Invertebrate Taxonomy 10: 507–616. Rosariyanto, E., H. van Mastrigt, H.S. Innah, and H. Yoteni. 2002. Butterflies of the Yongsu area, Papua, Indonesia. Pp. 61–62, 138–139 in Richards, S.J., and S. Suryadi (eds.) A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. Conservation International, Washington, D.C. Ross, E.S. 1948. The Embioptera of New Guinea. Pan-Pacific Entomologist 24: 97–116. Roth, L.M. 2003. Systematics and phylogeny of cockroaches (Dictyoptera: Blattaria). Oriental Insects 37: 1–186. Scheffrahn, R.H., J. Krecek, A.L. Szalanski, J.W. Austin, and Y. Roisin. 2005. Synonymy of two arboreal termites (Isoptera: Termitidae: Nasutitermitinae): Nasutitermes corniger from the Neotropics and N. polygynus from New Guinea. Florida Entomologist 88: 28–33.

................. 16157$

CH36

02-08-07 10:39:39

PS

PAGE 529

.

530 /

Scott, J.M., B. Csuti, J.J. Jacobi, and J.E. Estes. 1987. Species richness: a geographic approach to protecting future biological diversity. BioScience 37: 782–788. Segers, H., and L. De Meester. 1994. Rotifera of Papua New Guinea, with the description of a new Scaridium Ehrenberg, 1830. Archiv fu¨r Hydrobiologie 131: 111–125. Sekhran, N., and S.E. Miller (eds.). 1996. Papua New Guinea Country Study on Biological Diversity. Papua New Guinea Department of Environment and Conservation, Waigani. Simbolon, K., and A. Iswari. 1990. Protected Butterflies in Indonesia. Ministry of Forestry, Republic of Indonesia, Jakarta. Simon Thomas, R.T. 1962. De Plagen van enkele Cultuurgewassen in West Nieuw Guinea. [Checklist of pests of some crops in West Irian]. Meded. Econ. Zak. (Landbouwk. ser.) 1: 1–126. Sluys, R., and I.R. Ball. 1990. The aquatic triclads (Platyhelminthes, Tricladida) of the Bismarck Archipelago. Steenstrupia 16: 13–20. Smithers, C.N., and I.W.B. Thornton. 1981. The role of New Guinea in the evolution and biogeography of Psocopteran insects. Pp. 621–638 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Snelling, R.R. 1998. Social Hymenoptera. Pp. 39–47, 131–143 in Mack, A.L. (ed.) A Biological Assessment of the Lakekamu Basin, Papua New Guinea. Conservation International, Washington, DC. Snyder, T.E. 1949. Catalog of the termites of the world. Smithsonian Miscellaneous Collection 112: 1–490. Solem, A. 1984. A world model of land snail diversity and abundance. Pp. 6–22 in Solem, A., and A.C. van Bruggen, World-wide Snails: Biogeographical Studies on Non-marine Mollusca. E.J. Brill, Leiden. Steinmann, H. 1989. World Catalogue of Dermaptera. Kluwer Academic, Dordrecht. Stonedahl, G.M., and W.R. Dolling. 1991. Heteroptera identification: a reference guide, with special emphasis on economic groups. Journal of Natural History 25: 1027–1066. Sturm, H. 1999. Felsenspringer (Meinertellidae, Archaeognatha, Insecta) aus PapuaNeuguinea und Irian Jaya—Beschreibung einer neuen Gattung und drei neuer Arten [Bristle-tails (Meinertellidae, Archaeognatha, Insecta) from Papua New Guinea and Irian Jaya—description of a new genus and three new species]. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 13 (159): 1–12. Sturm, H., and R. Machida. 2001. Archaeognatha. Walter de Gruyter, Berlin. Sutton, S.L., and N.M. Collins. 1991. Insects and tropical forest conservation. Pp. 405–424 in Collins, N.M., and J.A. Thomas (eds.) The Conservation of Insects and Their Habitats. Academic Press, London. Takaoka, H. 2003. The black flies (Diptera: Simuliidae) of Sulawesi, Maluku and Irian Jaya. Kyushu University Press, Fukuoka, Japan. Tuxen, S.L. 1964. The Protura, a Revision of the Species of the World with Keys for Determination. Hermann, Paris. Ubaidillah, R. 1991. Hama Penting Pada Beberapa Lahan Pertanian di Wamena Jayawijaya Irian Jaya [The important pests of the agricultural area in Wamena, Jayawijaya, Irian Jaya]. Pp. 25–30 in Prosiding Seminar Hasil Penelitian dan Pengembangan Sumber Daya Hayati. Lembaga Ilmu Pengetahuan Indonesia (LIPI), Bogor. Ubaidillah, R., W.A. Noerdjito, and D. Peggie. 1994. Papilionidae Irian Jaya dan pulau disekitarnya [Papilionidae of Irian Jaya and adjacent islands]. Pp. 463–473 in Pratignjo, S.E., W.R. Farida, and Sunaryo (eds.) Prosiding Seminar Hasil Penelitian dan Pengembangan Sumber Daya Hayati, Puslitbang Biologi-LIPI, Bogor, 4 April 1994. Lembaga Ilmu Pengetahuan Indonesia (LIPI), Bogor.

................. 16157$

CH36

02-08-07 10:39:40

PS

PAGE 530

Insects of Papua / 531 van der Lande, V.M. 1993. Onychophora in New Guinea: a review. Science in New Guinea 19: 3–10. van der Lande, V.M. 1994. Haemadipsid leeches of New Guinea: a review of their biology and a guide to identification. Science in New Guinea 20: 9–22. van Herwaarden, H.C.M. 1998. A guide to genera of stick- and leaf-insects (Insecta: Phasmida) of New Guinea and the surrounding islands. Science in New Guinea 24: 55–114. van Mastrigt, H.J.G., and E. Rosariyanto. 2002. Butterflies and moths of the Dabra area, Mamberamo River Basin, Papua, Indonesia. Pp. 63–66, 140–143 in Richards, S.J., and S. Suryadi (eds.) A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. Conservation International, Washington, DC. van Mastrigt, H.J.G., and E. Rosariyanto. 2005. Buko panduan lapangan kupu-kupu untuk wilayah Mamberamo Sampai Pegunungan Cyclops. Conservation International Indonesia Program, Jakarta. van Steenis-Kruseman, M.J. 1950. Cyclopaedia of collectors. Flora Malesiana ser. I, 1: i–clii, 1–639 (supplements l.c. 5: ccxxxiv–cccxlii, 1958, and 8: 1–115, 1974). Willemse, L.P.M. 2001. Guide to the Pest Orthoptera of the Indo-Malayan Region. Backhuys Publishers, Leiden. Williams, D.F. (ed.). 1994. Exotic Ants: Biology, Impact, and Control of Introduced Species. Westview Press, Boulder, Colorado. Yeates, D.K., M.S. Harvey, and A.D. Austin. 2003. New estimates for terrestrial arthropod species-richness in Australia. Records of the South Australian Museum Monograph Series 7: 231–241. Yoshii, R., and Y.R. Suhardjno. 1992. Notes on the Collembolan fauna of Indonesia and its vicinities. II: Collembola of Irian Jaya and Maluku Islands. Acta Zoologica Asiae Orientalis 2: 1–52.

................. 16157$

CH36

02-08-07 10:39:40

PS

PAGE 531

4.4. Cicada Endemism in Papua . , loudly singing cicadas (Hemiptera, Cicadoidea) are well represented in New Guinea. New Guinea cicadas are classified in four tribes: Chlorocystini, Dundubiini (subtribe Cosmopsaltriina), Cicadettini, and Prasiini. The species of the first three tribes have been revised in the last forty years, so their taxonomy and distribution is well documented. The phylogenetic relationships among genera and species were analyzed for Chlorocystini (de Boer 1995b) and Cosmopsaltriina (Duffels and Turner 2002). The New Guinea Prasiini are in need of further revision. At present 155 species are described from New Guinea and surrounding islands (de Boer and Duffels 1997; de Boer 1999, 2000).

T

Cicada Endemism New Guinea cicadas show a high rate of endemism, and not a single species is distributed throughout the island. Most species are restricted to a part of New Guinea or to one or more nearby islands (de Boer 1995a; de Boer and Duffels 1997). There are two possible explanations for this cicada endemism. The first lies in the biology of these insects. Cicadas spend the greater part of their lives as larvae underground, where they feed on the roots of plants. These larvae probably remain within the root system of a single plant, and certainly do not cover large distances. Individuals live in the winged adult stage for at most a couple of weeks. This means that only a short part of the cicada lifecycle is available for dispersal. Distribution patterns, however, suggest that even the winged insect does not disperse very far. Another explanation for the high endemicity lies in the complex geotectonic history of New Guinea (Holloway and Hall 1998). Southern New Guinea, the Fly River platform, is an integral part of the Australian continental plate, but the rest of New Guinea consists of more than thirty allochtonous terranes, of both continental and island arc origin, which have accreted to the Australian plate margin during the last 25 million years. The isolation of these terranes has apparently greatly favored speciation and endemism in cicadas.

Plotting the distributions of the individual New Guinean cicada species on the map, it appears that many of these distributions coincide. From this, five areas of endemism can be identified (Figure 4.4.1). Four of these areas of endemism correspond to groups of island arc terranes that successively accreted to the Australian continental plate: the central mountain ranges (25 million years ago), the Papuan Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

532

................. 16157$

CH37

02-08-07 10:39:56

PS

PAGE 532

Cicada Endemism in Papua / 533

Figure 4.4.1. Areas of endemism (circled) in the west Pacific. Papua is subdivided in four areas of endemism. peninsula (15 mya), the northern mountain ranges (10 mya), and the Vogelkop peninsula (also 10 mya). Each of the cicada genera is more or less concentrated in one of the areas of endemism, but none of them is strictly endemic to that area. This can be explained by supposing that the New Guinean cicada genera evolved in isolation prior to the accretion event, while after accretion some species dispersed into adjacent areas. The fifth area of endemism, south New Guinea, is part of the Australian continental plate, but not a single Australian species occurs there. South New Guinea shows some endemism at the species level only; the endemics belong to genera that are centered elsewhere in New Guinea.

Papua can be subdivided into four main areas of endemism, each with its own distinct biota: the Vogelkop peninsula, the northern mountain ranges, the central mountain ranges, and south New Guinea. The latter three continue into Papua New Guinea. A subdivision of Papua into four areas of endemism is rather coarse, but some remarks on more local endemicity will be made.

The Vogelkop Area The Vogelkop has a highly characteristic cicada fauna and can be recognized as the most distinctive area of endemism in New Guinea. This area comprises the Vogelkop proper and the nearby islands of Misool, Salawati, Waigeo, and Roon. No less than three genera are centered in this area: Arfaka, Rhadinopyga, and Aedeastria. Arfaka contains three species, which are all endemic to the Vogelkop. Of the genus Rhadinopyga (four described and four undescribed species), seven are endemic in the area and one occurs in the Vogelkop and on Bacan Island (northern Maluku). The genus Aedeastria (12 species) has three endemic species in the Vogelkop, while two species are found there and elsewhere in New Guinea;

................. 16157$

CH37

02-08-07 10:40:14

PS

PAGE 533

#

534 /

.

the genus as a whole has a predominantly western New Guinea-north Maluku distribution. In total 37 cicada species (of 13 genera) occur in the Vogelkop area, 19 species (51%; of eight genera) are endemic there. Most of the Vogelkop endemics are restricted to one of the islands: Misool (Arfaka: 1 species, Thaumastopsaltria: 1), Salawati (Rhadinopyga: 1), Waigeo (Aedeastria: 1 and Rhadinopyga: 2), while one Aedeastria species is endemic to Waigeo and Misool. Most peculiar is the endemism on the small island of Roon, which lies just north of the Wandammen peninsula in the Cenderawasih Bay and is only separated from New Guinea proper by a narrow stretch of water. Two species of Baeturia and two undescribed species of Rhadinopyga are endemic to Roon, while a species of Aedeastria has a strongly deviating form on the island. The Vogelkop proper has six endemics (Aedeastria: 1; Lembeja: 1; Rhadinopyga: 3; Toxopeusella: 1), but this area is certainly undercollected. This high rate of endemism suggests a long history of isolation for the Vogelkop terranes. The Vogelkop is of composite geological origin: it comprises microcontinents as well as island arc fragments. This composite nature is reflected in the phylogenetic relationships of the genera centered in the Vogelkop. Aedeastria and Rhadinopyga indicate relationships of the Vogelkop to the Papuan Peninsula and the Solomon Islands respectively, while Arfaka possibly has its nearest relatives in Sulawesi.

Central Mountain Ranges Two groups of cicadas, the large mimica complex of the genus Cosmopsaltria (17 species) and the nasuta group of the genus Baeturia (12 species) show a main representation in the central mountain ranges and characterize the cicada fauna of this area. The ranges of several species of these groups extend into the Papuan peninsula and some also occur in other adjacent areas. Altogether 48 cicada species belonging to ten genera have been recorded from the central mountain ranges (Figures 4.4.2,3). The cicada fauna of the area has a fairly high rate of endemism; 22 species (46%) are found only there. Most endemics of the central mountain ranges have very restricted distributions; only two endemics are widespread and recorded from Papua and Papua New Guinea. The Papua part has 34 species belonging to eight genera. A total of 11 species (32%) are endemic here. Most of these endemics (9 species) have been recorded from the northern slopes of the central mountains, south of the Idenburg ( Taritatu) River, in an area centered around 139 00 N and 3 30 S. The area was visited by Dr L. J. Toxopeus during the Third Archbold Expedition in 1939. A very interesting collection of cicadas was made in a wide range of habitats, and at altitudes ranging from the foothills of the central mountains (Bernhard Camp: 50 m) to Araucaria Camp (700–900 m), Rattan Camp (1,200 m), the moss forest of Mist Camp, and Top Camp (1,800 m and 2,100 m, respectively), and even as high as Lake Habbema (3,250 m). Since a total of 23 cicada species (68% of the known fauna of the central mountain ranges of Papua), including nine endemics, have been recorded from these sites, the area can be regarded as a biodiversity hotspot for cicadas. It

................. 16157$

CH37

02-08-07 10:40:15

PS

PAGE 534

Cicada Endemism in Papua / 535

Figure 4.4.2. Baeturia hamiltoni de Boer of the tribe Chlorocystini, an endemic species of Central New Guinea. Left: male. Right: female. Body length male: 28 mm. Drawings: Dick Langerak, from de Boer (1994).

................. 16157$

CH37

02-08-07 10:41:26

PS

PAGE 535

536 /

#

.

Figure 4.4.3. Cosmopsaltria papuensis Duffels of the subtribe Cosmopsaltriina, an endemic species of Central New Guinea. The populations of this species differ remarkably in marking on head and thorax. Left: male from Star Range. Top right: male from Baliem Camp. Bottom right: male from Araucaria Camp. Body length (left): 41 mm. Drawings: J. Zaagman, from Duffels (1983).

................. 16157$

CH37

02-08-07 10:42:17

PS

PAGE 536

Cicada Endemism in Papua / 537

is most likely that this really is a hotspot, and not just the result of very intensive collecting, because the Star Mts for instance, which were visited by the Netherlands New Guinea Expedition in 1959, have a fairly high number of recorded species (17) but no endemics at all. Two other areas at the western end of the central mountain ranges should be mentioned: the Wissel Lakes area and the nearby Mappia area. Each of these areas has one endemic cicada species.

Northern Mountain Ranges The cicada fauna of northern New Guinea is fairly diverse; the 74 species occurring there belong to 14 genera. Altogether 21 species (28%) are endemic in this area. Furthermore, there are at least some undescribed endemic Lembeja species. Many north New Guinea cicadas have a comparatively wide distribution all along the northern mountain ranges to well into the Papuan peninsula. These distributions sometimes include the islands of Biak, Yapen, and Numfoor, which are therefore included in the north New Guinea area of endemism. The genera Guineapsaltria and Mirabilopsaltria are more or less centered in the north. Furthermore the speciose genus Baeturia has many species in this area and several of its species groups are centered there. A monophyletic subgroup of the Baeturia viridis group consists of four species occurring in successive areas along the northern mountain ranges. The Papua part of northern New Guinea has 50 species belonging to 14 genera. A total of eight species (16%) is endemic here; these endemics belong to six genera. One species of Baeturia is widespread in the area. Another species of this genus and one species of Scottotympana are recorded from the lowland between Cape d’Urville and Jayapura. Furthermore, one species of Cosmopsaltria is recorded from the Geelvink (now Cenderawasih) Bay area, and a species of Mirabilopsaltria is endemic to the surroundings of Jayapura. The islands of Biak and Numfoor each have an undescribed endemic Diceropyga species. Yapen Island has no endemic cicadas.

South New Guinea None of the genera is centered in the southern lowlands of New Guinea. South New Guinea is the area of endemism with the poorest cicada fauna: 23 species occur here and seven (30%) are endemic. The Papua part has two endemic (sister) species, that belong to the genus Baeturia. In total 14 species belonging to no less than nine genera occur in southern Papua. The relatively high number of genera relative to the low number of species is most likely due to several independent invasions. The islands of Kai have two endemic cicadas, a Cosmopsaltria and an Aedeastria species, but the nearby Aru Islands have no cicada endemics.

Literature Cited de Boer, A.J. 1994. The taxonomy and biogeography of the loriae group of the genus Baeturia Sta˚l, 1866 (Homoptera, Tibicinidae). Tijdschrift voor Entomologie 137: 1–26. de Boer, A.J. 1995a. Islands and cicadas adrift in the West-Pacific, biogeographic patterns related to plate tectonics. Tijdschrift voor Entomologie 138: 169–244.

................. 16157$

CH37

02-08-07 10:42:17

PS

PAGE 537

538 /

#

.

de Boer, A.J. 1995b. The phylogeny and taxonomic status of the Chlorocystini (sensu stricto) (Homoptera, Tibicinidae). Contributions to Zoology 65: 201–231. de Boer, A.J. 1999. Taxonomy and biogeography of the New Guinean Cicadettini (Hemiptera, Tibicinidae). Deutsche Entomologische Zeitschrift 46: 115–147. de Boer, A.J. 2000. The cicadas of Mt. Bosavi and the Kikori Basin, southern Papua New Guinea. Bishop Museum Occasional Papers 61: 1–23. de Boer, A.J., and J.P. Duffels. 1997. Biogeography of Indo-Pacific cicadas east of Wallace’s Line. Pp. 297–330 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Duffels, J.P. 1983.Taxonomy, phylogeny and biogeography of the genus Cosmopsaltria, with remarks on the historic biogeography of the subtribe Cosmopsaltriaria (Homoptera: Cicadidae). Pacific Insects Monograph 39: 1–127. Duffels, J.P., and H. Turner. 2002. Cladistic analysis and biogeography of the cicadas of the Indo-Pacific subtribe Cosmopsaltriina. Systematic Entomology 27: 235–261. Holloway, J.D., and R. Hall. 1998. SE Asian geology and biogeography: an introduction. Pp. 1–23 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden.

................. 16157$

CH37

02-08-07 10:42:18

PS

PAGE 538

4.5. Marine Wood-Boring Invertebrates of New Guinea and Its Surrounding Waters . of a complex of crystalline arrays of cellulose, embedded in hemicelluloses and lignin (lignocellulose), which is a rich source of potential energy, but a rather refractory substrate. Only a limited range of microorganisms and invertebrates can exploit this resource. On land, wood breakdown is achieved mainly by beetles, termites, basidiomycete fungi, and, at a rather slower rate, by cellulolytic bacteria. Where regular tidal inundation excludes such organisms, another suite of organisms takes over. Here, bacterial degradation still occurs, but the fungi are mainly ascomycetes; the major degradation, however, is due to wood-boring crustaceans and bivalves. These borers belong to the isopod families Sphaeromatidae and Limnoriidae, the amphipod family Cheluridae, and the bivalve families Pholadidae and Teredinidae. On the coasts of New Guinea and the waters surrounding it, these organisms inhabit natural and anthropogenic niches. They readily colonize the large quantities of both living and dead wood in the intertidal zone on coasts with a low wave energy regime where mangrove forests thrive. The quantities of dead wood brought down to the sea by large rivers also provide a suitable substrate, either where retained at the coastline, or when carried out to sea. Driftwood also arrives on ocean currents and waterlogging leads to deposition of quantities of wood on the seabed, particularly off the mouths of large rivers. These organisms can cause extremely rapid degradation of wood. For example, a survey of the piers constructed around New Guinea during the Pacific War revealed that at many of the sites surveyed, these borers brought piers to the point of structural failure within eighteen months (Shillinglaw and Moore 1947). Furthermore, planks of the dense timber Hopea exposed at sites along the north coast of Papua New Guinea lost up to two-thirds of their weight within four months due to teredinid activity (Cragg, pers. obs.). This chapter will consider the taxonomy of these borers, their biogeography, the factors that control their distribution, their ecological roles, and the economic implications of their activities. The goals of this chapter are to summarize and make accessible information from a wide variety of sources, many of which are not readily accessible to the wider scientific readership. While more information is available from Papua New Guinea, there is sufficient evidence of similar patterns of borer colonization and activity in Papua for the findings to be applicable around the whole island.

W

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

539

................. 16157$

CH38

02-08-07 10:39:58

PS

PAGE 539

.

540 /

Borer Species from New Guinea and Adjacent Waters Collections of wood borers from the New Guinea area have been reported by a considerable number of authors (Tables 4.5.1,2). The marine wood-boring fauna of New Guinea includes crustaceans of all of the main families of borers mentioned above. There are also other wood-inhabiting isopods from the families Corallanidae and Cirolanidae that may or may not form their own burrows. The teredinids present a particular problem for taxonomists because, unlike most bivalves, the shell shows more variation within species than between species (Turner 1966). Species identification within this family relies on the characteristics of the stalked calcified paddle-shaped structures (pallets) that serve as retractable stoppers to the entrance of the animal’s tunnel (Figure 4.5.1). Because these structures are affected by the age of the animal, the nature of the tunnelled wood, and by local water chemistry, there is much scope for creation of synonyms. In her comprehensive review of the family, that involved re-examination of much type material, Turner (1966) clarified the nomenclature by reducing many species names to synonyms. For example, the following species given by Roch (1961) as occurring in and around New Guinea are recorded in Turner’s catalog as synonyms of the species shown in parentheses: Teredo arenaria (Kuphus polythalamia), Teredo bataviana (Spathoteredo obtusa), Teredo malaccana (Lyrodus pedicellatus), Teredo milleri (Lyrodus affinis), Bankia penna-anseris (Nausitora dunlopei), Teredo hermitensis (T. clappi). Turner (1971) provided a key to the family based almost exclusively on pallet characteristics, though species pairs with similar pallet morphology have now been recognized that can be distinguished by characteristics of the larval brooding strategy (Turner and Calloway 1987). The species list in Table 4.5.1 is based on Turner’s catalog and key. However, Rayner (1983) reported that Nausitora globosa came to be considered by Turner to be a separate species, rather than a synonym of N. dunlopei, and Lyrodus tristi to be no longer considered a synonym of L. pedicellatus. Turner and Rayner recognized two unnamed teredinid species in the collections from New Guinea, one from the genus Nausitora and the other from Lyrodus, but unfortunately these species have not been formally described. Specimens of a teredinid were found in seagrass roots at Motupore Island near Port Moresby. They had typical teredinid shells and calcified tunnel linings, and females carried broods of larvae developed beyond the straight hinge stage. The pallets were pale yellow and resembled those of Zachsia zentkewitschi (Cragg, pers. obs.). As this species was found on the Pacific coast of Russia, it is unlikely that the specimens that were found in the tropical waters of Motupore Island belong to the same species. The taxonomy of the Limnoriidae was put on a firm footing by Menzies (1957), who included descriptions of species that occur around New Guinea, and Cookson (1991), who brought subsequent descriptions of new species into an overhauled taxonomic structure in which Menzies’ subgenera were no longer supported by

................. 16157$

CH38

02-08-07 10:39:59

PS

PAGE 540

x x x x x

Bankia bipalmulata (Lamarck)

Bankia bipennata (Turton)

Bankia campanellata Moll & Roch

Bankia carinata (Gray)

Bankia gracilis Moll

x

Bankia barthelowi Bartsch

................. 16157$

CH38

02-08-07 10:39:59

PS

x

x

x

x

x

Bankia australis (Calman)

(x)

x

driftwood (x)

x

x

x

x

x

(x)

x

sunken wood Nypa roots

Substrate

mangrove

Bactronophorus thoracites (x) (Gould)

Teredinidae

Species

timber

marine and brackish

marine

marine

marine

marine

marine

marine

marine and brackish

Salinity1,2

Table 4.5.1. Wood-boring bivalves in waters around New Guinea

Bismarck, Solomon, and Coral seas2

Gulf of Papua1; Bismarck, Solomon, and Coral seas2

Gulf of Papua1; Daru2

Gulf of Papua1; Ponam Island, Rabaul2

Manokwari3; Bismarck and Solomon seas, Kupiano2

Bismarck and Solomon seas2

Bismarck and Solomon seas2

Merauke3; Gulf of Papua1; Bismarck, Solomon, and Coral seas2

Sites

(continued)

Indo-Pacific tropical and subtropical

Circumtropical and subtropical

Circumtropical

Circumtropical and subtropical

Indo-Pacific tropical and subtropical

Philippines

Australia, New Zealand

Australia, Southeast Asia, India

Distribution6

Marine Wood-Boring Invertebrates of New Guinea / 541

PAGE 541

................. 16157$

CH38

02-08-07 10:39:59

PS

x

(x)

marine

Bismarck, Solomon, and Coral seas2

x

euryhaline marine

Lyrodus massa (Lamy)

(x)

Bismarck, Solomon, and Coral seas2

x

x

Jayapura, Fakfak, Merauke, Muna, Seegarbin3; Bismarck, Solomon, and Coral seas2

Lyrodus bipartitus (Jeffreys)

marine and brackish

Merauke3; Rabaul, Coral Sea2

Bismarck, Solomon, and Coral seas2

Fakfak3; Rabaul, Lae2

Bismarck, Solomon, and Coral seas2

New Guinea3

(x)

marine and brackish

marine

marine

marine

Sites

Lyrodus affinis (Deshayes)

x

(x)

(x)

(x)

Salinity1,2

Batjan3

x

Dicyathifer manni (Wright)

x

(x)

mangrove

Kuphus polythalmia (Linnaeus)

x

Bankia rochi Moll

x

x

Bankia nordi Moll

Bankia philippinensis Bartsch

x

timber

Bankia johnsoni Bartsch

Species

driftwood

Substrate

sunken wood Nypa roots

Table 4.5.1. (Continued)

Circumtropical and subtropical

Circumtropical and subtropical

Indo-Pacific tropical and subtropical

Philippines, Sumatra, Solomon Islands

Indo-Pacific tropical and subtropical

Indo-Pacific tropical and subtropical

Philippines, Malaysia

Indo-Pacific tropical and subtropical

Philippines

Distribution6 542 /

.

PAGE 542

(x) x x x x x x

Nausitora sp. nov.

Nototeredo edax (Hedley)

Spathoteredo obtusa (Sivickis)

Teredo clappi Bartsch

Teredo fulleri Clapp

Teredo furcifera von Martens

Teredo johnsoni Clapp

x

Nausitora dunlopei Wright

x

x

Lyrodus sp. nov.

Nausitora hedleyi Schepman

x

Lyrodus tristi (Ireldale)

x

................. 16157$

Nausitora globosa (Sivikis)

x

Lyrodus pedicellatus (Quartrefages)

CH38

02-08-07 10:40:00

PS

x

x

x

x

x

x

x

x

x

(x)

x

(x)

x

(x)

x

(x)

(x)

x

(x)

x

marine

euryhaline marine

marine

marine and brackish

marine

euryhaline marine

marine

brackish

brackish

brackish

marine

marine

euryhaline marine

2

Bismarck, Solomon, and Coral seas2

Fakfak3

Bismarck and Solomon seas, Port Moresby2

Bismarck, Solomon, and Coral seas2

Fakfak, Ralum-New Britain3; Solomon and Bismarck seas2

Bismarck, Solomon, and Coral seas

Wewak, Lae2

Merauke4; New Britain3; Bismarck, Solomon, and Coral seas2

Bismarck and Coral seas, Lae2

Bismarck, Solomon, and Coral seas 2

Bismarck, Solomon, and Coral seas2

Oro Bay, Madang, Vanimo2

Bismarck, Solomon, and Coral seas2

(continued)

Circumtropical and subtropical

Circumtropical and subtropical

Circumtropical and subtropical

Circumtropical and subtropical

Indo-Pacific tropical

Indo-Pacific tropical to warm temperate

No other reports

Indo-Pacific tropical

Indo-Pacific tropical to warm temperate

No other reports

Australia

Cosmopolitan, temperate to tropical

Marine Wood-Boring Invertebrates of New Guinea / 543

PAGE 543

................. 16157$

CH38

02-08-07 10:40:01

x

(x)

Teredora princesae (Sivikis)

Teredothyra excavata (Jeffreys)

x

Teredo somersi Clapp

x

x

Teredo poculifer Ireldale

Teredothyra dominicensis (Bartsch)

x

timber

Teredo mindanensis Bartsch

Species

mangrove x

(x)

x

(x)

driftwood

Substrate

x

sunken wood Nypa roots

Table 4.5.1. (Continued)

PS

marine

marine

marine

marine

brackish

marine

Salinity1,2

Rabaul, Lae2

Rabaul, Lae, Alotau2

Madang, Ponam Island2

Bismarck and Solomon seas, Port Moresby2

Marshall Lagoon2

Gulf of Papua1; Bismarck, Solomon, and Coral seas2

Sites

Circumtropical and subtropical

Western Atlantic tropical to subtropical

Indo-Pacific tropical to temperate

Western Atlantic tropical to subtropical

Queensland (Australia)

Java, tropical Australia

Distribution6

544 /

.

PAGE 544

................. 16157$

CH38

x

02-08-07 10:40:01

PS

x

Martesia striata (Linnaeus)

marine and brackish

marine and brackish Gulf of Papua1

Gulf of Papua1

Gulf of Papua1

Gulf of Papua1

Port Moresby5

Ponam Island3

Port Moresby3

Rabaul, Port Moresby2

Jayapura3; Gulf of Papua1; Rabaul, Lae, Marshall Lagoon2

Cosmopolitan, temperate to tropical

West Atlantic, New Jersey to Brazil

India, Borneo

No other reports

Indo-Pacific tropical

Indo-Pacific tropical

Indo-Pacific tropical and subtropical

Circumtropical and subtropical

Notes: ‘‘x’’ indicates species present in substrate; ‘‘(x)’’ indicates species is rare in substrate. Species collected at sites around the coast of Papua New Guinea are listed by sea coast of collection (Coral, Solomon, or Bismarck seas). Sources: 1Cragg and Aruga 1988; 2Rayner 1983; 3Roch 1961; 4Schepman 1919; 5see text; 6Turner 1971.

x

Martesia cuneiformis (Say)

brackish

Lignopholas rivicola (Sowerby)

marine

marine

marine

marine

marine

brackish x

x

Barnea obturamentum Hedley

Pholadidae

Zachsia sp.

(x)

Uperotus rehderi (Nair)

(x)

(x)

(x)

(x)

Teredothyra smithi (Bartsch)

x

Uperotus clavus (Gmelin)

x

Teredothyra matocotana (Bartsch)

Marine Wood-Boring Invertebrates of New Guinea / 545

PAGE 545

................. 16157$

CH38

02-08-07 10:40:01

PS

x

Limnoria pfefferi Stebbing

x

x

x

Limnoria multipunctata Menzies

x

x x

x

Limnoria kautensis Cookson & Cragg

Limnoria insulae Menzies

x

Limnoria indica Becker & Kampf

x

x x

x

Limnoria foveolata Menzies

Limnoria andamanensis Rao & Ganapati

Limnoriidae

Isopoda

Species

mangrove timber driftwood sunken wood

Wood type

marine

marine

marine

marine

Salinity

Port Moresby, Rabaul, Madang7

Kai Islands,1 Milne Bay7

Kaut6

Port Moresby, Motupore Island, Alotau7

Lorengau7

Kai Islands, Vogelkop2

Buka Passage, Madang, Rabaul, Alotau7

Site

Table 4.5.2. Burrow-inhabiting isopods from wood in waters around New Guinea

Puerto Rico, Belize, Panama, Florida, Andaman Islands, Aldabra, Philippines, Australia7

Saipan, Canton Island, Guam,1 Puerto Rico, Jamaica, Japan,2 Belize,18 Australia7

No other reports

Fiji, Guam, Palmyra Island, Ponape, Andaman Islands, Kenya, Australia7

Madras, Hong Kong, Manila, north Queensland, Andaman Islands, Japan, Malaysia, Belize7

Belize, Andaman Islands7

Distribution 546 /

.

PAGE 546

x

x

Limnoria tripunctata Menzies

Limnoria unicornis Menzies

................. 16157$

CH38

02-08-07 10:40:02

PS

x

Corallana estuaria Jones et al.

Corallana glabra Nierstrasz

x

Corallana bidentata Jones et al.

Corallanidae x

marine

brackish

euryhaline

Port Moresby3; Gulf of Papua5

Port Moresby3; Gulf of Papua5

Kaut6

Kai Islands10

Galley Reach, Waipara River, Sewa Bay (Milne Bay)4

Port Moresby4; Gulf of Papua5

Central Province9

x

euryhaline

euryhaline

marine

Port Moresby, Madang, Lorengau7

Wewak, Lae7

Manus Island7

Admiralty Islands2

Pistorius bidens Harrison & Holdich

x

x

x

Central Province9

x

Sphaeroma triste Heller

x

x

x

x

x

Sphaeroma intermedium (Baker)

x

Sphaeroma terebrans Bate

Sphaeromatidae

Paralimnoria asterosa Cookson & Cragg

Paralimnoria andrewsi (Calman)

x

Limnoria platycauda Menzies

Murray River (Queensland)

(continued)

Philippines, India, Sri Lanka, Australia15

Heron Island (Australia)14

North Australia12

India to Australia,12 Manila13

Circumtropical12

No other reports

Ghana, Puerto Rico, Japan, Hawaii, Samoa, Florida, Philippines, Cocos Islands7

Ponape1; Caroline Is., Andaman Is., San Salvador, Belize, N. Australia7

Uruguay, Colombia, Panama, Puerto Rico, Florida, Turkey, Hong Kong, Ghana, Japan2

Puerto Rico, Belize, Cuba, Andaman Islands, Aldabra, India, Japan, Thailand7

Marine Wood-Boring Invertebrates of New Guinea / 547

PAGE 547

................. 16157$

CH38

02-08-07 10:40:03

PS

10

Gulf of Papua, Milne Bay, Bougainville Island8; New Ireland5

Gonema River11; Galley Reach9; Gulf of Papua5

Labu Lakes, Lae10

Gulf of Papua5

Port Moresby, Manus Island4

Port Moresby, Milne Bay, Oro Bay4; Gulf of Papua5

Aru Islands

Site

Caribbean, tropical Pacific islands, Japan, Thailand2

Malaysia17

East Africa to Queensland, (Australia)13

Queensland, Sri Lanka, east Africa, Thailand16

No other reports

India, Sri Lanka, Australia, Philippines15

Distribution

Source: 1Menzies 1957; 2Kuhne 1976; 3Cragg and Levy 1978; 4Jones et al. 1982; 5Cragg and Aruga 1988; 6Cookson and Cragg 1988; 7Cookson 1991, 8Rayner 1976; 9see text; 10 Bruce 1982b; 11Bowman 1977; 12Harrison and Holdich 1984; 13Holdich et al. 1981; 14Harrison and Holdich 1982; 15Bruce 1982a; 16Bruce 1986; 17Bruce 1993; 18Kensley and Schotte 1987.

Chelura insulae Calman

Cheluridae

Amphipoda

x

Ceratolana papuae Bowman

brackish

brackish

x

Anopsilana pustulosa (Hale)

marine brackish

x

Anopsilana willeyi (Stebbing)

Cirolanidae

Corallana tridentata Jones et al.

euryhaline

x

Corallana nodosa Schioedte & Meinert

x

marine

Salinity

Corallana leopoldi (Nierstrasz)

Species

Wood type

mangrove timber driftwood sunken wood

Table 4.5.2. (Continued)

548 /

.

PAGE 548

Marine Wood-Boring Invertebrates of New Guinea / 549

Figure 4.5.1. The teredinid Dicyathifer manni with shell above and partially retracted siphons between calcareous pallets below. Photograph courtesy of Suzanne Rayner.

the anatomical evidence. The key developed from Menzies’ descriptions was superseded by the more comprehensive key of Cookson (1991). The species Limnoria foveolata is based on a single collection with only female specimens. It may prove to be a synonym of either L. indica or L. saseboensis (Cookson, 1997). The anatomy of the Sphaeroma species reported here has been detailed by Harrison and Holdich (1984); uropod features (Figure 4.5.2) and telson sculpturing are diagnostic features. The sphaeromatid isopod Pistorius bidens was originally recorded from under slabs of intertidal rock in Heron Island (Harrison and Holdich 1982b). The original description of this species was based on ethanol-preserved specimens, in which the striking coloration of the live animal had not been retained. They were reported to be cream or red-brown with black chromatophores. Live animals were found in the middle to high intertidal in burrows in a stump of the mangrove of Avicennia, in Central Province, Papua New Guinea (Cragg, pers. obs.). These were found to be bright red above with black chromatophores along the midline of the dorsal surface, but green below. Like other sphaeromatids, these animals inhabit tight fitting burrows, can roll into a tight ball with the telson towards the burrow opening, but are also capable of vigorous swimming. Beetle larvae, tentatively identified as Eobia sp., are common in stumps of Avicennia in the medium to high intertidal of mangrove forests around Port Moresby (Cragg, pers. obs.; Figure 4.5.3). They have heavily sclerotized mandibles (Figure 4.5.4), which are used to excavate galleries that are oval in cross-section. These are

................. 16157$

CH38

02-08-07 10:40:07

PS

PAGE 549

.

550 /

Figure 4.5.2. Dorsal view of Sphaeroma triste showing the characteristically broad, toothed exopods of the uropods.

Figure 4.5.3. Stump of the mangrove Avicennia with openings to the galleries of wood larvae of the beetle Eobia.

................. 16157$

CH38

02-08-07 10:40:14

PS

PAGE 550

Marine Wood-Boring Invertebrates of New Guinea / 551

Figure 4.5.4. Ventral view of the head of a larva of Eobia, showing the heavily sclerotosed mandibles. packed with damp frass or rejectimenta, suggesting that these larvae are subject to tidal inundation, as has been found to be the case for larvae of the intertidal woodboring weevil Pselactus (Sawyer and Cragg 1995).

Wood-boring invertebrates have a wide range of life history strategies. Limnoriids and sphaeromatids often occur in burrows as a male and female pair. Females carry a small number of developing embryos (up to 100 for Sphaeroma, but six or fewer in tropical limnoriids) in a brood pouch formed by flanges (oostegites) developing from the walking legs. Newly released juveniles remain within the parental burrow. Dispersal to new wood is undertaken by late juveniles or adults (Cragg 2003). Teredinids are protandric hermaphrodites that produce very large numbers of eggs 40–60 m in diameter. These are fertilized after broadcast spawning or in the epibranchial chamber (a mantle-lined chamber containing the gills). Zygotes may be released into the sea or, in the case of the genera Teredo and Lyrodus, retained on the gills until the veliger or pediveliger stage (respectively the planktonfeeding and settlement stages of larval development) (Turner and Johnson, 1971). Development in pholads is, like that of non-brooding teredinids, planktotrophic (relying on planktonic food to fuel growth). Planktotrophic larvae can become very widely dispersed, while the dispersal potential of species that brood to the pediveliger stage is much lower. The life histories of the wood-boring isopods show characteristics of K-selection, while those of the bivalves are r-selected. The prediction from this is that the bivalves should be more effective at locating new sources of wood (which is cer-

................. 16157$

CH38

02-08-07 10:40:17

PS

PAGE 551

.

552 /

tainly evidenced by the numbers of teredinid species colonizing test panels within a short period of exposure), but that the isopods should be strong competitors once established. However, the boring pattern of the two borer taxa may permit coexistence, as teredinids tend to burrow deeper than the isopods (Si et al. 2000).

As is common with a wide range of invertebrate taxa in the waters of the Indonesian archipelago, wood-boring invertebrates exhibit high diversity. Of the approximately 30 wood-boring members of the Limnoriidae recognized by Cookson (1991) and in subsequent species descriptions, 12 occur around New Guinea. There are approximately 70 wood-boring teredinids recognized by Turner in her catalog of 1966, or subsequently removed from synonymy by her. Thirty-eight of these have been reported from New Guinea waters, together with two undescribed species of wood borer, an undescribed seagrass borer, and the mud-burrowing Kuphus. Thus, a substantial portion of the diversity in the two main borer families is represented around New Guinea. Furthermore, six of the 15 recognized species of Corallana have been found associated with wood, including four from the waters around New Guinea (Jones et al. 1983). Though some of the species are separated by local environmental gradients, particularly in salinity, many species occur sympatrically. Indeed, pieces of fallen wood or wooden pilings may contain more than one species of borer from the same family. For example, Sphaeroma terebrans and S. triste have been found in the same timber piling (Cragg and Levy 1979) or piece of mangrove, though one species or the other predominates with nearby sites often having the reverse species predominant (Cragg, pers. obs.). Sympatry may be even more marked in the teredinids: wood panels exposed as part of a survey of the factors controlling teredinid colonization often became colonized within three months by up to five and, at Port Moresby, by up to 11, different teredinid species (Cragg, pers. obs.).

The huge island of New Guinea is surrounded by the Solomon, Bismarck, Coral, Arafura, Banda, and Seram Seas, and West Caroline Basin. There is an extensive continental shelf off the southern coast. Seasonal ocean currents move waters westward then eastward between these basins and the adjoining Indian and Pacific oceans. Species with planktotrophic development are likely to broadly dispersed by these currents, so it is not surprising that the extensive study by Rayner (1983) of teredinids in the waters of Papua New Guinea found that most teredinid species were present in all three sea coasts examined (Coral, Bismarck, and Solomon seas). Further collection would probably reveal island-wide distribution in many cases. Indeed, most of these species have Indo-Pacific or even circumtropical distributions (Table 4.5.1). However, even the isopods, with their much smaller capability for dispersal, show broad patterns of distribution (Table 4.5.2), with some having an apparently discontinuous Indo-Pacific plus Caribbean distribution. In inter-

................. 16157$

CH38

02-08-07 10:40:18

PS

PAGE 552

Marine Wood-Boring Invertebrates of New Guinea / 553

preting these distributions, it should noted that borers may disperse in driftwood or, in the past, in hulls of wooden ships. Of the borers with limited known distributions, Paralimnoria asterosa, Limnoria kautensis, and Limnoria foveolata have been found only at depths that are rarely sampled specifically for wood borers, while Zachsia sp. occurs in the rather inaccessible habitat of seagrass roots. The dispersal of Teredo poculifer may be limited by its requirement for brackish water, though it may have been dispersed from estuary to estuary in hulls of dugout canoes (Rayner 1979).

While the limits of distribution of borers are determined by species dispersal capabilities and by tolerance of environmental variables, such as temperature, that vary on a geographic scale, distribution within these limits is determined by local variations in factors such as the availability and type of substrate for boring, salinity, and tidal height or depth in the water column. There is no indication that water temperature is a key factor for determining local distribution, though as temperature decreases with depth, some species, such as the limnoriids L. kautensis, L. foveolata, and P. asterosa, which have only been found at depths below 10 m, may be restricted to below the typical level of the local thermocline. Tidal height may be a critical factor determining distribution of some organisms. Figure 4.5.5 demonstrates the typical vertical distribution of burrows of Sphaeroma triste with respect to tidal level. Virtually all burrows occur in the middle intertidal, where, at low tide, the animals will be subject to desiccation and cannot suspension feed. Perhaps this level of desiccation cannot be readily tolerated by their predators. Limnoria insulae has only been found intertidally (Cookson 1991), but other limnoriids have only been found subtidally (Ku¨hne 1976; Cookson and Cragg 1988) or even at considerable depths (Cookson 1991). Records of deepwater limnoriids are lacking for the New Guinea region, perhaps due to limited collecting in this environment. Table 4.5.3 gives details of teredinids collected at depths of tens of meters in the Gulf of Papua. All species collected have also been found in shallow water around New Guinea (Table 4.5.1). Field observations of borers indicate that salinity is a key determinant of distribution. Some species, indicated as marine in Tables 4.5.1 and 4.5.2, have only been found at sites with little or no freshwater influence. They may be capable of tolerating lowered salinity for short periods, but they do not seem to be capable of reaching breeding condition in freshwater. The absence of the teredinids Bankia bipennata, Teredothyra matocotana, and Teredo mindanensis from the Gulf of Papua delta complex, though they were collected in the more saline waters offshore, indicates that these may be stenohaline marine species (Cragg and Aruga 1988). Rayner (1979) examined the colonization of bait blocks distributed along a well-defined gradient of salinity in an estuary to the east of Port Moresby, and followed up with laboratory experiments with the species found in that estuary. On this evidence, she was able to characterize the teredinids Teredo poculifer as a

................. 16157$

CH38

02-08-07 10:40:18

PS

PAGE 553

.

554 /

Figure 4.5.5. Vertical distribution of tunnels of Sphaeroma triste in four wood preservative (CCA)-treated pilings at Koke Settlement, Port Moresby, expressed as a percent of the total number of tunnels for that piling. N number of pilings per tunnel; MSL mean sea level; LWS low water spring tide level.

................. 16157$

CH38

02-08-07 10:41:28

PS

PAGE 554

Marine Wood-Boring Invertebrates of New Guinea / 555

stenohaline brackish water species; Nausitora dunlopei, N. hedleyi, and N. globosa as euryhaline brackish water species; and Dicyathifer manni, Bankia gracilis, B. rochi, and Lyrodus bipartitus as euryhaline marine species. Even at the same site, borer species may be separated by their salinity tolerances. For example, at the main wharf in Lae, a clear salinocline often forms, with the turbid waters of the Markham River lying above waters close to full marine salinity. The exact nature of the salinocline varies considerably through the year and even during a single day (Cragg, pers. obs.). The vertical distribution of the most common borers at that site reflects this, with Nausitora hedleyi, Nausitora sp. nov., and Lyrodus bipartitus colonizing panels placed above the typical position of the salinocline, and Bankia bipalmulata, B. carinata, and Lyrodus pedicellatus below (Figure 4.5.6). The isopods Sphaeroma terebrans and S. triste are widely distributed in New Guinea, ranging from full salinity to very brackish waters. S. triste has been found at salinities as low as 0.2 psu (Cragg and Aruga 1988). Other known estuarine isopod wood borers are Anopsilana pustulosa (Holdich et al. 1981) and Corallana estuaria (Jones et al. 1983). The distribution of limnoriids, on the other hand, suggests that they have limited tolerance of lowered salinity. They occur to the east and west of the delta complex of the Gulf of Papua, but were not found during extensive collecting in the mangroves of the complex itself, which is, due to considerable riverine input, to the landward of the 20 psu isohaline (Cragg and Aruga 1988). Elsewhere, limnoriids have been found in mangrove wood (Kensley and Schotte 1987), so it is likely that it is low salinity, rather than the nature of the mangrove wood, that excludes these organisms from the deltas. Collections in the deltas to the west of Merauke could be used to test this hypothesis. The distribution of borers is strongly influenced by the availability of wood. Mangrove forests retain large quantities of fallen wood as many mangrove species have very dense (non-floating) timber. These become riddled by teredinids (Figure 4.5.7) and sometimes by sphaeromatids. The large rivers of New Guinea are prone to meandering in their flood plains, and because meanders involve the undercutting of river banks, considerable quantities of trees from rainforests get carried down to coastal areas (Cragg 1983). Some are carried offshore where they may become colonized by borers and waterlogged (see Table 4.5.3 for rainforest timbers collected during prawn trawling on the continental shelf around New Guinea). Wharf construction results in substantial input of wood into marine environments where borers are active. Most of the species listed in Tables 4.5.1 and 4.5.2 are dependent on wood for shelter, and in the case of teredinids and limnoriids, for food. Sphaeromatids, on the other hand, may burrow into other substrates (Harrison and Holdich 1984; Cragg 2003) and anopsilanids are cryptozoic, exploiting cavities of various sorts (Holdich et al. 1981; Bruce 1986, pers. obs.). Of the corallanids, some occur in colonies of Sphaeroma and only Corallana nodosa has been found in colonies on its own (Jones et al. 1983). All teredinids except Zachsia and Kuphus are obligate wood borers. Kuphus is a mud borer (Turner 1966). A probably undescribed species of Zachsia was found

................. 16157$

CH38

02-08-07 10:41:29

PS

PAGE 555

.

556 /

Figure 4.5.6. Vertical distribution of teredinids in a vertical array of panels of timber exposed at Lae for 113 days, expressed as a percentage of the total number of individuals identified for the teredinid species in question. Three replicate panels were retrieved at 0.5 m depth and two panels at each of the other depths. Total number of individuals identified: B. bipalmulata—916; B. carinata—30; L. bipartita—67; L. pedicellatus—161; N. hedleyi—171; N. sp. nov.—219. in rhizomes of the seagrasses Enhalus acoroides and Cymodocea. Rhizomes that were colonized were often close to the burrows of callianassid shrimps, where burrowing activity may have exposed the usually buried structure (Cragg, pers. obs.). Dicyathifer and Bactronophorus appear to be mangrove specialists, only rarely occurring in fixed wood or driftwood (Rayner 1983). Uperotus clavus appears to be restricted to palm nuts (Turner 1971).

Ecological Roles of Borers Teredinid borers dominate the niche created by intertidal wood, partly because they grow larger than the other borers, but also because they have the capacity to

................. 16157$

CH38

02-08-07 10:42:22

PS

PAGE 556

Marine Wood-Boring Invertebrates of New Guinea / 557

Figure 4.5.7. Mangrove log riddled with teredinid tunnels lodged among prop roots of the mangrove Rhizophora. produce large numbers of offspring. They pass more than 50% of the wood volume through their guts before fragmentation, tidal export, or burial takes place. Microbial degradation becomes more significant after fragmentation. Because approximately half of mangrove primary production is converted into woody tissue, and most of this is recycled after falling to the forest floor, teredinids play a key role in energy flow in undisturbed mangrove ecosystems where wood is not extracted for commercial use (Cragg 1993). Tunneled wood (see Figure 4.5.7) provides refuge for a wide range of organisms in the mangrove intertidal. In particular, juvenile crabs and small fish exploit the protection from desiccation and predators offered by such wood (Cragg, pers. obs.). The activities of some borers create conditions favorable to activities of others. The amphipod Chelura insulae has been reported to be found associated with colonies of Limnoria tripunctata, L. platycauda, L. sexcarinata, and Paralimnoria andrewsi (Ku¨hne 1976) and with L. kautensis and P. asterosa (Cookson and Cragg 1988). The Cirolana species that occur with Sphaeroma may take over tunnels created by the sphaeromatids, or they may prey on them. Anopsilana willeyi has been

................. 16157$

CH38

02-08-07 10:42:26

PS

PAGE 557

Location (lat/long)

8 58 /143 52

8 53 /143 56

8 45 /144 10

8 37 /144 12

8 37 /144 15

8 32 /144 20

8 21 /145 08

7 58 /145 14

7 58 /145 15

not available;

Site

A

................. 16157$

B

CH38

C

D

E

F

02-08-07 10:42:27

G

PS

H

I

Note: na

34.8

35

na

34.4

na

34.4

na

na

na

Salinity (PSU)

indicates species present at the location.

55

50

60

30-38

27-30

25-30

30

10-24

29–30

Depth (m)

Alstonia ?spectabilis R. Br.

Unidentified species

Ficus sp.

Unidentified species

Hernandia sp.

Bruguiera sp.

Hopea sp.

Unidentified species

Pterocymbium beccarii K.Schum.

Wood species

Teredo mindanensis

Martesia striata

Table 4.5.3. Wood-boring bivalves found in wood trawled from the sea bed in the Gulf of Papua

558 /

.

PAGE 558

Bankia bipennata Bankia carinata

Bankia campanellata

Teredothyra matocotana

Marine Wood-Boring Invertebrates of New Guinea / 559

observed to feed on teredinids and Sphaeroma (Cherian 1980). A group of 15 to 20 of these cirolanids can consume a teredinid in a few minutes. As A. pustulosa can be trapped using fish bait (Bruce 1982b), it too is likely to feed on carrion.

Economic Consequences of Borer Activity Wood is a valuable construction material: it has a lower energy cost than alternative materials, a good strength to weight ratio, and is readily shaped without the need for special tools. It is widely used for coastal construction. In traditional village houses, posts are used to raise the house clear of the reach of tides. Wood is also a favored material for traditional boats. Modern construction often uses timber for wharf components: fender and load-bearing piling, walings between pilings, decking, and curbing. Any timber exposed in the intertidal zone and below, in seawater, or in brackish waters in New Guinea waters is at risk from to attack by the borers described above. The rate of attack is often too great to permit the economic use of untreated timber (Shillinglaw and Moore 1947). Wood preservatives such as creosote and CCA (a mixture of water-soluble copper, chromium and arsenic compounds that form a leach-resistant bond when impregnated under pressure into wood) have been tested in New Guinea waters (Tamblyn et al. 1978; Eaton et al. 1989). The resistance of the wood to teredinids and limnoriids is considerably enhanced by such treatments, but Sphaeroma spp. are capable of causing devastating damage even to heavily treated timbers (see Figures 4.5.8 and 4.5.9, and Cragg and Levy 1979). Timbers treated with creosote enhanced by contact insecticides have been tested in New Guinea waters and have not proved immune to attack by Sphaeroma (Eaton and Cragg 1996). This organism continues to be a threat to marine timber construction in subtropical and tropical waters worldwide (Cragg 2003). However, because its attack is focused in the lower intertidal zone (Cragg and Levy 1979), additional protection in the form of a physical barrier of plastic, copper, or even concrete may provide protection (Figure 4.5.10).

Literature Cited Bowman, T.E. 1977. Ceratolana papuae, a new genus and species of mangrove-boring cirolanid isopod from Papua New Guinea. Proc. Biol. Soc. Wash. 90: 819–825. Bruce, N.L. 1982a. On the genus Corallana Dana, 1952 (Isopoda, Corallanidae) with description of a new species from western Australia. Crustaceana 42: 241–249. Bruce, N.L. 1982b. Records of isopod crustacea (Corallanidae, Cirolanidae) from Papua New Guinea with the description of a new species. J. Crust. Biol. 2: 612–618. Bruce, N.L. 1986. Cirolanidae of Australia. Rec. Aus. Mus. Sup. 6. Bruce, N.L. 1995. Range extension of the mangrove dwelling isopod genus Ceratolana Bowman (Cirolanidae). Crustaceana 68: 123–125. Cherian, C.J. 1980. Cirolana spp. (Isopoda) associated with marine wood-boring and fouling organisms. Current Sci. 49: 293.

................. 16157$

CH38

02-08-07 10:42:28

PS

PAGE 559

.

560 /

Figure 4.5.8. Piling destroyed by Sphaeroma tunneling in the intertidal zone, Port Moresby, Papua New Guinea.

Figure 4.5.9. Cross-section of a pile superficially riddled with burrows of Sphaeroma. Note how burrows penetrate perpendicularly from the surface.

................. 16157$

CH38

02-08-07 10:42:34

PS

PAGE 560

Marine Wood-Boring Invertebrates of New Guinea / 561

Figure 4.5.10. Timber pilings protected by jacket of concrete in the low intertidal zone, Port Moresby. Cookson, L.J. 1991. Australasian species of Limnoriidae (Crustacea: Isopoda). Mem. Mus. Victoria 52: 137–362. Cookson, L.J. 1997. Additions to the taxonomy of the Limnoriidae (Crustacea: Isopoda). Mem. Mus. Victoria 56: 129–143. Cookson, L.J., and S.M. Cragg. 1988. Two new species of Limnoriidae (Isopoda) from Papua New Guinea. J. Nat. Hist. 21 (6): 1501–1514. Cragg, S.M. 1983. The mangrove ecosystem of the Purari River. Pp. 295–324 in Petr, T. (ed.) The Purari-Tropical Environment of a High Rainfall River Basin. Monographiae Biologicae. W. Junk, The Hague. Cragg, S.M. 1988. The wood-boring isopod Sphaeroma, a threat to maritime structures in warm waters. Pp. 727–732 in Houghton, D., R.N. Smith, and H.O.W. Eggins (eds.) Biodeterioration 7. Elsevier, London. Cragg. S.M. 1993. Wood break-down in mangrove ecosystems: a review. PNG J. Agric. For. Fish. 36: 30–39. Cragg, S.M. 2003. Marine wood boring arthropods: ecology, functional anatomy and control measures. Pp. 272–286 in Goodell, B., D.D. Nicholas, and T.P. Schultz (eds.) Wood Deterioration and Preservation: Advances in Our Changing World. American Chemical Society, Oxford University Press, Oxford. Cragg, S.M., and J. Aruga. 1988. Intertidal and subtidal wood-boring faunas from the Gulf of Papua, Papua New Guinea. Pp. 236–244 in Advances in Aquatic Biology and

................. 16157$

CH38

02-08-07 10:42:38

PS

PAGE 561

.

562 /

Fisheries (N. Balakrishnan Nair Felicitation Volume). St. Josephs Press, Trivandrum, India. Cragg, S.M., and C.R. Levy. 1979. Attack by the crustacean Sphaeroma on CCA-treated softwood in Papua New Guinean waters. Int. J. Wood Pres. 1 (4): 161–168. Eaton, R.A., F. Ampong, J. Beesley, J.D. Bultman, L.J. Cookson, S.M. Cragg, J. De Palma, A. Gambetta, B. Henningson, M. Levi, C.R. Levy, T. Nilsson, and E. Orlandi. 1989. An international collaborative marine trial to investigate the effect of timber substrate on the efficacy of CCA and CCB wood preservatives. Material und Organismen 24: 51–79. Eaton, R.A., and S.M. Cragg. 1996. Evaluation of creosote fortified with synthetic pyrethroids as wood preservatives for use in the sea. Part 1: Efficacy against marine wood-boring molluscs and crustaceans. Material und Organismen 29: 211–229. Ellison, A.M., and E.J. Farnsworth. 1990. The ecology of Belizean mangrove-root fouling communities. I Epibenthic fauna are barriers to isopod attack of red mangrove roots. J. Exp. Mar. Biol. Ecol. 142: 91–104. Harrison, K., and D.M. Holdich. 1982. New eubranchiate sphaeromatid isopods from Queensland waters. Mem. Queensland Mus. 20: 421–446. Harrison, K., and D.M. Holdich. 1983. Sphaeromatid isopods (Crustacea) from brackish waters in Queensland, Australia. Zool. Scripta 12: 127–140. Harrison, K., and D.M. Holdich. 1984. Hemibranchiate sphaeromatids (Crustacea: Isopoda) from Queensland, Australia, with a world wide review of the genera discussed. Zool. J. Linn. Soc. 81: 275–387. Holdich, D.M., K. Harrison, and N.L. Bruce. 1981. Cirolanid isopod crustaceans from the Townsville region of Queensland, Australia, with descriptions of six new species. J. Nat. Hist. 15: 555–605. Jones, D.A., J.D. Icely, and S.M. Cragg. 1983. Some corallanid isopods associated with wood from Papua New Guinea. Journal of Natural History 17: 837–847. Kensely, B., and M. Schotte. 1987. New records of isopod crustacea from the Caribbean, the Florida Keys, and the Bahamas. Proc. Biol. Soc. Wash. 100: 216–247. Ku¨hne, H. 1976. Zur geographisches Verbreitung holzzersto¨render Crustaceen und Systematik der Untergattung Limnoria s. str. Menzies. Material und Organismen Suppl. 3: 543–553. Menzies, R.J. 1957. The marine borer family Limnoriidae (Crustacea: Isopoda). Bull. Mar. Sci. Gulf Caribb. 7: 101–200. Rayner, S.M. 1979. Comparison of the salinity range tolerated by teredinids (Mollusca: Teredinidae) under controlled conditions with that observed in an estuary in Papua New Guinea. Aust. J. Mar. Freshwater Res. 30: 521–533. Rayner, S.M. 1983. Distribution of teredinids (Mollusca: Teredinidae) in Papua New Guinea. Records of the Australian Museum 35: 61–76. Roch, G.F. 1961. Die Terediniden der Sunda-Inseln und Neu Guineas. Beaufortia 9: 7–48. Sawyer, G.S., and Cragg, S.M. 1995. Attack by the wood-boring weevil, Pselactus spadix on timbers in the intertidal and splash zones in ports in the U.K. Material und Organismen 29: 67–79. Schepman, M.M. 1919. Nova Guinea 13, Zoologie, p. 195, pl. 7, fig. 3. Shillinglaw, A.W., and D.D. Moore. 1947. Report of marine borer survey in New Guinea waters. Council for Scientific and Industrial Research. Bll. 223. Si, A., C.G. Alexander, and O. Bellwood. 2000. Habitat partitioning by two wood-boring invertebrates in a mangrove system in tropical Australia. J. Mar. Biol. Ass. UK 80: 1131–1132. Tamblyn, N., S.M. Rayner, and C.R. Levy. 1978. Field and marine tests in Papua New

................. 16157$

CH38

02-08-07 10:42:39

PS

PAGE 562

Marine Wood-Boring Invertebrates of New Guinea / 563 Guinea. 1: Performance of creosote and copper-chrome-arsenic in pine and eucalypt timbers in tropical marine waters. J. Inst. Wood Sci. 8 (2): 1–6. Turner, R.D. 1966. A Survey and Illustrated Catalogue of the Teredinidae (Mollusca: Bivalvia). Museum of Comparative Anatomy, Harvard University, Cambridge Massachusetts. Turner, R.D. 1971. Identification of marine wood-boring molluscs. Pp. 17–64 in Jones, E.B.G., and S.K. Eltringham (eds.) Marine Borers and Fouling Organisms of Wood. OECD, Paris. Turner, R.D., and C.B. Calloway. 1987. Species pairs in the Teredinidae. International Research Group on Wood Preservation Doc. No. IRG/WP/4142. Turner, R.D., and A.C. Johnson 1971. Biology of marine wood-boring molluscs. Pp. 259–301 in Jones, E.B.G., and S.K. Eltringham (eds.) Marine Borers and Fouling Organisms of Wood. OECD, Paris.

................. 16157$

CH38

02-08-07 10:42:39

PS

PAGE 563

4.6. Herpetofauna of Papua of Papua includes 23 families, 109 genera, and 371 species of frogs, crocodiles, turtles, lizards, and snakes (Table 4.6.1). The majority of species ( 340) are found primarily on land or in freshwater, but 24 are primarily marine (sea snakes and marine turtles). Approximately 28% of the herpetofauna is endemic to Papua and 61% of the species are endemic to New Guinea and its satellite islands. Those species that are extralimital to New Guinea are mostly shared with Australia or Maluku, or represent taxa that are widely distributed in the Indo-Pacific region. (See Appendix 8.4 for a preliminary checklist of species from the Papua and the nearby Aru Islands.) All of the satellite islands associated with Papua are politically part of that province and are included in this treatment. In addition, the Aru Islands, which are politically part of Maluku, are also included because they are biogeographically associated with New Guinea and were once connected by land to that island. It is instructive to compare the herpetofauna of Papua with that of eastern New Guinea (the eastern half of mainland New Guinea plus satellite islands) and Papua New Guinea (eastern New Guinea plus the Admiralty and Bismarck archipelagos and Bougainville Island; Table 4.6.1). Papua is inhabited by a few taxa that belong to lineages that are widespread in Southeast Asia but just reach the western fringes of New Guinea. These include the snake family Cylindrophiidae, represented by a single species endemic to the Aru Islands, and the lizard family Dibamidae, represented by a widespread Southeast Asian species that reaches western New Guinea. Otherwise all three areas have the same 21 families of amphibians and reptiles. Papua and eastern New Guinea both have 109 genera, most of which are common to both areas. Papua New Guinea has slightly more genera (117), largely because it has some unique lineages that occur on the Bismarcks and on Bougainville Island (e.g., several ranid frog genera). The differences in numbers of species are much more dramatic. Papua New Guinea has many more species than does eastern New Guinea. Again, this reflects the inclusion of lineages from the Admiralty and Bismarck archipelagos and Bougainville Island in the Solomon Archipelago. However, when these areas are excluded, the number of species in east New Guinea (475) is almost a third greater than the 371 species reported from Papua (west New Guinea). These differences probably reflect a much higher level of field survey and species description activity for eastern New Guinea (mainland Papua New Guinea) than for Papua. This is apparent in Brown’s (1991) monograph of the lizard genus Emoia, for which less than an eighth of the roughly 10,000 museum specimens that he obtained from the New Guinea region were from Papua; the rest were from Papua New Guinea. It is clear from recent survey activity that the herpetofauna of Papua Province

T

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

564

................. 16157$

CH39

02-08-07 10:40:09

PS

PAGE 564

................. 16157$

CH39

4

5

1

6

7

23

Frogs

Turtles

Crocodiles

Lizards

Snakes

Totals

21

6

5

1

5

4

Families Eastern New Guinea

21

6

5

1

5

4

PNG

109

41

33

1

11

23

Papua

109

37

30

1

11

30

Genera Eastern New Guinea

117

40

32

1

11

33

PNG

371

83

141

2

15

130

Papua

475

90

152

2

16

215

Species Eastern New Guinea

553

103

183

2

16

248

PNG

Note: Eastern New Guinea includes the eastern half of mainland New Guinea and satellite islands; PNG includes the eastern half of mainland New Guinea plus the Admiralty and Bismarck archipelagos and Bougainville Island. Papua and PNG share 237 species, and Papua and Eastern New Guinea share 236 species. Taxa from the Aru Islands are included in the totals for Papua because this area is biogeographically part of New Guinea, although it is politically part of Maluku Province, Indonesia.

Papua

Taxon

Table 4.6.1. Herpetofauna (native species) of Papua, eastern New Guinea, and Papua New Guinea (PNG) Herpetofauna of Papua / 565

02-08-07 10:40:11

PS

PAGE 565

566 /

is very incompletely known; 63 species (17% of the herpetofauna) have been described since 1980 and recent fieldwork has demonstrated the existence of many additional new species (Allison and Dwiyahreni 1998; Richards, Iskandar, and Allison 2000; Richards, Iskandar, and Tjaturadi 2002). It is likely that the number of frog species will more than double, and perhaps triple, and that there will be significant increases in the number of lizards and modest increases in the number of turtles and snakes when all species have been discovered and described. It is also instructive to compare New Guinea, the world’s largest and highest tropical island, with Borneo, the world’s second largest tropical island. New Guinea has a land area of 790,000 km2; Borneo is only slightly smaller, with a land area of 743,330 km2. Borneo formed from ‘‘Mesozoic accretion of oceanic crustal material (ophiolite), marginal basin fill, island arc material and microcontinental fragments’’ onto a Paleozoic continental core (Moss and Wilson 1998: 137). These processes continued into the Cenozoic, accompanied by considerable erosion, and by the late Miocene the drainage systems on Borneo were similar to those today. New Guinea had a similar, but much more complex, geological history (Chapter 2.1), involving the accretion since the early Cenozoic of island arcs, sea mounts, and plate fragments, together with considerable uplift and volcanism (Dow 1977). The island today is composed of 32 geological terranes, many of which are centers of endemism (Pigram and Davies 1987; van Welzen 1997). Both islands support a rich diversity of tropical forests. The floras of the two islands are estimated to include a similar number of species, about 14,000 (Roos et al. 2004). Borneo is part of the Sunda Shelf and was connected to mainland Southeast Asia until the late Eocene or early Miocene, and subsequently has been repeatedly connected to mainland Southeast Asia during periods of lowered sea level, most recently ca 17,000 during the last glacial maximum (Voris 2000). Consequently, its biota is broadly representative of that of Southeast Asia, which has a similar climate. New Guinea has a more recent geological origin and has always been separated by sea from Southeast Asia. Its biota has been strongly influenced by both Australia and Southeast Asia. Although both islands are mountainous, the mountains in Borneo form something of a small central core surrounded by lowlands. New Guinea, in contrast, has numerous isolated mountain ranges, and isolated high mountains, separated by lowland basins. These differences in geological history and in physiognomy are reflected in the differences in the herpetofaunas of these two islands (Table 4.6.2). Borneo has a third more families and nearly a quarter more genera than does New Guinea, reflecting the biogeographic association of Borneo to mainland Southeast Asia and the relative isolation of New Guinea from source areas with a similar wet tropical climate. However, New Guinea has nearly a third more species than does Borneo. This likely resulted from a higher number of geographic isolating mechanisms and therefore a higher level of species radiation on New Guinea compared to Borneo. In order words, New Guinea has far higher beta diversity than does Borneo. Borneo does, however, have higher species richness in snakes than does New Guinea, reflecting the high diversity of snakes in the Southeast Asian region.

................. 16157$

CH39

02-08-07 10:40:11

PS

PAGE 566

Herpetofauna of Papua / 567

Table 4.6.2. Herpetofauna of Borneo and New Guinea Taxon

Families Borneo New Guinea

Genera Borneo New Guinea

Species Borneo New Guinea

Frogs

6

4

32

29

150

282

Caecelians

1

0

2

0

4

0

Turtles

5

5

13

11

13

17

Crocodiles

1

1

2

1

3

2

Lizards

8

6

36

35

113

193

Snakes

9

7

63

43

155

109

Totals

30

23

148

119

438

603

Source: Totals for New Guinea from Allison (1996, updated). Figures for Borneo from Inger and Tan (1996), Stuebing (1991, 1994), Das and Ismail (2001), and King and Burke (1989). Totals were updated to include species described subsequently to these publications.

History The first collection of amphibians and reptiles from Papua was made in 1824 by Rene´-P. Lesson on the voyage of the corvette Coquille, which sailed along the north coast of New Guinea under the command of Capt. Louis Duperrey and visited Waigeo Island (Chapter 1.2). R. P. Lesson’s specimens, which were deposited in the Paris Museum, formed the basis for new species descriptions by Pierre-Adolphe Lesson (1830). This work was followed during the next decade or so by visits from several additional global exploring expeditions, mostly under the command of French and Dutch navigators. Naturalists on these expeditions made small collections of specimens from costal regions of Papua and later deposited these in European museums. Much of this material was included in the comprehensive review of the world’s herpetofauna by A. M. C. Dume´ril and G. Bibron (1836, 1837, 1839) who described several taxa that are endemic to New Guinea and a number of Indo-Pacific taxa that occur in New Guinea. By the 1840s much of coastal New Guinea had been mapped and the era of major world exploring expeditions was coming to a close. In the New Guinea region, subsequent exploration work began focusing mainly on areas with early European mission settlements. Small opportunistic collections of specimens of amphibians and reptiles made in association with this work found their way to European museums where they were studied and reported on by taxonomists. However, by 1850, only about 80 (22%) of the species now known from Papua had been described. Most of these were widespread Indo-Pacific taxa that were named from specimens collected from areas outside Papua. During the second half of the 19th century a number of naturalists visited Papua and some made significant collections. In 1858 Alfred Russel Wallace spent four months near Dore´ Bay (Manokwari) collecting zoological specimens. He focused mainly on insects and birds but did collect a few herps. Adolf Meyer in the early

................. 16157$

CH39

02-08-07 10:40:12

PS

PAGE 567

568 /

1870s visited the Scholten Islands and areas on mainland New Guinea along Cenderawasih Bay, including adjacent mountains. He claimed to have crossed the Bird’s Neck region of New Guinea to Bintuni Bay but this now seems unlikely (Souter 1964). Meyer did, however, make significant zoological collections and later described 19 species of herps that are today accepted as valid (Meyer 1874). Meyer’s collections were deposited in the Dresden Museum and were mostly destroyed during World War II. In 1872–1873 Luigi D’Albertis and Odoardo Beccari collected in the Arfak Mountains, becoming the first naturalists to work in the interior of New Guinea. Their collections, which were mostly deposited in the Museo Civico di Storia Naturale ‘‘Giacomo Doria’’ in Genoa together with those made by the Dutch feather merchant A. A. Bruijn and deposited in Amsterdam, were studied by W. Peters and G. Doria (1878) who described 20 species recognized as valid today. By the end of the 19th century taxonomists had described 58 species that occur in Papua and are now regarded as endemic to the New Guinea region. However, most of these were described from eastern New Guinea, which was receiving considerable attention, mainly from European naturalists, especially Georges Boulenger of the British Museum who between 1885 and 1900 described 39 currently recognized species from New Guinea (Boulenger 1921). However, only one of these is endemic to Papua and overall, only 16 species endemic to Papua had been named by 1900. In the early 1900s the Dutch initiated a number of major exploring expeditions to the interior of Papua, including the massive series of Dutch Military Expeditions from 1907 to 1917. In 1903 the Dutch explorer H. A. Lorentz began the first of several expeditions to southern New Guinea with the ultimate aim of reaching the summit of the highest peak in New Guinea, known today as Mt Jaya (Lorentz 1913). In 1909–1910 the British Ornithologists’ Union conducted a huge expedition centered mainly in the Mimika River drainage, also with the aim of reaching the summit of Mt Jaya. The Mimika River proved to be a poor starting point from which to reach Mt Jaya and the British Ornithologists’ Union expedition, on account of many difficulties, only reached the foothills. One of the expedition members, A. F. R. Wollaston, returned to the area in 1912–1913 with a smaller group and managed to reach the glaciers on the Mt Jaya massif (Boulenger 1914; Wollaston 1912, 1916). Naturalists on these various expeditions made significant collections and by 1915 the number of recognized amphibian and reptile species endemic to Papua had doubled. Around the start of World War I, Nelly de Rooij of the Amsterdam Museum published a definitive two-volume comprehensive treatment of the reptiles of New Guinea and Southeast Asia (de Rooij 1915, 1917). P. N. van Kampen (1923) later produced a similar but far less critical treatment of amphibians. A decade or so later H. W. Parker of the British Museum published a very thorough and detailed revision of the frog family Microhylidae and placed the Indo-Australian species into a modern taxonomic framework (Parker 1934). These efforts, particularly de Rooij’s and Parker’s, produced a strong foundation for subsequent

................. 16157$

CH39

02-08-07 10:40:13

PS

PAGE 568

Herpetofauna of Papua / 569

taxonomic work on the New Guinea herpetofauna. Collecting activity declined during World War I (although Boulenger named 21 species between 1911 and 1920, including eight that are now regarded as endemic to Papua), then picked up again around 1920 and remained reasonably active until World War II (although only 14 species endemic to Papua were described between 1920 and 1950). In 1938–1939 members of the Third Archbold Expedition collected throughout the Mamberamo River basin, obtaining large samples of amphibians and reptiles. This material, together with that from Archbold Expeditions to other parts of New Guinea, was deposited mainly in the American Museum of Natural History in New York where, in the mid-1950s, R. G. Zweifel began working on the New Guinea herpetofauna, mainly frogs. In an extraordinary career spanning six decades, Zweifel has published more than 50 papers on New Guinea frogs and lizards, including a number of major monographs, and has described or co-described more than 70 species of currently recognized New Guinea species. Towards the end of the Dutch Administration, Prof. L. D. Brongersma, a herpetologist from the Leiden Museum, organized an enormous expedition to the Digul River drainage and the Star Mts that resulted in large collections of amphibians and reptiles (Brongersma 1962). This material has yet to be fully studied but has resulted in numerous new species descriptions (e.g., Brown 1991; Zug 2004). During the next several decades there was relatively little fieldwork in Papua and from 1950 to 1980 only eight endemic amphibians and reptiles were described. In subsequent decades the tempo of fieldwork has markedly increased. This includes a number of expeditions conducted by the Museum Zoologense Bogoriense, or under the auspices of Conservation International’s Rapid Assessment Program (Nasution et al. 1996; Richards, Iskandar, and Allison 2000; Richards, Iskandar, and Tjaturadi 2002). In 1997 biologists from the Bishop Museum and the Museum Zoologense Bogoriense undertook a major herpetological survey of the PT Freeport Contract of Work area, from sea level to ca 3,500 m (Allison and Dwiyahreni 1998). Beginning in the late 1990s Rainer Gu¨nther of the Berlin Museum began studying the frogs and lizards of Papua, conducting fieldwork around Nabire, on Biak and Yapen, the Wandammen Peninsula, and Fakfak Mountains, describing many new species (Gu¨nther 1999, 2000a,b, 2001, 2002a,b, 2003a,b,c,d, 2004a,b; Gu¨nther and Richards 2000; Gu¨nther, Richards, and Iskandar 2001; Gu¨nther and Ro¨sler 2002). Allen Allison and Hellen Kurniati collected in the Tangguh area of the Bomberai Peninsula in March 2002 (Allison and Kurniati, in prep.). Most recently, there has been something of an unfortunate trend to describe new taxa from specimens collected in association with the pet trade, often without specific locality information, which are deposited in museums when they die.

Origin and Composition of the Herpetofauna There are only four native families of frogs in New Guinea. The Myobatrachidae, with the fewest species, includes five genera and only seven New Guinea species,

................. 16157$

CH39

02-08-07 10:40:13

PS

PAGE 569

570 /

five of which occur in Papua. These latter taxa include three species of Lechriodus (Figure 4.6.1), all endemic to New Guinea (a fourth species is endemic to Australia), together with Limnodynastes convexiusculus and Crinia remota, both of which also occur in southwest Papua New Guinea and northern Australia. Uperoleia lithomoda also occurs in southwest Papua New Guinea and northern Australia but has not been reported from Papua. The seventh New Guinea species, Mixophyes hihihorlo, is endemic to the southern highlands of Papua New Guinea. The remaining 121 species of myobatrachids are all endemic to Australia. There is some uncertainty in the assignment of these frogs to family. The myobatrachids have classically been thought to be closely related to the leptodactylids of South America and are sometimes included in that family. However, most modern treatments recognize the Myobatrachidae as a full family (e.g., Duellman and Trueb 1986) with at least two subfamilies, the Myobatrachinae, which includes Crinia and Uperoleia, and the Limnodynastinae, which includes the remaining New Guinea species. However, Frost (2005) follows Zug, Vitt, and Caldwell (2001), in treating these subfamilies as full families and mentions the possibility that Limnodynastidae may be a sister taxon of the Heleophrynidae, a family of six species in the genus Heliophryne, which are all endemic to South Africa. It seems fairly clear that the Myobatrachids, in the broad sense, are of Gondwanan origin. They seem to have reached New Guinea relatively recently, as judged by their low diversity in New Guinea compared to Australia, and by the fact that only two species (Lechriodus aganoposis and L. platyceps; Figure 4.6.1) occur north of the central mountains in New Guinea.

Figure 4.6.1. Myobatrachidae: Lechriodus platyceps.

................. 16157$

CH39

02-08-07 10:41:28

PS

PAGE 570

Herpetofauna of Papua / 571

The family Hylidae is represented by only two genera in the New Guinea region: Litoria (Figure 4.6.2) and Nyctimystes, which have a combined total of 149 species distributed among Australia (68 species), Timor and the Lesser Sunda Islands (1), Solomon Islands (2), New Guinea (87), and Maluku (3). There are 37 species of Litoria recorded for Papua, of which 14 (38%) are endemic. There are five species of Nyctimystes in Papua; three of these (60%) are endemic. About ten species of Litoria occur in both New Guinea and Australia. In the west, Nyctimystes has reached Halmahera where it is represented by a single species, N. rueppelli. Two species of Litoria, L. amboinensis and L. vagabunda, occur on both New Guinea and Seram, and, as the name implies, L. amboinensis occurs westwards to Ambon. Litoria capitula is endemic to Tanimbar. In the east, two species of Litoria, L. thesaurensis and L. lutea, are found in the Solomon Islands where L. lutea is endemic. New Guinea hylids reach their greatest diversity in the central mountains (Allison, Kraus, and McShane 2004). Hylids occur throughout much of the world except for Africa and south and Southeast Asia. They are represented in the Australia-New Guinea region by an endemic subfamily, the Pelodryadinae, which some elevate to full family status. The absence of hylids in southern Asia, and their presence in Australia and South America, suggests that the family is of west Gondwanan origin (Tyler 1999). They likely reached New Guinea as that island began forming during the Miocene and have radiated extensively (Zweifel and Tyler 1982). Approximately half of the frogs in New Guinea are members of the nearly-

Figure 4.6.2. Hylidae: Litoria angiana.

................. 16157$

CH39

02-08-07 10:42:32

PS

PAGE 571

572 /

cosmopolitan family Microhylidae (Figure 4.6.3). The New Guinea taxa are currently classified into 19 genera within two subfamilies, Asterophryinae and Genyophryinae. Most of the genera are endemic to New Guinea but appear to be related to microhylid genera found in the Indo-Malaysian region and may be derived from a common ancestor from that area (Kuramoto and Allison 1989). Only two genera, Cophixalus and Austrochaperina, reach Australia, where the species are restricted to tropical regions of the northeast (Zweifel and Tyler 1982; Zweifel 1985, 2000) and are thought to have originated from ancestors in New Guinea (Zweifel and Tyler 1982). There are 15 genera and 63 species of microhylids known from Papua, of which 41 species are endemic. The four New Guinea genera not represented in Papua are Barygenys, a small clade of fossorial (burrowing) frogs distributed over much of eastern New Guinea (Zweifel 1972a, 1980); Pherohapsis, a monotypic genus endemic to upland savannas on the Sogeri Plateau of Papua New Guinea (Zweifel 1972a); Genyophryne, a monotypic genus occurring in northeastern New Guinea and offshore islands (Zweifel 1971); and Aphantophryne, which includes three leaf litter-inhabiting species that are endemic to the

a

b

Figure 4.6.3. Microhylidae: a. Sphenophryne cornuta; b. Xenobatrachus obesus; c. Xenorhina arboricola.

c

................. 16157$

CH39

02-08-07 10:42:37

PS

PAGE 572

Herpetofauna of Papua / 573

Owen Stanley Mts and associated ranges of Papua New Guinea (Zweifel and Parker 1989). New Guinea frogs in the family Ranidae are currently classified into three widely distributed Indo-Pacific genera: Limnonectes, Rana, and Platymantis. Limnonectes, which is sometimes treated as a subgenus of Rana, is represented in New Guinea by a single native species, L. grunniens, which also occurs in Java, Sulawesi, and parts of Maluku (Frost 2005). Its distribution in New Guinea is limited to far western Papua, suggesting that it may be a relatively recent immigrant. All other species of Limnonectes (ca. 50 total) are confined to south and Southeast Asia, southern China and southern Japan. There are ten species of Rana recorded from Papua, two of which are endemic to the province. The rest are shared with Papua New Guinea and are all endemic to the island of New Guinea except for Rana daemeli, which also occurs in northern Australia and in the Bismarck Archipelago (Menzies 1987). Rana has a nearly worldwide distribution but the species groups found in New Guinea appear to be fairly closely related to ranid clades from Southeast Asia, suggesting that they may have originated from common ancestors there. The third genus, Platymantis (Figure 4.6.4), includes six species from Papua, five of which are endemic. The sixth species, P. papuensis, is shared with Papua New Guinea and extends to New Ireland. Other members of the genus occur in Palau, the Philippines, the Bismarck and Admiralty archipelagos, the Solomon

Figure 4.6.4. Ranidae: Platymantis papuensis.

................. 16157$

CH39

02-08-07 10:43:41

PS

PAGE 573

574 /

Islands, and Fiji (Allison 1996) but some of these probably represent separate radiations.

There are three families of freshwater turtles occurring in New Guinea, all of which occur in Papua. The smallest family, Carettochelidae, has an extensive worldwide fossil record dating back to the Eocene (Ernst and Barbour 1989) but has only one extant species, Carettochelys insculpta, that is endemic to southern New Guinea and northern Australia and is clearly relictual (Cann 1998). Its fossil history in New Guinea dates back to the Miocene (Glaessner 1942). Carettochelidae is thought to be closely related to the family Trionychidae, which has a worldwide distribution and is represented in New Guinea by two closely related species in the genus Pelochelys (Rhodin, Mittermeier, and Hall 1993; Webb 1995, 1997, 2002). Other species of Pelochelys are found in Southeast Asia and are known to tolerate saltwater, suggesting that ancestral forms may have originally dispersed to New Guinea from points west (Rhodin, Mittermeier, and Hall 1993). All remaining freshwater turtles occurring in New Guinea are side-necked species in the family Chelidae, which is also represented by a large assemblage of species in Australia and one in Timor (Cann 1998; Iskandar 2000). There are three genera and six currently recognized species of chelid turtles known from Papua; one of these, Chelodina reimanni, is endemic. The other species are all shared with Papua New Guinea. Three of these are endemic to the island of New Guinea and two, Chelodinia rugosa (formerly C. siebenrock) and Emydura subglobosa, are also shared with Australia (Cann 1998; Iverson 1992, 1996; Iskandar 2000; Georges et al. 2002; McCord and Thomson 2002). All six species are found in swamps or slow-flowing rivers on the south coast of New Guinea and at least one of them, Elseya novaeguineae, is also found in similar habitats on the north coast. In addition, Iskandar (2000) lists three undescribed species of Elseya from Papua. Other members of the family Chelidae occur in South America and the family is almost certainly of Gondwanan origin (Ernst and Barbour 1989). There are six species of marine turtles reported from Papua: Caretta caretta (Loggerhead), Chelonia mydas (Green), Eretmochelys imbricata (Hawksbill), Lepidochelys olivacea (Olive Ridley), Natator depressus (Flatback), and Dermochelys coriacea (Leatherback). Four of these, Green, Hawksbill, Olive Ridley, and Leatherback, nest in the province, mainly in the Raja Ampat Islands, beaches on the Vogelkop, and the islands of Cenderawasih Bay. All of the marine turtles are widely distributed in the world’s oceans; they nest mainly on tropical beaches (Groombridge 1982; Ernst and Barbour 1989; Spotila 2004).

Crocodilians represent an ancient evolutionary lineage that dates back more than 200 million years to the Triassic period when there was a single supercontinent

................. 16157$

CH39

02-08-07 10:43:42

PS

PAGE 574

Herpetofauna of Papua / 575

called Pangea. As Pangea subsequently split into smaller landmasses and ultimately into today’s continents, crocodilians were dispersed around the world. They are now found mostly in tropical and subtropical areas. Although there is a rich fossil record, the geographical origin of the Australasian species remains obscure (Ross 1989). The two crocodiles that occur throughout New Guinea are Crocodylus porosus, which is found from Australia to India and readily enters salt water, and C. novaeguineae, which is endemic to the New Guinea region. It is thought to be closely related to taxa from Borneo and the Philippines (Ross 1989, 1990). Some specialists believe that the southern and northern New Guinea populations of C. novaeguineae, both of which occur in Papua, are genetically distinct with the southern population being as yet unnamed (Hall 1989).

There are six families of lizards in New Guinea, one of which, Dibamidae, is represented only by Dibamus novaeguineae, a species that is widely distributed in eastern Indonesia and the Philippines, but in New Guinea is restricted to the Vogelkop Peninsula and Waigeo Island. Other species in the family, all included in the genus Dibamus, are found mainly in south and Southeast Asia (Greer 1985). The remaining lizard families—Scincidae, Gekkonidae, Pygopodidae, Varanidae, and Agamidae—are all shared with Australia and all but the Pygopodidae are also shared with south and Southeast Asia. The zoogeographic affinities of these families are complex and still being elucidated. All of these families have had a long history in Australia and many New Guinea species and some genera have clear affinities to clades in Australia. There appears to have been considerable interchange of taxa between Australia and New Guinea since the early Miocene (Aplin, Baverstock, and Donnellan 1993). All but the Pygopodidae have likely had a long history in New Guinea and have radiated extensively there, particularly the skinks and geckos. Other New Guinea species and genera seem to have originated from areas to the west. The agamids are represented by six genera and approximately 18 species in Papua. These include Chlamydosaurus kingii, Diporiphora bilineata, and Lophognathis temporalis, which all occur in savanna areas of northern Australia and southern New Guinea, and Hydrosaurus amboinensis, which is widely distributed in Indonesia and reaches its easternmost range in the forested lowlands of Papua. The remaining agamids are all in the genus Hypsilurus (Figure 4.6.5), a clade that includes about 12 species in New Guinea and two species in eastern Australia. There are eight species reported from Papua. Four of these are endemic (Wermuth 1967; Urban 1999). Geckos in the pantropical family Gekkonidae are well represented in New Guinea with 36 native species recorded of which 26 occur in Papua (Bauer and Henle 1994). These are classified into eight genera, all of which occur throughout the Indo-Pacific region (Figure 4.6.6). The genus Cosymbotus is represented by a single species, C. platyurus. The closely related genus Hemidactylus includes two

................. 16157$

CH39

02-08-07 10:43:43

PS

PAGE 575

576 /

b

a

Figure 4.6.5. Agamidae: a. Hypsilurus dilopus; b. Hypsilurus papuensis.

a b

Figure 4.6.6. Gekkonidae: a. Cyrtodactylus mimikanus; b. Gekko vittatus.

................. 16157$

CH39

02-08-07 10:43:59

PS

PAGE 576

Herpetofauna of Papua / 577

species that occur in Papua, H. frenatus and H. garnotii. The genus Hemiphyllodactylus is represented by a single species, H. typus. All four of these species are human commensals that have spread throughout much of the Indo-Australian area and all but Cosymbotus occur throughout Oceania. Three additional Australasian genera, Gekko, Lepidodactylus, and Nactus each include two Papuan species; some of the six species are human commensals and the rest are local or regional endemics. The two largest genera, Cyrtodactylus, with seven species, and Gehyra, with eight species reported from Papua, include a mix of relatively widespread taxa as well as local endemics. Only three species of geckos, Cyrtodactylus aaroni, C. irianjayensis, and Gehyra leopoldi are endemic to Papua. However, several genera, particularly Gehyra, Gekko, and Nactus are in serious need of revision and probably include many more species than are currently recognized for Papua. The pygopodids are slender, legless lizards closely related to geckos and sometimes included in that family (Han, Zhou, and Bauer 2004; Kluge 1974; Zug, Vitt, and Caldwell 2001). As with geckos, they have fused transparent eyelids. Only two species are represented in New Guinea and both occur in Papua. One species, Lialis burtonis, is widely distributed in Australia but in New Guinea is restricted to savanna regions along the south coast. The other, L. jicari, is endemic to and widespread in New Guinea, but in Papua is known only from the northeast coast. Pygopodids are otherwise restricted to Australia where there is a modest radiation; the New Guinea taxa are almost certainly of Australian origin (Cogger 2000). The skinks are the largest lizard family in the Australasian region and include at least 124 species in New Guinea (65% of the lizard fauna) of which 87 occur in Papua (Figure 4.6.7). These species are all currently classified into the subfamily Lygosominae and include 16 genera found in Papua. The sole Papuan representative of the pantropical genus Mabuya is M. multifasciata, which occurs throughout southern and Southeast Asia, but is restricted on New Guinea to far western Papua where it may have originated as a human introduction. Two small skink genera, Papuascincus and Lobulia, are endemic to New Guinea and include small radiations of species, most of which occur in Papua. Several more, including Prasinohaema, Lygisaurus (sometimes included in Carlia), and Tribolonotus (Figure 4.6.7a) are endemic to the greater New Guinea-Solomon Islands region. At least two genera, Ctenotus and Tiliqua, are largely confined to Australia. They are each represented in Papua by single species: Ctenotus robustus, which is shared with Australia, and Tiliqua gigas (Figure 4.6.7b) which is widespread throughout much of eastern Indonesia and occurs throughout New Guinea. The genus Carlia is largely confined to New Guinea and Australia and has radiated extensively in both regions (Ingram and Covacevich 1989; Stuart, Hugall, and Moritz 2002; Zug 2004). The genus Lamprolepis is centered mainly in Southeast Asia and is represented in Papua by a single species, L. smargadina (Figure 4.6.7c) that occurs from the Solomon Islands across New Guinea and much of eastern Indonesia, the Philippines, and the western Pacific to southern Taiwan (Greer 1970a). The genera Cryptoblepharus and Lipinia, which are widespread in Southeast Asia, are each represented in Papua by a small number of mostly endemic species.

................. 16157$

CH39

02-08-07 10:44:00

PS

PAGE 577

578 /

a

b

c

d

e

f

Figure 4.6.7. Scincidae: a. Tribolonotus novaeguineae; b. Tiliqua gigas; c. Lamprolepis smargadina; d. Eugongylus unilineatus; e. Emoia longicauda; f. Sphenomorphus jobiensis. Both are also represented throughout Oceania by single wide-ranging species, Cryptoblepharus poecilopleurus and Lipinia noctua, respectively, but only L. noctua reaches Papua. Cryptoblepharus also occurs in Australia where there is a small radiation of species (Cogger 2000). Glaphyromorphus is represented in Papua by two species, G. crassicaudis and G. nigricaudis, both of which also occur in Papua New Guinea, the Torres Strait islands, and northern Australia. Other members of the genus are confined to Australia or to Timor and associated islands (Greer 1990). There are two species of Eugongylus (Figure 4.6.7d) reported from Papua.

................. 16157$

CH39

02-08-07 10:44:11

PS

PAGE 578

Herpetofauna of Papua / 579

One is a widespread species, E. rufescens, which occurs from northern Australia and the Solomon Islands across New Guinea to eastern Indonesia. The other, E. unilineatus, is endemic to the north coast of New Guinea. Other members of this small genus occur from Palau and the western Pacific to the Solomon Islands (Greer 1970b). The remaining two genera, Emoia (Figure 4.6.7e) and Sphenomorphus (Figure 4.6.7f), are both large ( 70 species), with at least half of the species occurring in the New Guinea region where there have been extensive radiations (Brown 1991; Greer 1970b). Emoia also occurs throughout the islands of Oceania, with many endemic species (Brown 1991), but barely reaches Australia, where it is represented by two widespread species, Emoia atrocostata, which inhabits coastal habitats over a large part of the Indo-Pacific region, and E. longicauda (Figure 4.6.7d), which inhabits lowlands throughout New Guinea, the Admiralty Archipelago, and Mussau Island in the Bismarck Archipelago (Brown 1991; Cogger 2000). Only ten of the 75 currently recognized species of Emoia are found outside the New Guinea–Pacific Islands region (Brown 1991; Ineich and Zug 1991; How et al. 1998; Zug and Ineich 1995). Two of these are endemic to the Lesser Sundas, two are endemic to Southeast Asia, one is endemic to northern Maluku, and one is endemic to Christmas Island. An additional species, E. reimschisseli, is largely restricted to northern Maluku but has recently been reported from the Kai and Aru islands (How et al. 1998), and another, E. sorex, is endemic to Sulawesi, Maluku, and Batanta Island off the western tip of New Guinea. The two remaining species have enormous ranges and include E. atrocostata, mentioned above, and E. caeruleocauda, which is found in Southeast Asia, New Guinea, and large parts of the western Pacific (Brown 1991). Sphenomorphus is largely confined to Southeast Asia and the New Guinea-Solomon Islands region. Both of these large genera likely contain a number of different lineages and almost certainly will be split into smaller genera, many of which are likely to be endemic to New Guinea, as the species groups become better understood. Emoia and Sphenomorphus are the two largest lizard genera in Papua with 28 and 20 species, respectively. There are 12 currently recognized species of Varanus in Papua and the Aru Islands, but only five of these are endemic (Bo¨hme 2003; Pianka, King, and King 2004; Chapter 4.7). The endemic species are all related to Varanus prasinus (Figure 4.7.6) and include: V. beccarii, which is endemic to the Aru Islands and is often treated as a subspecies of V. prasinus; V. kordensis, which is restricted to Biak; V. boehmei, recently described from Waigeo (Jacobs 2003); V. macraei, currently known only from Batanta Island (Bo¨hme and Jacobs 2001); and V. reisingeri, which is endemic to Misool (Eidenmu¨ller and Wicker 2005). The remaining species are all shared with Papua New Guinea and at least four of these also occur in mainland Australia: V. similis and V. panoptes, which are both found in savannas and open areas in northern Australia and southern New Guinea (Bo¨hme 1988, 1991, 2001); V. doreanus, which inhabits lowland forest in far northeastern Australia and throughout much of New Guinea (Bo¨hme, Horn, and Ziegler 1994); and V. indicus (Figure 4.7.3), which occurs from the coastal Northern Territory and

................. 16157$

CH39

02-08-07 10:44:12

PS

PAGE 579

580 /

northeastern Queensland, the Bismarck and Solomon archipelagos, across New Guinea west to many of the islands in Nusa Tenggara and Maluku. V. indicus is also found in the Caroline, Mariana, and Marshall islands in the Pacific, although some of these populations may represent human introductions or separate species. A sixth species, V. prasinus, reaches the high islands of the Torres Strait, and a closely related species, V. keithhornei, is found on the Cape York Peninsula. Varanus prasinus is also shared with Papua New Guinea, as are V. jobiensis (Figures 4.7.4,5) and V. salvadorii (Figure 4.7.7). All three species are widely distributed in New Guinea, although V. salvadorii appears to be absent from the eastern peninsula. Varanus finschi, which has been reported from the Kai Islands (from which it is commonly available in the pet trade), the Bismarck Archipelago, mainland Papua New Guinea, and northern Australia, can be expected to occur in Papua (Ziegler, Philipp, and Bo¨hme 1999) but has not yet been collected there. Varanids occur from Africa across Asia to New Guinea and Australia. Nearly two-thirds of the species occur in Australia. However, fossil evidence suggests that the family probably originated in the Northern Hemisphere, although it has been in Australia since at least the Miocene. Most of the New Guinean forms are probably most closely related to taxa in Asia while those restricted to the south coast represent an expansion of the Australian radiation (Jennings and Pianka 2004).

There are eight families of snakes found in New Guinea, including the Cylindrophiidae, represented by Cylindrophis aruensis, which is endemic to the Aru Islands (McDowell 1975). Eighty-two species of snakes occur in Papua and six of these are endemic. Six of the eight families (all except the Boidae and Cylindrophiidae) are represented in Australia and seven of the families (all except the Boidae) occur in Southeast Asia. Most families appear to have had a long history in the Indo-Australian region. As with the lizards, the zoogeographic affinities of the various taxa are complex. The snake family Typhlopidae (burrowing snakes) includes about 200 species found throughout tropical areas of the world. Typhlopids are well represented in Australasia (McDiarmid, Campbell, and Touraˆe 1999). They are, however, on account of their fossorial nature, more poorly known than most snakes. They appear to have had a long history in the Australia-New Guinea region, as evidenced by the large number of species present and by their extensive distribution throughout this region (Cogger 2000). The earliest fossil from Australia dates to late Oligocene or early Miocene (Archer, Hand, and Godthelp 1991). Ten species of burrowing snakes in two genera, Ramphotyphlops (Figure 4.6.8) and Typhlops, are known from Papua; three of the species (all in the genus Ramphotyphlops) are endemic. The rest are mainly shared with Papua New Guinea or Australia. As a group, the Papuan species probably have their closest affinities with other species and species groups occurring in Southeast Asia. One species, Ramphotyphlops braminus, is widespread in the Indo-Pacific region and has been introduced by human transport to many tropical areas throughout the world.

................. 16157$

CH39

02-08-07 10:44:12

PS

PAGE 580

Herpetofauna of Papua / 581

Figure 4.6.8. Typhlopidae: Ramphotyphlops braminus. The family Cylindrophiidae (pipesnakes) includes about ten species of primitive snakes, all in the genus Cylindrophis, which occur in southern and Southeast Asia. One species, C. aruensis, is endemic to the Aru Islands (McDowell 1975) and is thought to be closely related to C. boulengeri from Wetar (McDowell 1975). These species may have originated from a common ancestor in eastern Indonesia. Pythons (family Pythonidae) are thought to have originated in Australasia and may well have originated in Australia (Greer 1997). The six species found in Papua are widespread in New Guinea (Figure 4.6.9). All are shared with Papua New Guinea and four of these are also shared with Australia: Leiopython albertisii, Morelia amethistina, M. spilota, and M. viridis. The other two, Apodora papuana (Figure 4.6.9a) and Morelia boeleni (Figure 4.6.9b), are endemic to New Guinea. The generic classification of Australian and New Guinea pythons is somewhat controversial and two recent revisions (Underwood and Stimson 1990; Kluge 1993) reached very different conclusions. More recently Harvey et al. (2000) revised the Morelia amethistina species complex, recognizing populations from Australia, New

a

b

Figure 4.6.9. Pythonidae: a. Apodora papuana; b. Morelia boeleni.

................. 16157$

CH39

02-08-07 10:45:40

PS

PAGE 581

582 /

Guinea, Ambon/Seram, Tanimbar, and Halmahera as full species. Rawlings, Barker, and Donnellan (2004) demonstrated that Apodora is a sister taxon to the Australia-New Guinea lineage of Liasis. There are three species of boid snakes (family Boidae) in New Guinea, all in the genus Candoia (Smith et al. 2001; Figure 4.6.10). Two of these, C. carinata and C. aspera, occur in Papua. Candoia carinata occurs in the Sangi Archipelago (between Mindanao and Halmahera), on scattered islands in southern Maluku and throughout much of New Guinea, and the Admiralty and Bismarck archipelagos. Candoia aspera is found throughout New Guinea and its satellite islands as well as the Bismarck and Admiralty archipelagos. A third species, Candoia paulsoni, has a rather puzzling distribution and is found throughout the Solomon Islands, islands in Milne Bay Province and much of mainland Papua New Guinea, and then disjunctly in Halmahera and northern Sulawesi, but has not been reported from Papua. The two remaining species of Candoia are extralimital to New Guinea (Smith et al. 2001): C. superciliosa is restricted to Palau, and C. bibroni occurs throughout much of the western Pacific from the Solomon Islands to Samoa. Candoia bibroni is regarded as the most basal taxon in the genus (Austin 2000). Boids otherwise occur in the neotropical region and in Madagascar and adjacent islands in the Indian Ocean. The origin of New Guinea boids is obscure but they may be most closely related to those in Madagascar (Burbrink 2005). There are two species of file snakes, family Acrochordidae, found in Papua (McDowell 1979). Both species are widely distributed in the New Guinea region. Acrochordus granulatus is found in marine and estuarine habitats from India and Southeast Asia to the Solomon Islands and northern Australia. Acrochordus arafurae (Figure 4.6.11) is found in lakes, swamps, and slow-moving rivers through-

Figure 4.6.10. Boidae: Candoia aspera.

................. 16157$

CH39

02-08-07 10:46:36

PS

PAGE 582

Herpetofauna of Papua / 583

Figure 4.6.11. Acrochordidae: Acrochordus arafurae. out much of southern New Guinea and northern Australia. It is closely related to A. javanicus, which occurs in similar habitats throughout much of Southeast Asia. The Colubridae includes a rather diverse group of advanced snakes found throughout the world. Most species are harmless but a few are venomous and have grooved rear teeth for venom delivery. The absence of hollow fangs is the sole characteristic that all colubrids share and there is little consensus on the evolutionary relationships of the various genera. The group is almost certainly not monophyletic but is used in the traditional sense here for convenience and includes aquatic marine and brackish water snakes (subfamily Homalopsinae) and freshwater snakes in the genus Tropidonophis (subfamily Natricinae). The colubrids are the largest group of snakes in New Guinea with about 34 species in some 11 genera (Figure 4.6.12). About half these genera are members of the subfamily Homalopsinae whose members are widespread in Australasia and are generally found in swampy, often estuarine habitats. Those found in Papua

a

b

Figure 4.6.12. Colubridae: a. Tropidonophis picturatus; b. Fordonia leucobalia.

................. 16157$

CH39

02-08-07 10:47:15

PS

PAGE 583

584 /

include Cerberus rynchops, Enhydris polylepis, Fordonia leucobalia (Figure 4.6.12a), and Myron richardsoni, all of which occur in Australia, and Cantoria annulata and Heurnia ventromaculata which are both endemic to New Guinea (Gyi 1970; Voris et al. 2002). Heurnia ventromaculata is endemic to Papua and is known only from a single specimen collected in the lower Mamberamo River. All homalopsine snakes have enlarged, grooved teeth in the rear of the mouth and a well developed venom gland. They also all have a vertical pupil and appear to be nocturnal. The group occurs in south and Southeast Asia across Indonesia to southern New Guinea and northern Australia. The majority of species and much of the group’s diversity is found in Southeast Asia and it is likely that the group originated there. The genus Brachyorrhos, which some include within the Homalopsinae (McDowell 1987), is represented in Papua by two species: B. albus, which occurs from Timor throughout much of Maluku to New Guinea, and B. jobiensis which is endemic to Yapen Island. Brachyorrhos albus is known to be terrestrial (Voris et al. 2002), whereas all other homalopsines are aquatic, and was relegated to incertae sedis (of uncertain position) within the Homalopsinae by Zaher (1999). Its phylogenetic relationships are unclear. There are only four terrestrial or freshwater colubrid genera in New Guinea: the colubrines Boiga, Dendrelaphis, Stegonotus, and the natricine Tropidonophis (Figure 4.6.12b). These same four genera occur in Australia, where they are generally restricted to northern and eastern seaboards. This suggests that these species may be relatively recent arrivals from New Guinea, although they are represented in Australia as fossils in the Oligocene (Rage 1987). There is a colubrid fossil from the late Eocene in Southeast Asia and its presence is consistent with the origin of the group on the Asian continent (Rage et al. 1992). However, fossils of similar age, or perhaps slightly older, have recently been found in North America (Parmley and Holman 2003). In any case, the aforementioned colubrid genera are all well represented in Southeast Asia (in den Bosch 1985; David and Vogel 1996) and it is quite possible that the New Guinea lineages originated from there, but there is no definitive evidence of this. All except Boiga have speciated in New Guinea and are each represented there by a small to modest number of species (McDowell 1972, 1984; Malnate and Underwood 1988). The main areas of Boiga diversity are Southeast Asia, south Asia, and Africa (Tweedie 1983; Greer 1997). The genus Dendrelaphis ranges from India through Southeast Asia and New Guinea to the Solomon Islands and Australia. At least five species occur in New Guinea (McDowell 1984; O’Shea 1996) and are found in Papua. However, these species are poorly diagnosed and the group is in need of taxonomic revision. If, as some suggest, the group is related to Chrysopelea from Southeast Asia (Underwood 1967), it is likely that New Guinea lineages of Dendrelaphis originated from there. There are approximately 11 species currently included in Stegonotus and they range from Borneo and the southern Philippines to New Guinea (including the Bismarck Archipelago) and northern Australia. There are four species known from Papua. All of these are shared with Papua New Guinea and two of them, S. cuculla-

................. 16157$

CH39

02-08-07 10:47:15

PS

PAGE 584

Herpetofauna of Papua / 585

tus and S. parvus, also occur in Australia. Stegonotus is thought to be related to Dinodon from Southeast Asia (Underwood 1967; McDowell 1972) but the deeper phylogenetic relationships are unknown, as are the relationships among the species of Stegonotus. The group’s origin is therefore unclear. Malnate and Underwood (1988) recognized 17 species of Tropidonophis ranging from the southern Philippines through parts of Maluku to New Guinea, the Bismarck Archipelago, and northern Australia. Kraus and Allison (2004) recently described an additional species from the D’Entrecasteaux Archipelago off eastern New Guinea. Malnate and Underwood speculate that the group ‘‘diverged from SE Asian stock’’ across the Sunda Shelf to reach the Philippines and New Guinea. A total of 13 species has been reported from New Guinea and ten of these occur in Papua. The family Elapidae, as used here, includes the sea kraits (subfamily Laticaudinae), sea snakes (subfamily Hydrophiinae), and the terrestrial subfamily Elapinae. These three subfamilies have often been treated as full families (Greer 1997). There are nine genera and 15 species of elapine snakes known from Papua (Figure 4.6.13). Four of these genera, Demansia, Furina, Oxyuranus, and Rhinoplocephalus are each represented by single species that is restricted to southern New Guinea and northern Australia. The genus Pseudechis, as currently understood, appears to be represented by two species endemic to New Guinea and closely related to species in Australia (O’Shea 1996; Hoser 2000; Wu¨ster et al. 2005; Kuch

a

b

Figure 4.6.13. Elapidae: a. Acanthophis laevis; b. Micropechis ikaheka; c. Laticauda colubrine.

c

................. 16157$

CH39

02-08-07 10:47:20

PS

PAGE 585

586 /

et al. 2005). The death adders, genus Acanthophis (Figure 4.6.13a), are represented by two species in New Guinea: Acanthophis rugosus, which occurs in southern New Guinea and northern Australia, and Acanthophis laevis, which is widespread in New Guinea. Several additional species are endemic to Australia. The genus extends to Seram and is badly in need of taxonomic revision. The remaining genera, Aspidomorphus, Toxicocalamus, and Micropechis (Figure 4.6.13b), are all endemic to New Guinea. Micropechis is represented by a single, widespread species, M. ikaheka, which ranges from the lowlands up to about 1,500 m in the mountains (O’Shea 1996). The other two genera have undergone modest species radiations. There are currently three recognized species of Aspidomorphus (McDowell 1967), two of which occur in Papua (but are not endemic). This group is also in need of taxonomic revision and it is likely that additional species will be recognized. Toxicocalamus currently has nine recognized species, four of which occur in Papua, where T. grandis is endemic (McDowell 1969). As with Aspidomorphus this genus is in need of taxonomic revision and it is likely that the number of species will increase with further study. The sea kraits, subfamily Laticaudinae, include two species found in Papua, Laticauda colubrina (Figure 4.6.13c) and L. laticauda. Both species are widespread in the tropical seas of the Indo-Pacific (Heatwole 1999). Within the sea snakes (subfamily Hydrophiinae) there are 11 genera and 16 species found in Papua. Most of the species are widely distributed in the IndoAustralian region and none is endemic to Papua (Heatwole 1999). See Tomascik (1997) for a detailed treatment of this group in Indonesian waters.

Only three introduced species of amphibians have been reported from New Guinea. In addition, the Tokay Gecko (Gekko gecko) has been reported from the Aru Islands, probably representing a human introduction. Two of the introduced amphibians are toads, Bufo melanostictus and B. marinus, and one is a ranid frog, Limnonectes cancrivorus. All except Bufo marinus are known with certainty to occur in Papua. Bufo melanostictus is a common toad found throughout much of southern and southeastern Asia. It seems at present to have a limited distribution in Papua, where it is reported to occur in coastal areas around Manokwari (Menzies and Tapilatu 2000). It was probably introduced by transmigration settlers from nearby Warmare. This toad can be expected to spread to other areas of the province, aided by human transport. Phillips, Brown, and Shine (2003) have demonstrated that Bufo marinus has been responsible in Australia for population declines of frog-eating snakes and is widely thought to have similarly impacted some Papua New Guinea snakes such as the Papuan Black (Pseudechis papuanus). It is not known if Bufo melanostictus is having a similar impact in Papua. Limnonectes cancrivorus is native to Southeast Asia and has been introduced to the Philippines. It was documented from New Guinea by Menzies (1992, 1996),

................. 16157$

CH39

02-08-07 10:47:20

PS

PAGE 586

Herpetofauna of Papua / 587

who reported it from the Sorong and Jayapura areas in Papua. This species is something of a culinary delicacy in Southeast Asia and is often farmed there. It was presumably introduced to New Guinea as a food item.

Biology and Ecology of the Herpetofauna The frogs in Papua are found from the lowlands to alpine grassland above 3,200 m. Species richness varies geographically in relation to climate and geological history, but is generally highest in hill forest and upland forests (Allison, Kraus, and McShane 2004). Those known from Papua are typical of New Guinea frogs generally and include species that, when active, climb into vegetation (scansorial), remain on the forest floor (terrestrial), or live in burrows or cavities within the soil (fossorial). They range in size from ca 11.5 mm in Oreophryne minuta (Richards and Iskandar 2000) to more than 250 mm in Rana arfaki (Tyler 1976) and vary considerably in body form. Species that are primarily fossorial tend to have pointed, wedge-shaped snouts, wide bodies, small eyes, and short legs. Scansorial frogs mostly have relatively long legs, broad heads, large eyes, and adhesive terminal disks on the fingers and toes. Species that are primarily active on the forest floor tend to resemble scansorial species but are often more heavy-bodied, have shorter legs, and lack terminal disks on the fingers and toes. These various differences tend to cut across taxonomic groups, although all New Guinea myobatrachids are surface dwellers, as are most ranids. Most hylids are scansorial (Figure 4.6.2), although a few species, such as Litoria nasuta, occur on the ground along the banks of lakes and streams. Insofar as is known, no New Guinea hylids are fossorial. The microhylids are the most morphologically diverse group of New Guinea frogs. Most species are scansorial (Figure 4.6.3a) but a sizable number are surface dwellers or fossorial (Figure 4.6.3b). It is common to find such differences within a single frog genus. For example, most species of Callulops are surface dwellers or fossorial but Callulops slatteri is scansorial and highly arboreal. In something of an extreme case, almost all known species of Xenorhina are fossorial but X. arboricola (Figure 4.6.3c) lives high in the trees in the leaf litter that accumulates within epiphytic vegetation (Allison and Kraus 2000). This species has a pointed snout representative of fossorial frogs but has long legs and terminal toe and finger disks characteristic of scansorial frogs. Most New Guinea frogs are nocturnal and are most active during the first few hours of darkness, as judged by calling activity. A few species—primarily those that burrow—call mostly during the day. Some species (e.g., Copiula spp.), are crepuscular (active at dawn and dusk). Scansorial frogs generally shelter during the day in leaf litter and ascend into vegetation to call at night. A few species of microhylids and most hylids are entirely arboreal, although most arboreal hylids descend to the ground at some point to breed. Most species of New Guinea frogs call and breed throughout the year during periods of wet weather. However, a few species inhabiting savanna regions of

................. 16157$

CH39

02-08-07 10:47:20

PS

PAGE 587

588 /

southern New Guinea, where rainfall is highly seasonal, call and breed only during the wet season. All species of hylids have an aquatic tadpole stage. In most cases the frogs mate in water—in streams, rivers, lakes, or temporary pools, depending on the species. Species that breed in stagnant water generally have eggs that are pigmented and which they attach to aquatic vegetation or leave floating at the surface. Species inhabiting fast-moving streams (‘‘torrent dwellers’’), such as most species of Nyctimystes, attach their eggs to rocks and the tadpoles have sucker-like mouths with which to anchor themselves to stream substrate. The eggs of such species are generally unpigmented. In some hylid frogs, such as Litoria iris and allied species (Menzies 1976; Tyler 1976), females lay their eggs in small clumps with a protective layer of clear jelly. These are attached to leaves hanging over water and when the tadpoles hatch some two weeks later they drop into the water. All of the myobatrachid frogs from Papua have an aquatic tadpole stage. Four of these frogs (three species of Lechriodus, and Limnodynastes convexiusculus) produce small foam nests in shallow depressions at the edges of swamps. Females deposit the eggs into the nests, where they are fertilized by the male. The tadpoles depart for open water after they hatch. In Lechriodus fletcheri from Australia the tadpoles are known to be cannibalistic and this may very well be the case for the New Guinea species of Lechriodus (Menzies 1976). The only other myobatrachid frog occurring in Papua, Crinia remota, also inhabits northern Australia and is known to breed there in shallow pools and low-lying flooded areas (Cogger 2000). Ranid frogs practice two different reproductive modes. All species of New Guinea Rana lay their eggs in streams or lakes, in bunches or strands attached to aquatic vegetation. In contrast, species of Platymantis are terrestrial breeders and deposit their eggs within moist leaf litter or arboreal leaf axils. The eggs develop into small froglets without going through an aquatic tadpole stage (Menzies 1976, 1987). The breeding biology of all known species of New Guinea microhylids is similar to that of Platymantis, although some arboreal microhylids oviposit on the undersides of leaves in the trees that they inhabit and at least two species, Oreophryne anthonyi and Cophixalus riparius, oviposit in epiphytic ant plants (genus Hydnophytum). Other microhylids, primarily species in the genus Oreophryne, oviposit within the leaf axils of Pandanus, palms, and bananas, or within the hollow stems of bamboo. In the few species of microhylids that have been well studied, the male (rarely the female) guards the eggs until they hatch and, at least in some species, transports the froglets on his back until they can survive on their own—probably within a few days (Cogger 1964; Bickford 2004; Gu¨nther, Kapisa, and Tetzlaff 2001). Bickford (2004) showed that desiccation was the primary source of mortality for arboreal Oreophryne frog eggs, that predation was the main source of egg loss in terrestrial Hylophorbus, and that parental care reduced egg mortality in both cases. The food habits of New Guinea frogs are virtually unstudied. Most species are believed to be opportunistic feeders that take prey of appropriate size. Many spe-

................. 16157$

CH39

02-08-07 10:47:21

PS

PAGE 588

Herpetofauna of Papua / 589

cies of burrowing frogs apparently feed largely on earthworms (Zweifel 1972a). Some of the larger species, such as Asterophrys turpicola, are known at least occasionally to take small lizards, while Rana arfaki feeds on crabs and prawns (Zweifel and Tyler 1982). Tyler (1962) found aquatic snails and insects in stomachs of Rana grisea, and arthropods, mostly insects, in the stomachs of Litoria darlingtoni inhabiting a coffee plantation in the highlands of Papua New Guinea. Microhylids elsewhere are known to have a preference for ants (Mendes, Pinheiro, and Ruas 1994). There have been relatively few ecological studies of New Guinea frogs and none based in Papua. Allison and Dwiyahreni (1998) collected frogs along an altitudinal transect in the PT Freeport Contract of Work area (lowlands and mountains adjacent to Mt Jaya). These surveys were conducted in lowland, lower montane, montane, and subalpine habitats and included about 40 species of frogs. However, the survey coincided with a localized drought, included only cursory work in lower montane areas, and almost certainly undersampled the frogs. A total of 21 species was recorded from the lowlands, where considerable habitat diversity was sampled, and 14 species were reported from montane regions where habitat diversity was comparatively limited. This study, and a similar study of the Wapoga River area in northern Papua (Richards, Iskandar, and Allison 2000), demonstrated that ranid frogs are limited to lowland and lower montane sites, where more than five species may occur. This is also generally true of myobatrachids, although Lechriodus aganoposis is known to occur to elevations of 1,500 m (Zweifel 1972b) and Lechriodus platyceps has been reported as high as 1,890 m (Richards, Iskandar, and Allison 2000). These surveys, together with an additional survey of the Cyclops Mountains and southern Mamberamo Basin (Richards, Iskandar, and Tjaturadi 2002), demonstrate that there are generally at least four or five species of hylids and microhylids at lowland and hill forest sites and more than ten species at montane localities. Microhylids are generally the only frogs present in upper montane and subalpine habitats where permanent water is absent and more than five species may occur in mossy forest above 2,500 m elevation. These observations are in rough agreement with a preliminary analysis of overall species richness, which demonstrates that the richest region of New Guinea, the southern highlands of Papua New Guinea, has more than 35–38 species of frogs (Allison, Kraus, and McShane 2004). Richness is nearly as high along the central mountains and in the north coast ranges. The Lakekamu Basin in southern Papua New Guinea, probably the most intensively surveyed site in New Guinea for amphibians, produced only 30 species of frogs. These figures are far lower than sites in South America. For example, Duellman (1978) found 81 species of amphibians at Santa Cecilia, a lowland rain forest site in Amazonian Ecuador. However, species turnover (beta diversity) may be higher in New Guinea than in the neotropics (Allison, in prep.) and this may help to explain why overall species richness of frogs in New Guinea is so high.

................. 16157$

CH39

02-08-07 10:47:21

PS

PAGE 589

590 /

The various species of chelid turtles are generally found in swamps and slowmoving watercourses. Emydura subglobosa, probably the most common species, may be found in small streams but also in some of the larger rivers. Chelodina siebenrocki is often found in estuarine areas but most species of chelid turtles are found in freshwater; it is common to find several species in the same locality. These turtles tend to have fairly general diets and feed on fish, crayfish, invertebrates, and occasional vegetable matter. Most species reach a maximum size of 20–30 cm in carapace length (females tend to be slightly larger than males). Reproduction tends to be highly seasonal, coinciding with the beginning of the wet season, and females generally produce about ten eggs per clutch. Carettochelys insculpta is currently under study in Papua New Guinea and has been extensively studied in Australia where it is restricted to a few rivers in Arnhemland (Georges et al. 2000). Adults commonly reach 50 cm in carapace length and large individuals can reach 70 cm (Iskandar 2000). In Australia it is found in pools with an average depth of 2 m, sand or gravel bottoms, an accumulation of leaf litter and fallen trees, undercut banks, exposed tree roots, and other features that provide an abundance of refuges (Georges et al. 2000). Their habitat preferences in New Guinea are similar but include grassy lagoons, swamps, lakes, and waterholes (Rose, Parker, and Rhodin 1982). They generally prefer fresh water but are sometimes found in estuaries. Carettochelys insculpta nests gregariously during the dry season, twice per year every second year, laying their eggs in sandbanks (Doody, Georges, and Young 2003). Clutch size varies from 7–19 eggs (Georges et al. 2000) but may be higher in the New Guinea population where Iskandar (2000) gives a figure of 17–40 eggs. Carettochelys insculpta is omnivorous and its diet includes more plant material (e.g., water weeds) than animal material (e.g., aquatic insect larvae, shrimp, crayfish, mussels, clams, and carrion; Georges et al. 2000). The two species of Pelochelys in New Guinea are the largest freshwater turtles in the region and among the largest freshwater turtles in the world, reaching carapace lengths of 1 m or more. They are found in large rivers with sandy or muddy bottoms and are thought to feed on frogs, fish, and aquatic invertebrates. Little is known of their reproductive behavior but they are thought to produce clutches of 20–30 eggs (Iskandar 2000). The distribution and biology of the marine turtles has been extensively discussed in Groombridge (1982) and Tomascik et al. (1997) and will only be briefly summarized here. Green Turtles (Chelonia mydas) can attain 1.2 m in carapace length and a body mass of 200 kg. They have a circumtropical distribution, with a number of discrete geographic populations that are highly migratory. These turtles generally inhabit relatively shallow coastal waters and nest mainly in tropical and subtropical regions. Green Turtles are probably the most common species of marine turtle in Indonesian waters. They are largely herbivorous, feeding on algae and eel grass. Females have an average clutch size of 110 eggs and may lay from three to seven clutches per nesting season. Females probably breed at two- to four-

................. 16157$

CH39

02-08-07 10:47:21

PS

PAGE 590

Herpetofauna of Papua / 591

year intervals. They and their eggs are actively harvested at many sites throughout their range and many populations, including those in Papua, are highly depleted. The Olive Ridley (Lepidochelys olivacea) is the smallest of the sea turtles. Adults reach a maximum carapace length of about 80 cm and a body mass of 50 kg. They occur throughout the tropical oceans of the world and tend to be found mainly in the open ocean. Olive Ridleys characteristically form large nesting aggregations. Females typically nest from one to three times per season and may nest every year. They have an average clutch size of around 110 eggs. This species has a rather broad diet that mainly includes crabs, shrimp, and rock lobsters, and also jellyfish, tunicates, and algae. The Hawksbill (Eretmochelys imbricata) is intermediate in size to the Ridley and Green Turtles, reaching a maximum carapace length of around 1 m and a body mass of about 140 kg. These turtles are closely associated with coral reefs and have a circumtropical distribution. They generally nest from two to four times a season at intervals of three years. Clutch size increases with the body size of the female but is generally around 140 eggs. Hawksbills feed mainly on sponges. The Leatherback or Leathery Turtle (Dermochelys coriacea) is the largest of the marine turtles. Adults can reach a carapace length of nearly 2.5 m and a body mass exceeding 900 kg, although most individuals are smaller than this. They have a global distribution and are highly migratory. Leatherback Turtles nest on tropical beaches that are readily accessible from deep water and have been reported to nest in dense aggregations on beaches along the northern coast of the Vogelkop and on islands in Cenderawasih Bay, such as Yapen (Salm, Petocz, and Soehartono 1982). Females generally nest four to seven times per season, although some females may nest as many as 11 times. They are thought to nest at one- to three-year intervals. The average clutch size is 85 eggs. Leatherback Turtles feed primarily on jellyfish and planktonic tunicates. Globally, populations of the Leatherback are highly depleted. One radio-tracked specimen tagged on the Vogelkop was observed to visit the coast of California.

The Indo-Pacific or Saltwater Crocodile (Crocodylus porosus) is one of the largest and most dangerous crocodiles in the world, reaching a total length of 7 m (Ross 1989). It is generally found in coastal rivers and swamps, often in brackish water, as well as freshwater swamps and lakes. This species is also sometimes encountered in the open ocean (Jones 1909) and this probably accounts for its widespread distribution. Juveniles feed on aquatic invertebrates and small vertebrates; adults take a wide variety of vertebrate prey, particularly mammals, birds, and fish. During the wet season females construct a mound of rotting vegetation into which they deposit a clutch of 60–80 eggs. The female often protects the nest and has been reported in some cases to care for the young. The New Guinea Crocodile (Crocodylus novaeguineae) reaches a total length of about 3.3 m (Ross 1989). It generally inhabits freshwater rivers, swamps, and lakes and often co-occurs with the Indo-Pacific Crocodile. The southern populations of

................. 16157$

CH39

02-08-07 10:47:22

PS

PAGE 591

592 /

Crocodylus novaeguineae (which may represent a different species [Hall 1989]) generally nest during the wet season while the population north of the central ranges nests during the dry season. The nesting habits are otherwise similar to those of the Indo-Pacific Crocodile and involve the construction of a nest of rotting vegetation. This is often placed in a well-shaded place, such as at the base of a tree, and close to a water hole (Groombridge 1982). Females lay clutches of 27–45 eggs (Jelden 1981). New Guinea crocodiles are thought to feed primarily on fish, but they are opportunistic feeders and will also take aquatic invertebrates and vertebrates including frogs, snakes, and birds. Both species are widely exploited for their skins, and populations in many areas are known to be heavily depleted. Papua New Guinea has strictly regulated crocodile harvesting since the late 1960s and has also developed captive rearing programs. Similar programs have recently been developed in Papua.

Information on the biology and ecology of New Guinea reptiles is mostly available only from anecdotal reports embedded in the taxonomic literature. Most species have scarcely been mentioned since they were originally described, although the varanid lizards have recently been studied in some detail by Kai Philipp and colleagues (e.g., Philipp 1999; Philipp, Bo¨hme, and Ziegler 1999; Chapter 4.7). The biology of many lizard species, particularly those occurring in southern New Guinea, can be inferred from studies of these or closely related taxa in northern Australia. Information on Australian lizards is summarized in Greer (1989). The sole representative of the family Dibamidae, Dibamus novaeguineae, as is true of other species of Dibamus, probably inhabits leaf litter. It likely feeds on small arthropods. All species of New Guinea agamids are arboreal, although some perch on rocks along watercourses and others sometimes descend to the forest floor to forage. Hydrosaurus amboinensis, a large, distinctive species that is widely distributed in Indonesia, is generally restricted to rainforest riparian habitat in Papua. It eats both plant and animal material (e.g., arthropods, small mammals). Chlamydosaurus kingii, which is found in seasonal dry forest in southern New Guinea and northern Australia, feeds largely on Lepidoptera larvae, termites, and other small arthropods in Australia (Shine and Lambeck 1989; Griffiths and Christian 1996). Both Hydrosaurus and Chlamydosaurus, which can attain a meter or more in total length, have average clutch sizes of 7–8 eggs, although Chlamydosaurus in Australia has been reported to produce clutches of a many as 14 eggs (Bedford, Christian, and Griffiths 1993). In the genus Hypsilurus, studies of the two endemic Australian species have shown that they produce clutches of more than five to seven eggs and feed largely on insects (Shea 1991). The eight species from Papua likely have similar habits but some include considerable plant material in their diet (Allison, in prep). The biology and ecology of the 26 species of geckos known from Papua is likely to be similar to congeneric species from northern Australia, Papua New Guinea,

................. 16157$

CH39

02-08-07 10:47:22

PS

PAGE 592

Herpetofauna of Papua / 593

and Indonesia. All except some species of Nactus, which are usually found on the forest floor, appear to be arboreal. All are known to have a clutch size of two but probably produce multiple clutches per year. Many species oviposit into communal nests. All New Guinea geckos are ‘‘sit and wait’’ predators and most are insectivorous, although the larger species, such as Gekko vittatus (Figure 4.6.6b) and certain species of Cyrtodactylus, are also known to feed on other geckos. All species of geckos in Papua are nocturnal, although some individuals may occasionally call during the day. Several species of geckos, including Cosymbotus platyurus, Hemidactylus frenatus, Lepidodactylus lugubris, and Hemiphyllodactylus typus, are human commensals that have been transported by people throughout the Indo-Australian archipelago and beyond. Hemidactylus frenatus in particular is a common inhabitant of dwellings in lowland Papua. Most species of geckos produce territorial vocalizations and the chirping of Hemidactylus frenatus is a common feature of life in the IndoPacific region. In the family Pygopodidae most species are thought to be crepuscular or nocturnal but Lialis burtonis appears to also be active during parts of the day. It is a ‘‘sit and wait’’ predator that appears to prey entirely on small skinks (Patchell and Shine 1986a). The life history of Lialis jicari is unknown but it is probably similar to that of L. burtonis. As is true in geckos, many pygopodids, including Lialis, vocalize. The clutch size is generally two, but occasionally includes three eggs (Patchell and Shine 1986b). The skinks are the largest and most diverse group of lizards in the world with more than 1,300 currently recognized species. They are also the largest and most diverse group of lizards in New Guinea. They range in snout-vent length (SVL) from 25–30 mm to more than 30 cm in Tiliqua gigas and Sphenomorphus muelleri. Some species, such as Prasinohaema spp. and Lamprolepis smargadina, are highly arboreal but most are terrestrial and some, primarily species of Sphenomorphus, are fossorial. Most species are actively hunting predators on insects and small arthropods. Some of the larger species (e.g., Eugongylus spp.) are predators on other lizards. Many species are heliothermic and are commonly observed basking but some, such as species of Eugongylus, are crepuscular and others, such as the members of the Sphenomorphus jobiensis complex, which have particularly large eyes, appear to be partially nocturnal. Skinks occur from lowland and hill forest areas, where species richness is highest, to alpine grasslands where only a single species may be present (Greer, Allison, and Cogger 2005). New Guinea skinks of the genus Tribolonotus produce a single egg that is guarded by the female who also tends to guard the neonate. Other groups—such as Emoia spp. and Papuascincus spp.—produce a consistent clutch size of two eggs, but in many other New Guinea skinks the clutch size is variable and increases with body size. For example, the litter size in Tiliqua gigas evanescens ranges from 4 to 13 (Shea 2000). A number of species, particularly those from higher elevations such as species of Lobulia, produce live young (Greer, Allison,

................. 16157$

CH39

02-08-07 10:47:22

PS

PAGE 593

594 /

and Cogger 2005) although the live-bearing reproductive mode is also sometimes found in lowland lizards (e.g., Tiliqua gigas). The varanid lizards, together with the crocodiles, are the largest terrestrial predators in New Guinea and one species, Varanus salvadorii, had been documented to reach 2.5 m in total length (Horn 2004) and it is rumored that some individuals may exceed four meters. All species in Papua are scavengers and carnivores. Most will ascend trees when alarmed and some, such as Varanus prasinus, are almost exclusively arboreal (Greene 1986). Clutch size is variable and ranges from two to four eggs in some of the smaller species such as Varanus prasinus and its closely related congeners, to a dozen or so eggs in the larger species such as Varanus salvadorii (Horn 2004). See Chapter 4.7 for further details.

The various typhlopid species vary widely in length, girth, and head shape but are mostly small (total length mostly 30 cm or less) with minute cycloid body scales and very reduced eyes. They tend to be tan to dark brown-black in dorsal coloration and ventrally are white to tan or the same color as the dorsum. In the Australian species the females tend to be larger than the males (Shine and Webb 1990). Most species are assumed to be fossorial but are often found above ground on wet nights. Observations from the Solomon Islands and the Philippines demonstrate that some species of Ramphotyphlops ascend into trees (McCoy 1980; Gaulke 1995; Das and Wallach 1998) as does Typhlops inornatus in Papua New Guinea (F. Kraus, pers. comm.). Most Australian species of typhlopids are thought to feed on the larvae and pupae of ants and termites (Webb and Shine, 1993). Indeed, Webb and Shine (1992) showed that in Australia Ramphotyphlops nigrescens is able to follow ant trails. Webb and Shine also reported (1993) that Acutotyphlops subocularis from Papua New Guinea (as Ramphotyphlops subocularis; see Wallach 1995) apparently feeds on earthworms. All New Guinea typhlopids are known or are thought to be oviparous (egg laying). In Ramphotyphlops braminus the clutch size varies with body size and ranges from one to eight eggs (Ota et al. 1991; Kamosawa and Ota 1996). This appears to be a hermaphroditic, all-female species (McDowell 1974; Nussbaum 1980). It is this trait together with human transport that likely accounts for its nearly pantropical distribution. Pipesnakes (family Cylindrophiidae) are burrowing or leaf litter-inhabiting snakes that are mostly nocturnal and are known to feed on invertebrates and small vertebrates. They are ovoviviparous (eggs hatch inside the female’s oviducts) and are reported to produce 2 to 12 living young (Zug, Vitt, and Caldwell 2001). The species that is endemic to the Aru Islands, Cylindrophis aruensis, is apparently known only from two immature specimens (McDowell 1975) and nothing appears to be known about its life history. Pythons are the largest snakes in New Guinea, with maximum recorded lengths of 5 m for Morelia amethistina (Greer 1997 and references therein). Although

................. 16157$

CH39

02-08-07 10:47:22

PS

PAGE 594

Herpetofauna of Papua / 595

secretive and difficult to study in the field, New Guinea pythons are well studied in captivity (Ross and Marzec 1990). Some species such as Apodora papuana seem to be entirely terrestrial (and often associated with water; O’Shea 1988), but other species, such as Leiopython albertisi and Morelia boeleni, appear to be at least partially arboreal. In Australia, the Carpet Python, Morelia spilota mcdowelli, spends 45% of its time in trees, ascending to heights of nearly 10 m (Shine and Fitzgerald 1996). Morelia amethistina has been reported to reach heights of 22 m above the ground (Zweifel in McDowell 1984) and Morelia boeleni reaches at least 10 m (Allison, pers. obs.). The Green Tree Python, Morelia viridis, is highly arboreal (pers. obs.). Most species appear to be primarily nocturnal but sometimes bask during the day. Studies in Australia have shown that they tend to have relatively small home ranges (Shine and Fitzgerald 1996) but that Liasis fuscus sometimes travel 500 m in a single night (Madsen and Shine 1996a, 1996b). Pythons are mostly ‘‘sit and wait’’ predators, but some species apparently actively search for prey (Shine 1991a). They feed entirely on vertebrates, and at least one species, Apodora papuana, is known to feed mainly on other snakes, including other pythons. All pythons are oviparous and insofar as is known, the females brood the eggs, generating heat by muscle contractions (Ross and Marzec 1990). Clutch size varies with the species and with the size of the female. It is usually around 10–20 eggs, but can be as high as 40 eggs in some species (Greer 1997). The two species of Candoia (family Boidae) known from Papua are both medium-sized snakes that generally are less than 1.5 m in total length. Candoia carinata is fairly slender and at least partially arboreal although it is most commonly found on the ground. Candoia aspera is much more heavy-bodied and is exclusively terrestrial. It occurs throughout New Guinea from lowlands to above 1,000 m (McDowell 1979). Harlow and Shine (1992) studied the food habits and reproductive biology of Candoia. The taxon that they treat as Candoia carinata has subsequently been divided into several species by Smith et al. (2001) and many of the specimens that Harlow and Shine (1992) examined from the Australian Museum are now referable to Candoia paulsoni. However, the biologies of Candoia paulsoni and Candoia carinata appear to be quite similar, and the overall conclusions that Harlow and Shine (1992) reached would apply to Candoia carinata as redefined by Smith et al. (2001). Females in both C. carinata and C. aspera reach a larger body size than do males. As is characteristic of boids, both are ovoviviparous, producing on average 15–18 young. Litter size is correlated with female body size. The diets of both species are similar and include frogs, reptiles (mostly skinks), and mammals (Harlow and Shine 1992). File snakes (family Acrochordidae) are almost entirely aquatic. The larger of the two species found in New Guinea, Acrochordus granulatus, reaches a maximum body length of about 2 m. These snakes are fairly heavy-bodied and a large individual may exceed 4 kg in body mass. Acrochordus arafurae tends to be slightly smaller than A. granulatus. The skin of both species is very rough to touch, proba-

................. 16157$

CH39

02-08-07 10:47:23

PS

PAGE 595

596 /

bly an adaptation for holding slippery prey. They have relatively small eyes and are notable in being relatively slow moving and having a very low metabolic rate. Acrochordus arafurae is found exclusively in fresh water such as inland swamps and slow-moving watercourses but will ascend into freshwater creeks. In contrast, Acrochordus granulatus occupies marine and brackish habitat such as mangroves and mudflats. Both species are active foragers and prey mainly on fish. They are sometimes extremely abundant and in Australia can reach population densities of 400/ha of surface water (Houston and Shine 1994). Each tuberculate head scale has more than seven microscopic sensillae (sense organs) and each of the keeled body scales has a single sensilla (Povel and van der Kooij 1997). These organs are thought to aid in the detection of prey in the murky water inhabited by these snakes. Observations of captive individuals have demonstrated that fish are often caught in body coils (Lillywhite 1996). Both species produce live young. Acrochordus arafurae matures in about three years and females, on average, reproduce every 3–4 years (Madsen and Shine 2001). Acrochordus arafurae has an average litter size of around 16, and A. granulatus produces an average of six young per litter (Greer 1997). Snakes of the family Colubridae have diverse life histories. Boiga is represented in New Guinea and Australia by a single species, B. irregularis. This snake is common to abundant throughout lowland New Guinea from sea level to above 1,600 m. It is highly arboreal and nocturnal. It is also relatively large for a colubrid, reaching lengths in excess of 3 m, with females becoming slightly larger than males (Shine 1991b). Boiga irregularis has a slender, laterally flattened body, ridged ventral scales, and a prehensile tail, all features that make it an extremely good climber. It feeds largely on vertebrates and their eggs and is an active hunter but will also sit and wait to ambush prey. Young snakes tend to feed largely on frogs and lizards but switch to birds and mammals as they increase in size. The species is oviparous and produces clutches that average 5–6 eggs. Larger females tend to produce larger clutches, with the maximum number reported being 11 (Greer 1997). In is interesting to note that Boiga irregularis became established in Guam subsequent to World War II, probably from the Admiralty Islands, and has subsequently been responsible for the extinction of most of the native forest birds and several species of native reptiles on Guam (Rodda and Fritts 1992). It may have recently become established on Rota and Saipan in the northern Mariana Islands (Fritts, McCoid, and Gomez 1999). All members of the genus Dendrelaphis are long, reaching more than 2.5 m in some species. They are slender, fast-moving snakes with large eyes, are mainly arboreal, and hunt by day. Dendrelaphis generally inhabits vegetation within a few meters of the ground, but D. calligastra in New Guinea has been found as high as 23 m in trees (Loveridge 1948). Dendrelaphis calligastra in Australia feeds largely on frogs and lizards while Dendrelaphis punctulata feeds largely on frogs (Greer 1997 and references therein). All Dendrelaphis are oviparous. In Australia D. calligastra produces clutches that average seven eggs, and D. punctulata on average

................. 16157$

CH39

02-08-07 10:47:23

PS

PAGE 596

Herpetofauna of Papua / 597

produces around nine eggs (Greer 1997 and references therein). Parker (1982) reports that D. calligastra in Western Province, Papua New Guinea, deposits its eggs ‘‘in hollows and cracks in tree trunks, bamboo stems, and dead branches.’’ The four species of New Guinea Stegonotus, all of which are found in Papua, are medium-sized terrestrial snakes that are widespread in New Guinea. Two of the species, Stegonotus parvus and S. cucullatus, also occur in Australia, where they have been studied by Shine (1991b). Most are slate gray in color but range from brown to black, although some populations of Stegonotus cucullatus are almost white. McDowell (1972) noted that S. cucullatus sometimes closely resembles the venomous elapid, Micropechis ikaheka, suggesting mimicry. Species of Stegonotus appear to be crepuscular and at least some species, such as S. cucullatus, are nocturnal. They feed largely on vertebrates, including frogs, lizards, mammals, and the eggs of lizards and snakes (Shine 1991b; McDowell 1984). They are presumably active hunters. Males tend to reach a slightly larger size than do females. All species are oviparous. Stegonotus cucullatus produces clutches of 7–16 eggs (Shine 1991b). Parker (1982) reported finding a clutch of two eggs of S. diehli, but this may not have included a complete clutch; clutch size in other species has not been reported. The ten species of Tropidonophis known from Papua range, in maximum length, from about 55 cm to 1.3 m. They occur from lowlands to about 2,200 m (Malnate and Underwood 1988). All of the species have strongly keeled scales and tend have dull coloration, but populations of at least one species, Tropidonophis doriae, from northwestern Papua New Guinea, are brightly banded with black and orange-red (O’Shea 1996). Some species appear to be diurnal while others appear to be nocturnal. Species of Tropidonophis often occur in close proximity to water, but some, such as T. statisticus, occur in montane habitats some distance from permanent water. Tropidonophis preys primarily on frogs (Malnate and Underwood 1988; Shine 1991b). They tend to be relatively common. All of the species are oviparous. Tropidonophis mairii has a variable clutch size of 2–18 (Malnate and Underwood 1988; Brown and Shine 2002; Shine 1991b), but most species average 5–6 eggs per clutch. All the species of homalopsine colubrids are mostly aquatic but will sometimes venture onto land. Enhydris polylepis is found primarily in fresh water and in southwestern Papua New Guinea is found well inland from the coast to elevations of nearly 400 m (Parker 1982). Fordonia leucobalia tends to be found in brackish water, such as pools in mudflats. Cerberus rynchops is found mainly in mangrove habitats and has an enormous distribution, presumably because it can readily disperse across open ocean. Myron richardsoni tends to be found in the same habitat and has been recorded more than 2 km off the coast (Parker 1982). Cantoria annulata was reported by Parker (1982) to occur in mangrove and nipa palm habitats. Fordonia leucobalia feeds mainly on crabs and other crustaceans. Myron richardsoni is thought to take crabs and fish (Voris and Murphy 2002). Cerberus rynchops feeds mainly on fish but will also take frogs (Mori 1998). Enhydris polylepis appears

................. 16157$

CH39

02-08-07 10:47:23

PS

PAGE 597

598 /

to feed on fish and frogs, a diet that is similar to that of other species of Enhydris from Southeast Asia (Greer 1997 and references therein; Voris and Murphy 2002). The prey preferences of Cantoria annulata are unknown. All species of homalopsines are live-bearing and the litter size is quite variable. Cerberus rynchops produces from 1–35 young; Enhydris polylepis produces 7–27 young and Fordonia leucobalia produces 3–17 (Greer 1997 and references therein). Shine (1991b) reported that a single female Myron richardsoni produced a litter of six. Parker (1982) mentioned that a gravid Cantoria annulata contained five eggs, suggesting that it may be ovoviviparous. Within the family Elapidae the two sea kraits, Laticauda colubrina and L. laticauda, both reach a maximum size of about 1.5 m. Although highly aquatic, with laterally compressed tails, they spend significant amounts of time on land and can sometimes be found in large numbers sheltering under debris along rocky sea coasts or basking among rocks. They are oviparous and lay their eggs on land. Sea kraits feed entirely in the sea, are mainly nocturnal, and appear to exclusively eat eels (Greer 1997; Tomascik et al. 1997; Heatwole 1999). The sea snakes (subfamily Hydrophiinae) range in length from nearly a meter to as much as two meters. They are generally restricted to shallow water habitats, but Pelamis platurus is strictly pelagic, although it mostly feeds near the sea surface. Some species are diurnal, others are nocturnal, and some appear to be both. Most species feed on fish and in some cases specialize on particular fish taxa, but a few species such as Aipysurus eydouxi and Emydocephalus annulatus specialize in eating fish eggs. All species produce living young. Most species have small litters of fewer than ten young but large females of some species such as Enhydrina schistosa may produce more than 30 young per litter (Voris and Jayne 1979). The elapine snakes form a diverse assemblage. Demansia vestigiata is a slender, fast-moving diurnal snake that inhabits savanna woodland and reaches a total length of about 1.5 m. It is oviparous and feeds mainly on lizards and frogs (O’Shea 1988). Furina tristis is a secretive, nocturnal snake that reaches about a meter in length and feeds mainly on lizards, particularly Sphenomorphus. It is oviparous, with a clutch size of around six eggs. The Taipan, Oxyuranus scutellatus, is a large snake that may exceed 2.5 meters in length. It is diurnal and crepuscular and feeds largely on rodents. The Taipan can become very aggressive if aroused. Its venom is extremely toxic and the snake often inflicts multiple bites. For these reasons the taipan is one of the most dangerous venomous snakes in the world (Masci and Kendall 1995). Snakes of the genus Pseudechis are also aggressive if aroused and are considered extremely dangerous (Kuch et al. 2005). The two species recorded from Papua can exceed 2 m in length. They feed mainly on frogs, lizards, and mammals. Pseudechis rossignolii is probably similar to the Australian P. australis and is likely a savanna and woodland species; P. papuanus tends to prefer swamps and wetlands. The death adders (genus Acanthophis) are generally short (one meter or less in total length), heavy-bodied snakes that are mostly nocturnal. They tend to be rather sluggish, ‘‘sit and wait’’ predators that feed mainly on small vertebrates, particularly lizards, and small mammals (Shine 1980).

................. 16157$

CH39

02-08-07 10:47:24

PS

PAGE 598

Herpetofauna of Papua / 599

Death adders produce living young, with up to eight young reported for New Guinea species (O’Shea 1988). The remaining elapine genera are all endemic to New Guinea. Micropechis ikaheka can reach lengths of 2 m or more. It occurs in a variety of forest habitats and can also be found in disturbed areas such as coconut plantations. It is mostly nocturnal and semi-fossorial but is occasionally seen in the open during the day. Aspidomorphus and Toxicocalamus have undergone modest species radiations in New Guinea and are found from lowlands to about 1,500 m elevation. The two species of Aspidomorphus found in Papua are small terrestrial snakes, inhabiting forest and woodland habitat. Their biology is poorly known but they likely prey on lizards and perhaps invertebrates. They are probably oviparous. The four species of Toxicocalamus are mostly small, leaf litter-inhabiting snakes that prey mainly on soft-bodied invertebrates (O’Shea 1988). They are all apparently oviparous.

Endemism and Hotspots The major centers of faunal endemism in Papua involve a range of New Guinea geological terranes (e.g., Cyclops Mts), isolated mountain ranges (e.g., Fakfak Mts; Weyland Mts), satellite islands (e.g., Raja Ampat group and Schouten Islands), and lowland basins (e.g., southern savannas). These are generally also areas of high endemism and species richness for amphibians and reptiles (Allison 1998). As such, they were highlighted as important areas of endemism in the 1997 Irian Jaya Biodiversity Conservation Priority-setting Workshop (CPSW) (Allison 1998; Conservation International 1999). That workshop identified 19 areas as being important areas of endemism (Figure 4.6.14): southern savannas, Digul River drainage, Lorentz National Park, Charles Louis Mountains, Weyland Mountains, Kumawa Mountains, Fakfak Mountains, Batanta Island, Waigeo Island, Tamrau and Arfak Mountains (Vogelkop), Wandammen Peninsula, Numfoor Island, BiakSupiori Islands, Yapen Island, Van Rees Mountains, Mamberamo drainage, Foja Mountains, Cyclops Mountains, and Jayawijaya (Star) Mountains. In addition to these areas, it is convenient to add the Aru Islands because they are biogeographically part of the southern lowlands of southwest New Guinea, although they are politically part of Maluku Province. If we focus on the amphibian and reptile species endemic to Papua (109 species; Appendix 8.4) and exclude two species for which the range, and even the type locality, is unknown (Cyrtodactylus irianjayensis and Hypsilurus nigrigularis), we are left with 107 endemic species. Ninety-one of those (85%) occur in at least one CPSW endemic area. Fifty-four (50%) are endemic (or nearly endemic) to a single area and seven are endemic or nearly endemic to two or more areas. Thirty additional species occur in at least one or more CPSW endemic areas but are also found well outside those areas. Only 16 (15%) of the amphibians and reptiles endemic to Papua do not occur in any of the aforementioned areas of endemism (Table 4.6.3). Half of these were

................. 16157$

CH39

02-08-07 10:47:24

PS

PAGE 599

600 /

Figure 4.6.14. Major areas of endemism of the amphibians and reptiles of Papua, based on the 1997 Conservation Priority-setting Workshop (see Allison 1998), with the addition of the Aru Islands. described prior to 1954 and the rest were described in 1988 or later. They are generally limited to single small areas scattered across Papua, and except in one possible instance do not appear to indicate additional areas of endemism. The only gecko in the group, Gehyra leopoldi, and one of the skinks, Lipinia venemai, occur in lowlands in the western part of the Vogelkop. The only snake, Heurnia ventromaculata, occurs in the lowlands of the Mamberamo Basin well outside the area of endemism identified by the CPSW. It is known only from a single specimen. The recently described monitor lizard, Varanus reisingeri, is endemic to Misool (Eidenmu¨ller and Wicker 2005) and is currently the only terrestrial vertebrate endemic to that island. At least six of the 107 endemic species occur just outside of at least one area of endemism and may very well occur within those areas. These include four frogs: Litoria mystax, which is found in coastal areas to the west of the Cyclops Mts; Litoria sanguinolenta, which is currently known only from just outside Lorentz National Park; Xenobatrachus scheepstrai, known from near the Jayawijaya Mts area of endemism, and Xenorhina adisca, which is known only from the type locality in mountains just outside Lorentz National Park; and two skinks: Sphenomorphus mimikanus and S. wollastoni, both known from areas just east of Lorentz

................. 16157$

CH39

02-08-07 10:47:51

PS

PAGE 600

Herpetofauna of Papua / 601

Table 4.6.3. Distribution of amphibian and reptile species endemic to Papua Number of species represented

Region Southern Lowlands Digul Drainage

Number of endemic species

2

2

5

1

19

7

9

0

Weyland Mountains

13

1

Kumawa Mountains

0

0

Fakfak Mountains

5

1

Batanta Islands

5

1

Waigeo Island

4

1

Lorentz National Park Charles Louis Mountains

Tamrau and Arfak Mountains (Vogelkop)

16

7

Wandammen Peninsula

21

13

Numfoor Island

2

0

Biak/Supiori Islands

4

2

Yapen Island

10

8

Van Rees Mountains

1

0

Mamberambo Drainage

9

1

Foja Mountains

0

0

Cyclops Mountains Jayawijaya Mountains Aru Islands

4

2

10

3

6

4

Note: 107 species were included in the analysis; the distribution of two species is unknown. The Aru Islands are biogeographically part of southern New Guinea and are included although they are politically part of Maluku Province.

National Park. It is very likely that all six of the aforementioned species occur within areas of endemism. If we include these species, the number of species found in CPSW areas of endemism increases to 97 (91%). Five recently described species of frogs from the Wapoga River drainage (Gu¨nther 2001; Gu¨nther, Richards, and Iskandar 2001; Richards 2001; Richards and Iskandar 2000) are not known from any of the areas of endemism. These include three microhylids: Hylophorbus sextus, Oreophryne minuta, and O. wapoga, and two hylids: Litoria macki and L. wapogaensis. An additional hylid frog, Litoria brongersmai, was described nearby from Doorman Top in 1946. In addition, the collections are thought to include 28 undescribed species. This suggests that the Wapoga Basin is rich in endemic species and perhaps should be recognized as an

................. 16157$

CH39

02-08-07 10:47:52

PS

PAGE 601

602 /

additional area of endemism. It is quite possible, however, that additional work will demonstrate that many of these species occur in the nearby Van Rees Mts endemic area and perhaps in other areas as well. Six areas of endemism have ten or more Papua endemic species, namely (in order of importance), Wandammen Peninsula, Lorentz National Park, Tamrau/ Arfak Mountains (Vogelkop), Weyland Mountains, Yapen Island and the Jayawijaya (Star) Mountains. These represent the richest areas of endemism in Papua (Table 4.6.4). The five most important areas for restricted range species—those restricted to a single CPSW endemic area—include, in order of importance, the Wandammen Peninsula, Yapen Island, Lorentz National Park, the Tamrau/Arfak Mountains (Vogelkop), and the Aru Islands. However, this analysis is somewhat misleading because some areas are virtually

Table 4.6.4. Species occurring outside the areas of endemism Taxon Frogs Family Hylidae Litoria brongersmai (Loveridge 1945) Litoria macki Richards 2001 Litoria mystax (van Kampen 1906) Litoria sanguinolenta (van Kampen 1909) Litoria wapogaensis Richards and Iskandar 2001 Family Microhylidae Hylophorbus sextus Gu¨nther 2001 Oreophryne minuta Richards and Iskandar 2000 Oreophryne wapoga Gu¨nther, Richards and Iskandar 2001 Xenobatrachus scheepstrai Blum and Menzies 1988 Lizards Family Gekkonidae Gehyra leopoldi Brongersma, 1930 Family Scincidae Lipinia venemai Brongersma, 1953 Sphenomorphus mimikanus (Boulenger 1914) Sphenomorphus wollastoni (Boulenger 1914) Family Varanidae Varanus reisingeri Eidenmu¨ller and Wicker 2005 Snakes Family Colubridae (Homolopsinae) Heurnia ventromaculata Jong 1926 Note: Areas of endemism identified at the 1997 Conservation Priority Setting Workshop (Conservation International 1999).

................. 16157$

CH39

02-08-07 10:47:52

PS

PAGE 602

Herpetofauna of Papua / 603

unsurveyed for amphibians and reptiles but, on the basis of better known groups such as birds, are judged to be important areas of endemism. Examples would include the Foja Mts on the north coast and the Kumawa Mts on the southwest corner of the Bomberai Peninsula. There are no endemic amphibians or reptiles recorded from either of these areas, primarily because they remain unsurveyed for these taxa. It is very likely, however, that further work will demonstrate that these areas are rich in endemic herp species. Moreover, at least two of the areas, the Wandammen Peninsula (Wondiwoi Mts) and Yapen Island, rank high in endemic species, largely because most of the frogs described from these areas are currently known only from their type localities and undoubtedly have wider distributions. This high ranking in endemics may therefore reflect an artifact that can be resolved only through additional fieldwork. Yapen, for example, was once connected to the New Guinea mainland (Chapter 2.1) and has no endemic birds or mammals but does have three species of endemic fish (Chapter 4.8). Nevertheless, the CPSW endemic areas are occupied, at a minimum, by 85% of the herp species endemic to Papua. These are also areas of high species richness (Allison, in prep.). At least nine of the areas include isolated mountain ranges, including north coast ranges such as the Cyclops Mts and the Wandammen Peninsula, that represent geological terranes. Six of the areas include islands (including the Aru Islands) which, because of their geographic isolation, tend to be centers of endemism. One of the areas, the Jayawijaya Mts, represents a major mountain range that, at a broad scale, is geographically isolated from other mountains. Three of the areas, Lorentz National Park, the Mamberamo drainage, and the Digul River drainage are areas that include a mixture of lowlands and highlands. Their relatively high endemism probably reflects both habitat diversity and geographic isolation. The remaining area, the southern lowlands, represents a climatic zone of highly seasonal rainfall that is dominated by open woodland-savanna vegetation. Only two species endemic to Papua are endemic to the southern lowlands. One is a turtle, Chelodina reimanni and the other is a hylid frog, Litoria quadrilineata. However, this low incidence of endemism is probably an artifact because the woodland-savanna vegetation occurs on either side of the border between Indonesia and Papua New Guinea and many of the species restricted to this vegetation are found in both countries, and, in some cases, northern Australia as well. Other analysis has shown that the woodland-savanna regions of New Guinea have very high herpetofaunal species richness (Allison, Kraus, and McShane 2004).

Conservation The most serious conservation challenges to the New Guinea herpetofauna involve land clearance, with resulting loss of habitat, and illegal collecting for the pet trade. In addition, for turtles, the harvesting of eggs and nesting females and, for crocodiles, the killing of adults for their skins, are significant sources of mortality.

................. 16157$

CH39

02-08-07 10:47:52

PS

PAGE 603

604 /

The impact of these factors on the herpetofauna is difficult to assess in the absence of definitive field survey data. However, it is clear that large tracts of forest in Papua are being logged (see Chapter 7.1) and that there is accelerating land clearance for agriculture, such as oil palm plantations, and for subsistence farming. These impacts are mostly in lowland areas, but as the demand for arable land increases, subsistence gardeners are moving upslope and are beginning to threaten important upland areas of endemism close to population centers, such as the Cyclops Mountains. Yuwono (1998) provides important information on the reptile trade in Indonesia. He divides the market into three categories: the zoo market and the ‘‘collector and breeder’’ market, both of which are thought to be relatively small, and the pet trade, which mostly involves common species and is responsible for most of the volume in the Indonesian reptile trade. According to Yuwono, Papua is the largest supplier of live reptiles and amphibians in Indonesia. In part this is because of high demand for taxa that are banned for export from Australia but are available from Papua. The major collecting areas are thought to be Sorong, Jayapura, Wamena, and Merauke. The live pet trade in Papua, according to Yuwono (1998) focuses mainly on 38 taxa: five frogs, seven turtles, 12 lizards, and 14 snakes. However, his list does not include a highly prized turtle, Carettochelys insculpta, and some recently described taxa. All the species treated by Yuwono (1998) are listed as abundant or common except a turtle, Chelodina parkeri; a skink, Tribolonotus gracilis; a monitor lizard, Varanus salvadorii; and two pythons, Apodora papuana and Morelia boeleni. These are all listed as rare. The legal trade in live reptiles and amphibians in Indonesia is strictly regulated, but there is thought to be a large and growing illegal trade, particularly involving rare species. Some rare species, such as Boelen’s Python (Morelia boeleni) or Salvadori’s Monitor (Varanus salvadorii) command a high price on the world market. Unscrupulous collectors working with local people in remote areas can readily extirpate such species from a large area. Harvesting of turtle eggs and nesting females is thought to be responsible for significant declines in the populations of target species, particularly sea turtles (Putrawidjaja 2000) and the Pig-nosed Turtle, Carettochelys insculpta (Samedi and Iskandar 2000). Although the harvesting of crocodiles is, in principle, strictly regulated, there is thought to be a flourishing illegal trade that has greatly reduced and perhaps even eliminated some populations. It is important to closely monitor and regulate the live pet trade to ensure that rare species are not collected to extinction. It is equally important to closely monitor and manage the harvest of marine and large species of freshwater turtles and their eggs. It is also important to monitor and manage the harvest of crocodiles. Finally, it is crucial to designate a network of protected areas. To support this it is essential to conduct faunal surveys throughout the province, particularly in the CPSW endemic areas, to discover and describe new species and better document the ranges of known species. This information is necessary for guiding conserva-

................. 16157$

CH39

02-08-07 10:47:53

PS

PAGE 604

Herpetofauna of Papua / 605

tion priority setting and the identification of species and areas that need formal protection. The recently completed Global Assessment of Amphibians (Stuart et al. 2004) demonstrated that about 150 frogs from the New Guinea and the Solomon Islands region were so poorly known that their conservation status could not be assessed. This is about half the frog species known from the region, making it the most poorly known area in the world. The situation for reptiles is somewhat better, but many species remain equally poorly known. Lack of knowledge is a particularly acute problem in Papua. Based on experience from Papua New Guinea, it is likely that there are hundreds of undescribed frog and reptile species in Papua. These species will be overlooked by conservation planners until they are discovered and scientifically named. The scientific community, conservation organizations, and the Indonesian government need to work in partnership to conduct field surveys and incorporate the findings into conservation planning and policy setting to ensure that the extraordinary herpetofauna of Papua is protected and preserved.

Acknowledgments I would like to thank Fred Kraus and Dan Polhemus for critically reviewing the penultimate draft; Carla Kishinami for critically reviewing all the drafts; and Brad Evans for help in preparing the figures. This material is based upon work supported by the U.S. National Science Foundation.

Literature Cited Allison, A. 1996. Zoogeography of amphibians and reptiles of New Guinea and the Pacific region. Pp. 407–436 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia. SPB Academic Publishing, Amsterdam. Allison, A. 1998. Taxa group summary: reptiles and amphibians. Pp. 47–48 in Burnett, J.B., Y. de Fretes, and P. Kramadibrata (eds.) The Irian Jaya Biodiversity Conservation Priority-Setting Workshop: Final Report. Conservation International, Washington, D.C. Allison, A., and A.A. Dwiyahreni. 1998. The amphibians and reptiles of the PT Freeport Indonesia Contract of Work Mining Project Area, Irian Jaya, Indonesia. Biodiversity Study Series, vol. 8. P.T. Hatfindo, Bogor, Indonesia. Allison, A., and F. Kraus. 2000. A new species of frog of the genus Xenorhina (Anura: Microhylidae) from the north coast ranges of Papua New Guinea. Herpetologica 56 (3): 285–294. Allison, A., F. Kraus, and M. McShane. 2004. Patterns of species richness in the Papuan region: a preliminary assessment using amphibians and reptiles. Report prepared for The Nature Conservancy. Bishop Museum, Honolulu. Aplin, K.P., P.R. Baverstock, and S.C. Donnellan. 1993. Albumin immunological evidence for the time and mode or origin of the New Guinea terrestrial mammal fauna. Science in New Guinea 19: 131–145. Archer, M., S. Hand, and H. Godthelp. 1991. Riversleigh. Reed Books, Balgowlah, New South Wales.

................. 16157$

CH39

02-08-07 10:47:53

PS

PAGE 605

606 / Austin, C.C. 2000. Molecular phylogeny, population structure, and historical biogeography of Pacific Island boas (Candoia). Copeia (2): 341–352. Bauer, A.M., and K. Henle. 1994. Familia Gekkonidae (Reptilia, Sauria). Part 1 Australia and Oceania. Das Tierreich. Bedford, G.F., K.A. Christian, and A.D. Griffiths. 1993. Preliminary investigations on the reproduction of the frillneck lizard Chlamydosaurus kingii in the Northern Territory. Pp. 127–131 in Lunney, D., and D. Ayers (eds.) Herpetology in Australia: A Diverse Discipline. Royal Zoological Society of New South Wales; distributed and sold by Surrey Beatty & Sons, Mosman, New South Wales, Australia; Chipping Norton, New South Wales, Australia. Bickford, D.-P. 2004. Differential parental care behaviors of arboreal and terrestrial microhylid frogs from Papua New Guinea. Behavioral Ecology and Sociobiology 55 (4): 402–409. Bo¨hme, W. 1988. Der Arguswaran (Varanus panoptes Storr, 1980) auf Neuguinea: V. panoptes horni ssp. n. Salamandra 24 (2/3): 87–101. Bo¨hme, W. 1991. The identity of Varanus gouldii (Gray, 1838) and the nomenclature of the V. gouldii-species complex. Mertensiella 2: 38–41. Bo¨hme, W. 2003. Checklist of the living monitor lizards of the world (family Varanidae). Zoologische Verhandelingen 341: 1–43. Bo¨hme, W., H.-G. Horn, and T. Ziegler. 1994. Zur Taxonomie der Pazifikwarane (Varanus-indicus Komplex): Revalidierung von Varanus doreanus (A.B. Meyer, 1874) mit Beschreibung einer neuer Unterart. Salamandra 30 (2): 119–142. Bo¨hme, W., and H.J. Jacobs. 2001. Varanus macraei sp. n., eine neue Waranart der V. prasinus-Gruppe aus West Irian, Indonesien. Herpetofauna 23 (133): 5–10. Boulenger, G.A. 1914. An annotated list of the batrachians and reptiles collected by the British Ornithologists’ Union Expedition and the Wollaston Expedition in Dutch New Guinea. Transactions of the Zoological Society of London 20: 247–266. Boulenger, G.A. 1921. Liste des publications ichthyologiques et herpe´tologiques (1877–1920). Annales Socie´te´ Royale et Malacologique de Belgique 52: 11–88. Brongersma, L.D., and G.F. Venema. 1962. To the Mountains of the Stars. [English translation.] Hodder and Stoughton, London. Brown, G.P., and R. Shine. 2002. Reproductive ecology of a tropical natricine snake Tropidonophis mairii (Colubridae). Journal of Zoology (London) 258: 63–72. Brown, W.C. 1991. Lizards of the genus Emoia (Scincidae) with observations on their ecology and biogeography. Memoirs of the California Academy of Sciences 15: 1–94. Burbrink, F.T. 2005. Inferring the phylogenetic position of Boa constrictor among the Boinae. Molecular Phylogenetics and Evolution 34: 167–180. Cann, J. 1998. Australian Freshwater Turtles. Privately published, Singapore. Cogger, H.G. 1964. A reptile collecting expedition to New Guinea. Australian Natural History 14 (11): 363–368. Cogger, H.G. 2000. Reptiles & Amphibians of Australia. 6th ed. Reed New Holland, Sydney. Conservation International. 1999. Peta Lokakarya Prioritas Konservasi di Irian Jaya. Okakarya Penentuan Daerah Prioritas Konservasi Keanekaragaman Hayati Irian Jaya. The Irian Jaya Biodiversity Conservation Priority-Setting Workshop, 7–12 January, Biak, Irian Jaya [Map]. Conservation International, Washington, D.C. 1:1,575,000 Das, D.I., and G. Ismail. 2001. Lizards of Borneo. Available at http://www.arbec.com.my/ lizards/. Das, I., and V. Wallach. 1998. Scolecophidian arboreality revisited. Herpetological Review 29 (1): 15–16.

................. 16157$

CH39

02-08-07 10:47:53

PS

PAGE 606

Herpetofauna of Papua / 607 David, P., and G. Vogel. 1996. A synthetic list of the snakes of Sumatra. Pp. 24–28 in David, P., and G. Vogel (eds.) The Snakes of Sumatra. Edition Chimaira, Frankfurt am Main. de Rooij, N. 1915. The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E.J. Brill, Leiden. de Rooij, N. 1917. The Reptiles of the Indo-Australian Archipelago. II. Ophidia. E.J. Brill, Leiden. Doody, J.S., A. Georges, and J.E. Young. 2003. Twice every second year: reproduction in the pig-nosed turtle, Carettochelys insculpta, in the wet-dry tropics of Australia. Journal of Zoology (London) 259: 179–188. Dow, D.B. 1977. A geological synthesis of Papua New Guinea. Bureau of Mineral Resources, Geology and Geophysics Bulletin, Australian Government Publishing Service, Canberra. Duellman, W.E. 1978. The biology of an equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publications of the Museum of Natural History, University of Kansas 65: 1–352. Duellman, W.E., and L. Trueb. 1986. Biology of Amphibians. McGraw-Hill, New York. Dume´ril, A.M.C., and G. Bibron. 1836. Erpe´tologie Ge´ne´rale ou Histoire Naturelle Comple`te des Reptiles. Vol. 3. Roret, Paris. Dume´ril, A.M.C., and G. Bibron. 1837. Erpe´tologie Ge´ne´rale ou Histoire Naturelle Comple`te des Reptiles. Vol. 4. Roret, Paris. Dume´ril, A.M.C., and G. Bibron. 1839. Erpe´tologie Ge´ne´rale ou Histoire Naturelle Comple`te des Reptiles. Vol. 5. Roret, Paris. Eidenmu¨ller, B., and R. Wicker. 2005. Eine weitere neue Waranart aus dem Varanaus prasinus-Komplex von der Insel Misol, Indonesien. Sauria 27 (1): 3–8. Ernst, C.H., and R.W. Barbour. 1989. Turtles of the World. Smithsonian Institution Press, Washington, D.C. Fritts, T.H., M.J. McCoid, and D.M. Gomez. 1999. Dispersal of snakes to extralimital islands: incidents of the brown treesnake (Boiga irregularis) dispersing to islands in ships and aircraft. Pp. 209–223 in Rodda, G.H., Y. Sawai, D. Chiszar, and H. Tanaka (eds.) Problem Snake Management: Habu and Brown Tree Snake Examples. Cornell University Press, Ithaca, New York. Frost, D.R. 2005. Amphibian Species of the World: An Online Reference. Version 3.0 (22 August 2004). Electronic Database accessible at http://research.amnh.org/herpetology/ amphibia/index.html. American Museum of Natural History, New York. Gaulke, M. 1975. Observations on arboreality in a Philippine blind snake. Asiatic Herpetological Research 6: 45–48. Georges, A., M. Adams, and W. McCord. 2002. Electrophoretic delineation of species boundaries within the genus Chelodina (Testudines: Chelidae) of Australia, New Guinea and Indonesia. Zoological Journal of the Linnean Society 134: 401–421. Georges, A., S. Doody, J. Young, and J. Cann. 2000. The Australian Pig-Nosed Turtle. Robey, Canberra. Glaessner, M.F. 1942. The occurrence of the New Guinea turtle (Carettochelys) in the Miocene of Papua. Records of the Australian Museum 21: 106–109. Greene, H.W. 1986. Diet and arboreality in the emerald monitor, Varanus prasinus, with comments on the study of adaptations. Fieldiana, Zoology 31: 1–12. Greer, A.E. 1970a. The relationships of the skinks referred to the genus Dasia. Breviora 348: 1–30. Greer, A.E. 1970b. A subfamilial classification of scincid lizards. Bulletin of the Museum of Comparative Zoology 139: 151–183.

................. 16157$

CH39

02-08-07 10:47:54

PS

PAGE 607

608 / Greer, A.E. 1985. The relationships of the lizard genera Anelytropsis and Dibamus. Journal of Herpetology 19 (1): 116–156. Greer, A.E. 1989. The Biology and Evolution of Australian Lizards. S. Beatty, Chipping Norton, Australia. Greer, A.E. 1990. The Glaphyromorphus isolepis species group (Lacertilia: Scincidae): diagnosis of the taxon and description of a new species from Timor. Journal of Herpetology 24 (4): 372–377. Greer, A.E. 1997. The Biology and Evolution of Australian Snakes. Surrey Beatty & Sons, Chipping Norton, New South Wales. Greer, A.E., A. Allison, and H.G. Cogger. 2005. Four new species of Lobulia (Lacertilia: Scincidae) from high altitude in New Guinea. Herpetological Monographs 19: 153–179. Griffiths, A.D., and K.A. Christian. 1996. Diet and habitat use of frill neck lizards in a seasonal tropical environment. Oecologia 106: 39–48. Groombridge, B. 1982. The IUCN Amphibia-Reptilia Red Data Book. Part 1. Testudines, Crocodylia, Rhynchocephalia. International Union for Conservation of Nature and Natural Resources, Gland, Switzerland. Gu¨nther, R. 1999. Morphological and bioacoustic characteristics of frogs of the genus Platymantis (Amphibia, Ranidae) in Irian Jaya, with descriptions of two new species. Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin. Zoologische Reihe 75 (2): 317–335. Gu¨nther, R. 2000a. Albericus laurini species nova, the first record of the genus Albericus (Anura, Microhylidae) from the west of New Guinea. Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin. Zoologische Reihe 76 (2): 167–174. Gu¨nther, R. 2000b. In alten Sammlungen aus Neuguinea entdeckt: zwei neue Arten der Gattung Lipinia (Squamata: Scincidae). Salamandra 36 (3): 157–174. Gu¨nther, R. 2001. The Papuan frog genus Hylophorbus (Anura: Microhylidae) is not monospecific: description of six new species. Russian Journal of Herpetology 8 (2): 81–104. Gu¨nther, R. 2002a. Beschreibung einer neuen Copiula-Art (Amphibia, Anura, Microhylidae) von der Insel Yapen im Nordwesten von Papua, Indonesien. Zoologische Abhandlungen 52: 77–86. Gu¨nther, R. 2002b. Westernmost records of the Papuan frog genus Copiula with descriptions of two new species (Amphibia: Anura: Microhylidae). Zoologische Abhandlungen Staatliches Museum fu¨r Tierkunde Dresden 23 (2): 35–58. Gu¨nther, R. 2003a. First record of the microhylid frog genus Cophixalus from western Papua, Indonesia, with descriptions of two new species. Herpetozoa 16 (1/2): 3–21. Gu¨nther, R. 2003b. Further new species of the genus Oreophryne (Amphibia, Anura, Microhylidae) from western New Guinea. Zoological Abhandlungen (Dresden) 53: 65–85. Gu¨nther, R. 2003c. Sexual colour dimorphism in ranid frogs from New Guinea: description of two new species (Amphibia, Anura, Ranidae). Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin. Zoologische Reihe 79 (2): 207–227. Gu¨nther, R. 2003d. Three new species of the genus Oreophryne from western Papua, Indonesia. Spixiana 26 (2): 175–191. Gu¨nther, R. 2004a. Description of a new treefrog species from western New Guinea showing extreme color polymorphism (Anura, Hylidae, Litoria). Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin. Zoologische Reihe 80 (2): 251–260. Gu¨nther, R. 2004b. Two new treefrog species of the genus Litoria (Anura: Hylidae) from the west of New Guinea. Zoologische Abhandlungen (Dresden) 54: 163–175.

................. 16157$

CH39

02-08-07 10:47:54

PS

PAGE 608

Herpetofauna of Papua / 609 Gu¨nther, R., M. Kapisa, and I. Tetzlaff. 2001. Ein seltense Brutpflegeverhalten bei Froschlurchen: Ma¨nnchen von Sphenophryne cornuta transportiert Jungtiere auf seinem Rucken (Anura, Microhylidae). Herpetofauna 23 (135): 14–24. Gu¨nther, R., and S.J. Richards. 2000. A new species of the Litoria gracilenta group from Irian Jaya. Herpetozoa 13 (1/2): 27–43. Gu¨nther, R., S.J. Richards, and D. Iskandar. 2001. Two new species of the genus Oreophryne from Irian Jaya, Indonesia. Spixiana 24 (3): 257–274. Gu¨nther, R., and H. Ro¨sler. 2002. Eine neue Art der Gattung Cyrtodactylus Gray, 1827 aus dem Westen von Neuguinea (Reptilia: Sauria: Gekkonidae). Salamandra 38 (4): 195–212. Gyi, K.K. 1970. A revision of colubrid snakes of the subfamily Homalopsinae. Miscellaneous Publications of the Museum of Natural History, University of Kansas 20 (2): 47–223. Hall, P.M. 1989. Variation in geographic isolates of the New Guinea Crocodile (Crocodylus novaeguineae Schmidt) compared with the similar, allopatric, Philippine Crocodile (C. mindorensis Schmidt). Copeia 1989 (1): 71–80. Han, D., K. Zhou, and A.M. Bauer. 2004. Phylogenetic relationships among gekkotan lizards inferred from C-mos nuclear DNA sequences and a new classification of the Gekkota. Biological Journal of the Linnean Society 83 (3): 353–368. Harlow, P., and R. Shine. 1992. Food habits and reproductive biology of the Pacific Island boas (Candoia). Journal of Herpetology 26 (1): 60–66. Harvey, M.B., D.G. Barker, L.K. Ammerman, and P.T. Chippindale. 2000. Systematics of pythons of the Morelia amethistina complex (Serpentes: Boidae) with the description of three new species. Herpetological Monographs 14: 139–185. Heatwole, H. 1999. Sea Snakes. 2nd ed. University of New South Wales Press, Sydney, Australia. Horn, H.-G. 2004. Varanus salvadorii. Pp. 234–243 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Hoser, R. 2000. A new species of snake from Irian Jaya (Serpentes: Elapidae). Litteratura Serpentium 20 (6): 178–186. Houston, D., and R. Shine. 1994. Population demography of Arafura filesnakes (Serpentes: Acrochordidae) in tropical Australia. Journal of Herpetology 28 (3): 273–280. How, R.A., B. Durrant, L.A. Smith, and N. Saleh. 1998. Emoia (Reptilia: Scincidae) from the Banda Arc islands of eastern Indonesia: variation in morphology and description of a new species. Records of the Western Australian Museum 19 (2): 131–139. in den Bosch, H.A.J. 1985. Snakes of Sulawesi: checklist, key and additional biogeographical remarks. Zoologische Verhandelingen (Leiden) 217: 3–50. Ineich, I., and G.R. Zug. 1991. Nomenclatural status of Emoia cyanura (Lacertilia, Scincidae) populations in the central Pacific. Copeia 4: 1132–1136. Inger, R.F., and F.L. Tan. 1996. Checklist of the frogs of Borneo. Raffles Bulletin of Zoology 44 (2): 551–574. Ingram, G., and J.A. Covacevich. 1989. Revision of the genus Carlia (Reptilia, Scincidae) in Australia with comments on Carlia bicarinata of New Guinea. Memoirs of the Queensland Museum 27 (2): 443–490. Iskandar, D.T. 2000. Turtles and Crocodiles of Insular Southeast Asia and New Guinea. Department of Biology, Institute of Technology, Bandung, Indonesia. Iverson, J. 1992. Species richness maps of the freshwater and terrestrial turtles of the world. Smithsonian Herpetological Information Service 88: 1–18.

................. 16157$

CH39

02-08-07 10:47:54

PS

PAGE 609

610 / Iverson, J.B. 1986. A Checklist with Distribution Maps of the Turtles of the World. Privately published, Richmond, Indiana. Jacobs, H.J. 2003. A further new emerald tree monitor lizard of the Varanus prasinus species group from Waigeo, West Irian (Squamata: Sauria: Varanidae). Salamandra 39 (2): 39–64. Jelden, D.C. 1981. Preliminary studies on the breeding biology of Crocodylus porosus and Crocodylus n. novaeguineae on the middle Sepik, Papua New Guinea. Amphibia-Reptilia 1 (3/4): 353–358. Jennings, W.B., and E.R. Pianka. 2004. Tempo and timing of the Australian Varanus radiation. Pp. 77–87 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Jones, F.W. 1909. The fauna of the Cocos-Keeling Atoll, collected by F. Wood Jones. Proceedings of the Zoological Society of London 1909 (3): 132–160. Kamosawa, M., and H. Ota. 1996. Reproductive biology of the brahminy blind snake (Ramphotyphlops braminus) from the Ryukyu Archipelago, Japan. Journal of Herpetology 30: 9–14. King, F.W., and R.L. Burke. 1989. Crocodilian, Tuatara, and Turtle Species of the World: A Taxonomic and Geographic Reference. Association of Systematics Collections, Washington, D.C. Kluge, A.G. 1974. A taxonomic revision of the lizard family Pygopodidae. Miscellaneous Publications Museum of Zoology, University of Michigan 147: 1–221. Kluge, A.G. 1993. Aspidites and the phylogeny of pythonine snakes. Records of the Australian Museum Supplement 19: 1–77. Kraus, F., and A. Allison. 2004. A new species of Tropidonophis (Serpentes: Colubridae: Natricinae) from Fergusson Island, Papua New Guinea. Proceedings of the Biological Society of Washington 117: 303–310. Kuch, U., J.S. Keogh, J. Weigel, L.A. Smith, and D. Mebs. 2005. Phylogeography of Australia’s king brown snake (Pseudechis australis) reveals Pliocene divergence and Pleistocene dispersal of a top predator. Naturwissenschaften 92 (3): 121–127. Kuramoto, M., and A. Allison. 1989. Karyotypes of microhylid frogs of Papua New Guinea and their systematic implications. Herpetologica 45 (2): 250–259. Lesson, R.P. 1830. Zoologie. Pp. 1–65 in Duperrey, L.I. (ed.) Atlas de zoologie, Voyage autour du monde, exe´cute´ par ordre du roi, sur la corvette de sa Majeste´, La Coquille, pendant les anne´es 1822, 1823, 1824 et 1825. Vol. 2 (1), Arthus Bertrand, Paris. Lillywhite, H.B. 1996. Husbandry of the little file snake, Acrochordus granulatus. Zoo Biology 15: 315–327. Lorentz, H.A. 1913. Zwarte Menschen, Witte Berge: Verhaal van den Tocht naar het Sneeuwgebergte van Nieuw-Guinea. Brill, Leiden. Loveridge, A. 1948. New Guinean reptiles and amphibians in the Museum of Comparative Zoology and the United States National Museum. Bulletin of the Museum of Comparative Zoology 101 (2): 305–430. Madsen, T., and R. Shine. 1996a. Determinants of reproductive output in female water pythons (Liasis fuscus: Pythonidae). Herpetologica 52: 146–159. Madsen, T., and R. Shine. 1996b. Seasonal migration of predators and prey: a study of pythons and rats in tropical Australia. Ecology 77: 149–156. Madsen, T., and R. Shine. 2001. Conflicting conclusions from long-term versus shortterm studies on growth and reproduction of a tropical snake. Herpetologica 57: 147–156. Malnate, E.V., and G. Underwood. 1988. Australasian natricine snakes of the genus

................. 16157$

CH39

02-08-07 10:47:55

PS

PAGE 610

Herpetofauna of Papua / 611 Tropidonophis. Proceedings of the Academy of Natural Sciences, Philadelphia 140 (1): 59–201. Masci, P., and P. Kendall. 1995. The Taipan: The World’s Most Dangerous Snake. Kangaroo Press, Kenthurst, New South Wales, Australia. McCord, W.P., and S.A. Thomson. 2002. A new species of Chelodina (Testudines: Pleurodira: Chelidae) from northern Australia. Journal of Herpetology 36: 255–267. McCoy, M. 1980. Reptiles of the Solomon Islands. Wau Ecology Institute, Wau, Papua New Guinea. McDiarmid, R.W., J.A. Campbell, and T.S.A. Touraˆe. 1999. Snake Species of the World: A Taxonomic and Geographic Reference. Herpetologists’ League, Washington, D.C. McDowell, S.B. 1967. Aspidomorphus, a genus of New Guinea snake of the family Elapidae, with notes on related genera. Journal of Zoology, London 151: 497–543. McDowell, S.B. 1969. Toxicocalamus, a New Guinea genus of snakes of the family Elapidae. Journal of Zoology, London 159: 443–511. McDowell, S.B. 1972. The species of Stegonotus (Serpentes, Colubridae) in Papua New Guinea. Zoologische Mededelingen 47: 6–26. McDowell, S.B. 1974. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum. Part I. Scolecophidia. Journal of Herpetology 8 (1): 1–57. McDowell, S.B. 1975. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum. Part II. Aniliodea and Pythoninae. Journal of Herpetology 9 (1): 1–79. McDowell, S.B. 1979. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum. Part III. Boinae and Acrochordoidea. Journal of Herpetology 13 (1): 1–92. McDowell, S.B. 1984. Results of the Archbold Expeditions. No. 112. The snakes of the Huon Peninsula, Papua New Guinea. American Museum Novitates 2775: 1–28. McDowell, S.B. 1987. Systematics. Pp. 3–50 in Seigel, R.A., J.T. Collins, and S.S. Novak (eds.) Snakes: Ecology and Evolutionary Biology. Macmillian, New York. Mendes, L.F., M. Pinheiro, and C. Ruas. 1994. Resultados das misso˜es de estudo realizadas em Macau pelo Centro de Zoologia do IICT. IV.–Nota sobre os conteu´dos ga´stricos de anuros. 2-Microhylidae, Ranidae, Rhacophoridae. Concluso˜es. Garcia de Orta Se´rie de Zoologia 20 (1–2): 41–48. Menzies, J.I. 1976. Handbook of Common New Guinea Frogs. Wau Ecology Institute, Handbook 1, Wau, Papua New Guinea. Menzies, J.I. 1987. A taxonomic revision of the Papuan Rana (Amphibia: Ranidae). Australian Journal of Zoology 35: 373–418. Menzies, J.I. 1992. Ecological and taxonomic notes on ranid frogs (Amphibia: Ranidae) from far western New Guinea. Science in New Guinea 18 (3): 115–122. Menzies, J.I. 1996. Unnatural distribution of fauna in the East Malesian region. Pp. 31–38 in Kitchener, D.J., and D. Suyanto (eds.) Proceedings of the First International Conference on Eastern Indonesian-Australian Vertebrate Fauna. Published by the Western Australian Museum for Lembaga Ilmu Pentegahuan Indonesia (Indonesian Institute of Sciences), Perth. Menzies, J.I., and R.F. Tapilatu. 2000. The introduction of a second species of toad (Amphibia: Bufonidae) into New Guinea. Science in New Guinea 25 (1–3): 70–73. ¨ ber die von ihm auf Neu-Guinea und den Inseln Jobi, Mysore und Meyer, A.B. 1874. U Mafoor im Jahre 1873 gesammelten amphibien. Monatsbericht der Ko¨niglich Preussischen Akademie der Wissenschaften zu Berlin 1874: 128–140.

................. 16157$

CH39

02-08-07 10:47:55

PS

PAGE 611

612 / Mori, A. 1998. Prey-handling behavior of three species of homalopsine snakes: features associated with piscivory and Duvernoy’s glands. Journal of Herpetology 32: 40–50. Moss, S.J., and M.E.J. Wilson. 1998. Biogeographic implications of the Tertiary paleogeographic evolution of Sulawesi and Borneo. Pp. 133–163 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. Nasution, R.E., R. Jusuf, M. Siregar, M. Mansur, A. Sawito, and H. Kurniati. 1996. Eksplorasi flora dan fauna di Cagar Alam Pegunungan Cyclops, Irian Jaya. Pp. 466–497 in Arief, A.J., D.S. Said, Y. Jamal, S. Paryanti, H. Julistiono, Rochadi, and E. Harsono (eds.) Laporan Teknik Proyek Penelitian, Pengembangan dan Pendayagunaan Biota darat Tahun 1995/1996. Pusat Penelitian dan Pengembangan Biologi, Lembaga Ilmu Pengetahuan Indonesia, Bogor, Indonesia. Nussbaum, R.A. 1980. The brahminy blind snake, Ramphotyphlops braminus, in the Seychelles Archipelago, Indian Ocean: distribution, variation and further evidence for parthenogenesis. Herpetologica 36 (3): 215–221. O’Shea, M. 1996. A Guide to the Snakes of Papua New Guinea. Independent Publishing, Port Moresby, Papua New Guinea. O’Shea, M.T. 1988. Liasis papuanus (Papuan Olive Python) feeding. Herpetological Review 19 (2): 36. Ota, H., T. Hikida, M. Matsui, A. Mori, and A.H. Wynn. 1991. Morphological variation, karyotype and reproduction of the parthenogenetic blind snake, Ramphotyphlops braminus, from the insular region of East Asia and Saipan. Amphibia-Reptilia 12: 181–193. Parker, F. 1982. The snakes of Western Province. Wildlife in Papua New Guinea 82 (1): 1–78. Parker, H.W. 1934. A Monograph of the Frogs of the Family Microhylidae. British Museum (Natural History), London. Parmley, D., and J.A. Holman. 2003. Nebraskophis Holman from the Late Eocene of Georgia (USA), the oldest known North American colubrid snake. Acta Zoologica Cracoviensia 46: 1–8. Patchell, F.C., and R. Shine. 1986a. Feeding mechanisms in pygopodid lizards: how can Lialis swallow such large prey? Journal of Herpetology 20 (1): 59–64. Patchell, F.C., and R. Shine. 1986b. Food habits and reproductive biology of the Australian legless lizards (Pygopodidae). Copeia 1986 (1): 30–39. Peters, W., and G. Doria. 1878. Catalago dei Retilli e dei Batraci racolti da O. Beccari, L.M. D’Albertis e A.A. Bruijn nella Sotto-Regione Austro-Malese. Annali del Museo Civico di Storia Naturale di Genova 13: 323–450. Philipp, K.M. 1999. Niche partitioning of Varanus doreanus, V. indicus and V. jobiensis in Irian Jaya: preliminary results. Mertensiella 11: 307–316. Philipp, K.M., W. Bo¨hme, and T. Ziegler. 1999. The identity of Varanus indicus: redefinition and description of sibling species coexisting at the type locality. Spixiana 22 (3): 273–287. Phillips, B.L., G.P. Brown, and R. Shine. 2003. Assessing the potential impact of cane toads on Australian snakes. Conservation Biology 17: 1738–1747. Pianka, E.R., D. King, and R.A. King. 2004. Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. BMR Journal of Australian Geology 10 (3): 193–211. Povel, D., and J. van der Kooij. 1997. Scale sensillae of the file snake (Serpentes: Acro-

................. 16157$

CH39

02-08-07 10:47:55

PS

PAGE 612

Herpetofauna of Papua / 613 chordidae) and some other aquatic and borrowing snakes. Netherlands Journal of Zoology 47: 443–456. Putrawidjaja, M. 2000. Marine turtles in Irian Jaya, Indonesia. Marine Turtle Newsletter 90: 8–10. Rage, J.-C. 1987. Fossil history. Pp. 51–76 in Siegel, R.A., J.T. Collins, and S.S. Novak (eds.) Snakes: Ecology and Evolutionary Biology. Macmillan, New York. Rage, J.-C., E. Buffetaut, H. Buffetaut-Tong, Y. Chaimanee, S. Ducrocq, J.-J. Jaeger, and V. Suteethorn. 1992. A colubrid snake in the Late Eocene of Thailand: the oldest known Colubridae (Reptilia: Serpentes). Comptes Rendus de l’Academia des Sciences Serie´ II Me´canique Physique Chimie Sciences de l’Univers Sciences de la Terre 314: 1085–1089. Rawlings, L.H., D. Barker, and S.C. Donnellan. 2004. Phylogenetic relationships of the Austro-Papuan Liasis pythons (Reptilia: Macrostomata), based on mitochondrial DNA. Australian Journal of Zoology 52 (2): 215–227. Rhodin, A.G.J., R.A. Mittermeier, and P.M. Hall. 1993. Distribution, osteology, and natural history of the Asian giant softshell turtle, Pelochelys bibroni, in Papua New Guinea. Chelonian Conservation and Biology 1 (1): 19–30. Richards, S. 2001. A new species of torrent-dwelling frog (Anura: Hylidae: Litoria) from the mountains of Indonesian New Guinea (West Papua). Memoirs of the Queensland Museum 46: 733–739. Richards, S., and D. Iskandar. 2000. A new minute Oreophryne (Anura: Microhylidae) from the mountains of Irian Jaya, Indonesia. Raffles Bulletin of Zoology 48 (2): 257–262. Richards, S., D.T. Iskandar, and A. Allison. 2000. Amphibians and reptiles of the Wapoga River area, Irian Jaya, Indonesia. In Mack, A.L., and L.E. Alonso (eds.) A biological assessment of the Wapoga River area of northwestern Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14: 54–57, 113–120. Richards, S., D.T. Iskandar, and B. Tjaturadi. 2002. Amphibians and reptiles of the Dabra area, Mamberamo River Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25: 76–79, 164–167. Rodda, G.H., and T.H. Fritts. 1992. The impact of the introduction of the colubrid snake Boiga irregularis on Guam’s lizards. Journal of Herpetology 26 (2): 166–174. Roos, M.C., P.J.A. Kessler, S.R. Gradstein, and P. Baas. 2004. Species diversity and endemism of five major Malesian islands: diversity-area relationships. Journal of Biogeography 31 (12): 1893–1908. Rose, M., F. Parker, and A.G.J. Rhodin. 1982. New Guinea plateless turtle or pitted shell turtle (Fly River or pig-nosed turtle), Carettochelys insculpta Ramsay 1886. Pp. 243–246 in Groombridge, B. (ed.) The IUCN Amphibia-Reptilia Red Data Book. Part 1. Testudines, Crocodylia, Rhynchocephalia. International Union for the Conservation of Nature and Natural Resources, Gland, Switzerland. Ross, C.A. 1989. Crocodiles and Alligators. Golden Press Pty, Silverville, New SouthWales. Ross, C.A. 1990. Crocodylus raninius Mu¨ller and Schlegel, a valid species of crocodile (Reptilia: Crocodylidae) from Borneo. Proceedings of the Biological Society of Washington 103 (4): 955–961. Ross, R.A., and G. Marzec. 1990. The Reproductive Husbandry of Pythons and Boas. Institute for Herpetological Research, Stanford, California. Salm, R.V., R.G. Petocz, and T. Soehartono. 1982. Survey of coastal areas in Irian Jaya. UNDP/FAO and WWF/IUCN. FO/INS/78/061 Special Report. Bogor. Samedi and D.T. Iskandar. 2000. Freshwater turtle and tortoise conservation and utilization in Indonesia. Chelonian Research Monographs 2: 106–111. Shea, G.M. 1991. Nesting of the southern rainforest dragon Hypsilurus spinipes (Squamata: Agamidae). Herpetofauna 21 (2): 7–10.

................. 16157$

CH39

02-08-07 10:47:56

PS

PAGE 613

614 / ¨ kologie Shea, G.M. 2000. Die Neuguinea-Blauzunge, Tiliqua gigas (Schneider, 1801): O ¨ bersicht u¨ber die Unterarten nebst Beschreibung einer neuen Unterart, Tiliqua und U gigas evanescens subsp. nov. Pp. 177–189 in Hauschild, A., K. Henle, R. Hitz, G.M. Shea, and H. Werning (eds.) Blauzungenskinke. Beitra¨ge zu Tiliqua und Cyclomorphus. Natur und Tier-Verlag, Munich. Shine, R. 1980. Ecology of the Australian death adder Acanthophis antarcticus (Elapidae): evidence for convergence with the Viperidae. Herpetologica 36 (3): 281–289. Shine, R. 1991a. Australian Snakes: A Natural History. Cornell University Press, Ithaca, NewYork. Shine, R. 1991b. Strangers in a strange land: ecology of the Australian colubrid snakes. Copeia 1991 (1): 120–131. Shine, R., and M. Fitzgerald. 1996. Large snakes in a mosaic rural landscape: the ecology of carpet pythons, Morelia spilota (Serpentes: Pythonidae) in coastal eastern Australia. Biological Conservation 76: 113–122. Shine, R., and R. Lambeck. 1989. Ecology of frill neck lizards Chlamydosaurus kingii (Agamidae) in tropical Australia. Australian Wildlife Research 16: 491–500. Smith, H.M., D. Chiszar, K. Tepedelen, and F. van Breukelen. 2001. A revision of the bevelnosed boas (Candoia carinata complex) (Reptilia: Serpentes). Hamadryad 26 (2): 283–315. Smith, M., and M. Plane. 1985. Pythonine snakes (Boidae) from the Miocene of Australia. Journal of Australian Geology and Geophysics 9: 191–195. Souter, G. 1966. New Guinea: The Last Unknown. Taplinger, New York. Spotila, J.R. 2004. Sea Turtles: A Complete Guide to Their Biology, Behavior, and Conservation. Johns Hopkins University Press, Baltimore. Stuart, F.D.M., A.F. Hugall, and C. Moritz. 2002. A molecular phylogeny of rainbow skinks (Scincidae: Carlia): taxonomic and biogeographic implications. Australian Journal of Zoology 50 (1): 39–51. Stuart, S.N., J.S. Chanson, N.A. Cox, B.E. Young, A.S.L. Rodrigues, D.L. Fischman, and R.W. Waller. 2004. Status and trends of amphibian declines and extinctions worldwide. Science 306: 1783–1786. Stuebing, R.B. 1991. A checklist of the snakes of Borneo. Raffles Bulletin of Zoology 39 (2): 323–362. Stuebing, R.B. 1994. A checklist of the snakes of Borneo: addenda and corrigenda. Raffles Bulletin of Zoology 42 (4): 931–936. Tomascik, T., A.J. Mah, A. Nontji, and M.K. Moosa. 1997. The Ecology of the Indonesian Seas. Part 2. Periplus Editions, Hong Kong. Tweedie, M.W.F. 1983. The Snakes of Malaya. Singapore National Printer, Singapore. Tyler, M.J. 1962. Observations on the influence of frogs on the ecology of coffee plantations in the Western Highlands of New Guinea. Papua New Guinea Agricultural Journal 14 (4): 151–152. Tyler, M.J. 1976. Frogs. Australian Naturalist Library. Collins, Sydney. Tyler, M.J. 1999. Distribution patterns of amphibians and reptiles. Pp. 541–563 in Duellman, W.E. (ed.) Patterns of Distribution of Amphibians. Johns Hopkins University Press, Baltimore. Underwood, G. 1967. A Contribution to the Classification of Snakes. British Museum (Natural History), London. Underwood, G., and A.F. Stimson. 1990. A classification of pythons (Serpentes: Pythoninae). Journal of Zoology, London 221 (4): 565–604. Urban, H. 1999. Eine neue Agemenart der Gattung Gonocephalus aus Papua-New Guinea (Squamata: Sauria: Agamidae). Herpetozoa 11 (3–4): 185–188.

................. 16157$

CH39

02-08-07 10:47:56

PS

PAGE 614

Herpetofauna of Papua / 615 van Kampen, P.N. 1923. The Amphibia of the Indo-Australian Archipelago. E.J. Brill, Leiden. van Welzen, P.C. 1997. Increased speciation in New Guinea: tectonic causes. Pp. 363–387 in Dransfield, J., M.J.E. Coode, and D.A. Simpson (eds.) Plant Diversity in Malesia III: Proceedings of the Third International Flora Malesiana Symposium 1995. Royal Botanic Gardens, Kew, England. Voris, H.K., M.E. Alfaro, D.R. Karns, G.L. Starnes, E. Thompson, and J.C. Murphy. 2002. Phylogenetic relationships of the Oriental-Australian rear-fanged water snakes (Colubridae: Homalopsinae) based on mitrochondrial DNA sequences. Copeia 2002: 906–915. Voris, H.K., and B.C. Jayne. 1979. Growth, reproduction and population structure of a marine snake Enhydrina schistosa (Hydrophiidae). Copeia 1979: 307–318. Voris, H.K., and J.C. Murphy. 2002. The prey and predators of homalopsine snakes. Journal of Natural History 36: 1621–1632. Wallach, V. 1995. A new genus for the Ramphotyphlops subocularis species group (Serpentes: Typhlopidae), with description of a new species. Asiatic Herpetological Research 6: 132–150. Webb, J.K., and R. Shine. 1992. To find an ant: trail-following in Australian blind snakes (Typhlopidae). Animal Behaviour 43: 941–948. Webb, J.K., and R. Shine. 1993. Dietary habits of Australian blindsnakes (Typhlopidae). Copeia 1993 (3): 762–770. Webb, R.G. 1995. Redescription and neotype designation of Pelochelys bibroni from southern New Guinea (Testudines: Trionychidae). Chelonian Conservation and Biology 4 (1): 301–310. Webb, R.G. 1997. Geographic variation in the giant softshell turtle, Pelochelys bibroni. Linnaeus Fund research project. Chelonian Conservation and Biology 2 (3): 450. Webb, R.G. 2002. Observations on the giant softshell turtle, Pelochelys cantorii, with description of a new species. Hamadryad 27 (1): 99–107. Wermuth, H. 1967. Liste der rezenten Amphibien und Reptilien; Agamidae. Das Tierreich 86: 1–127. Wollaston, A.F.R. 1912. Pygmies & Papuans, the Stone Age To-day in Dutch New Guinea. Smith, Elder & Co., London. Wollaston, A.F.R. 1916. Introduction. Pp. 1–22 in Ogilvie-Grant, W.R. (ed.) Reports on the Collections Made by the British Ornithologists’ Union Expedition and the Wollaston Expedition in Dutch New Guinea, 1910–13. Francis Edwards, London. Wu¨ster, W., A.J. Dumbrell, C. Hay, C.E. Pook, D.J. Williams, and B.G. Fry. 2005. Snakes across the Strait: trans-Torresian phylogeographic relationships in three genera of Australasian snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus, and Pseudechis). Molecular Phylogenetics and Evolution 34: 1–14. Yuwono, F.B. 1998. The trade of live reptiles in Indonesia. Mertensiella 9: 9–15. Zaher, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubrid hemipenes. Bulletin of the American Museum of Natural History 240: 1–168. Ziegler, T., K.M. Philipp, and W. Bo¨hme. 1999. Zum Artstatus und zur Genitalmorphologie von Varanus finschi Bo¨hme, Horn et Ziegler, 1994, mit neuen Verbreitungsangapen fu¨r V. finschi und V. doreanus. Zoologische Abhandlungen Staatliches Museum fu¨r Tierkunde Dresden 50 (17): 267–279. Zug, G.R. 2004. Systematics of the Carlia ‘‘fusca’’ lizards (Squamata: Scincidae) of New Guinea and nearby islands. Bishop Museum Bulletin in Zoology 5: i–viii, 1–83.

................. 16157$

CH39

02-08-07 10:47:56

PS

PAGE 615

616 / Zug, G.R., and I. Ineich. 1995. A new skink (Emoia: Lacertilia: Reptilia) from the forest of Fiji. Proceedings of the Biological Society of Washington 108 (3): 395–400. Zug, G.R., L.J. Vitt, and J.P. Caldwell. 2001. Herpetology: An Introductory Biology of Amphibians and Reptiles. 2nd ed. Academic Press, San Diego. Zweifel, R.G. 1971. Results of the Archbold Expeditions. No. 96. Relationships and distribution of Genyophryne thomsoni, a microhylid frog of New Guinea. American Museum Novitates 2469: 1–13. Zweifel, R.G. 1972. Results of the Archbold Expeditions. No. 97. A revision of the frogs of the subfamily Asterophryinae Family Microhylidae. Bulletin of the American Museum of Natural History 148: 415–546. Zweifel, R.G. 1972. A review of the frog genus Lechriodus (Leptodactylidae) of New Guinea and Australia. American Museum Novitates 2507: 1–41. Zweifel, R.G. 1980. Description and relationships of a microhylid frog, Barygenys parvula, new species, from Papua New Guinea. Pacific Science 34 (3): 269–275. Zweifel, R.G. 1985. Australian frogs of the family Microhylidae. Bulletin of the American Museum of Natural History 182 (3): 265–388. Zweifel, R.G. 2000. Partition of the Australopapuan microhylid frog genus Sphenophryne with descriptions of new species. Bulletin of the American Museum of Natural History 253: 1–130. Zweifel, R.G., and F. Parker. 1989. New species of microhylid frogs from the Owen Stanley Mountains of Papua New Guinea and resurrection of the genus Aphantophryne. American Museum Novitates 2954: 1–19. Zweifel, R.G., and M. Tyler. 1982. Amphibia of New Guinea. Pp. 759–801 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Vol. 42, Monographiae Biologicae. W. Junk, The Hague. Additional information and updated species checklists are available at www.bishopmu seum.org/research/pbs/pngherps/.

................. 16157$

CH39

02-08-07 10:47:57

PS

PAGE 616

4.7. The Monitor Lizards of Papua .

.

Varanidae consists of about 80 recent species and subspecies (Bo¨hme 2003). They are all arranged in a single genus, Varanus, which is divided into nine subgenera. Eleven species are recorded for Papua (Table 4.7.1). Most of them are virtually unknown. The aim of this contribution is to introduce all species of Papua’s monitor lizards and summarize the available information. The species are grouped in a phylogenetic order, as proposed by Bo¨hme (2003).

T

Monitor Species

Argus Monitor Lizard Varanus panoptes horni The Argus Monitor Lizard is generally dark to reddish brown above, with a dorsal pattern consisting of alternating transverse rows of pale spots and larger dark blots. This pattern extends to the basal portion of the tail. The distal part of the tail is regularly banded (Bo¨hme 1988; Cogger 1994). The largest reported males grow to about 140 cm (H.-G. Horn, pers. comm.). This ground-dwelling species is restricted to the lowlands in south Papua where it inhabits a wide variety of habitats, from coastal sclerophyll forest to seasonal dry savanna. When searching for food V. p. horni ranges over large areas. They feed on insects, reptiles, birds, mammals, and even carrion (Cogger 1994). Larger specimens often stand on their hind legs to obtain a better view of their surroundings. Ritual combats have been observed in this species. In one case, the clinch phase (the bipedal stance with mutual embrace) took place in the water (Horn et al. 1994).

: Blue-tailed Monitor Lizard Varanus doreanus The Blue-tailed Monitor Lizard is a large monitor lizard, growing to 160 cm total length, eventually more (A. Allison, pers. comm.; H.-G. Horn, pers. comm.). The tail is strongly compressed and shows blue coloration interspersed with black. Adult V. doreanus have a brownish or blackish ground color with a dense speckling of yellow spots that may form larger flecks, arranged in irregular cross rows. The throat is marbled black and yellow. The coloration of juveniles and subadults is more striking (Bo¨hme et al. 2004). V. doreanus inhabits monsoon forest and dense Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

617

................. 16157$

CH40

02-08-07 10:43:15

PS

PAGE 617

Latin Varanus panoptes horni Varanus doreanus Varanus indicus Varanus jobiensis Varanus boehmei Varanus kordensis Varanus macraei Varanus prasinus prasinus Varanus reisingeri Varanus similis Varanus salvadorii

Subgenus

Varanus

Euprepiosaurus

Odatria

Papusaurus

Table 4.7.1. Papua’s Varanus species

................. 16157$

CH40

02-08-07 10:43:16

Papua Monitor Lizard

Biawak Papua

#

Biawak pohon berbintik

Biawak ekor biru Biawak Pasifik Biawak leher merah Biawak pohon bercak kuning Biawak hijau Biak Biawak pohon bercak biru Biawak hijau Biawak kuning

Biawak coklat

Indonesian

.

Spotted Tree Monitor Lizard

Blue-tailed Monitor Lizard Pacific Monitor Lizard Peach-throated Monitor Lizard Golden Speckled Tree Monitor Lizard Biak Emerald Monitor Lizard Blue Speckled Tree Monitor Lizard Emerald Monitor Lizard Yellow Tree Monitor Lizard

Argus Monitor Lizard

English

618 /

.

PS

PAGE 618

The Monitor Lizards of Papua / 619

primary rainforest, such as mixed alluvial forest and mixed hill forest (Philipp 1999a; Bo¨hme et al. 2004). Within these vegetation types V. doreanus prefers rather open areas, like natural clearings resulting from tree falls or dry river banks. V. doreanus appears to be restricted to intact, undisturbed forests and avoids humans. The Blue-tailed Monitor Lizard exhibits remarkable intraspecific niche segregation. Adults are largely terrestrial, whereas juveniles and subadults use the upper strata of the canopy (Philipp 1999a). This strategy is likely to reduce food competition between young and adult stages of V. doreanus. A remarkably high density is reported by Philipp (1999a). Thirteen individuals were encountered within an area of about 600 m2 around a fallen tree in a seasonally dry river bed. The stomachs of four preserved museum specimens contained insects, skinks, skink eggs, and one small snake. The stomachs of four other specimens were empty (Table 4.7.2). A juvenile V. doreanus with a snout-vent length of approximately 23 cm regurgitated a skink (presumed to be Lygosoma sp.) that had a headbody length of almost 17 cm (!) (K. Philipp, pers. obs.). V. doreanus is found throughout the lowlands of Papua’s mainland and the offshore islands, Salawati and Biak.

Pacific Monitor Lizard Varanus indicus V. indicus is rather eurytopic occupying a wide variety of habitats, but is most common in (salt) water-influenced habitats such as beach woodland, littoral forests, and mangroves. Infrequently, this species is found in forests further away from the coast. V. indicus inhabits primary as well as secondary forests and sometimes can be seen in the vicinity of villages (Philipp 1999a; Dryden and Ziegler 2004). Because of its adaptations to saltwater, the Pacific Monitor Lizard has an enormous geographic distribution, from the Moluccas to the Solomon Islands and from Australia to Micronesia. Across its large range, V. indicus shows considerable color variation. The populations in Papua are rather homogeneous in coloration. They show a dorsal color pattern consisting of irregularly scattered small whitish to yellowish dots. The tail is strongly compressed and banded dark with intervening whitish to yellowish bands. The coloration of juveniles and subadults is more striking. The largest individuals in Papua grow to about 150 cm total length. Stomach content analyses of preserved museum vouchers revealed that V. indicus feeds on Onchidiida, Malacostraca, insects, and reptiles (Philipp 1999b). Pacific Monitor Lizards are opportunistic feeders, eating almost everything they can catch, but the high percentage of crustaceans (63%; see Table 4.7.2) in their diet is remarkable. Around villages V. indicus often feed on chicks of domestic fowl, so this species is often hunted in the vicinity of villages. On several occasions eagles have been observed to hunt and feed on V. indicus (Beehler et al. 1992; K. Tindige, pers. comm.).

................. 16157$

CH40

02-08-07 10:43:16

PS

PAGE 619

#

.

620 /

.

Table 4.7.2. Composition of nutrition of 35 monitor lizards of the Varanus indicus group Prey items

V. doreanus

V. indicus

V. jobiensis

Mollusca Onchidiida

—

9%

—

Aviculariidae

—

—

10%

Atyidae

—

8%

—

Brachyura

—

55%

—

Arthropoda Arachnida Malacostraca Insecta

Blattodea

—

5%

Ensifera

—

6%

26%

Caelifera

14%

2%

10%

Rhynchota

—

—

2%

Coleoptera

14%

4%

26%

Hymenoptera

—

—

2%

Lepidoptera

—

—

2%

Teleostei

—

6%

—

Amphibia

Anura

—

—

12%

Reptilia

Scincidae

29%

—

—

Chordata Osteichthyes

Eier

29%

9%

5%

Serpentes

14%

2%

—

8 (4)

20 (4)

7 (0)

Number of specimens

Note: Percentages indicate percent of stomach contents by item. The last column is the number of investigated museum vouchers; the number in parentheses indicates the number of specimens with an empty stomach.

V. indicus is a skillful swimmer (Figure 4.7.1). Alarmed animals often take refuge in water and disappear there. Pacific Monitors are also good climbers. A gravid female collected in August 1907 contained six eggs (ca 55 30 mm; Philipp 1999b). V. indicus is reported to inhabit huge areas of the mainland of Papua, and is recorded for the Papuan islands Rawak, Waigeo, Salawati, and Biak. A distribution on many more islands around Papua might be expected.

Peach-throated Monitor Lizard Varanus jobiensis The Peach-throated Monitor Lizard is a strikingly colored endemic species of New Guinea. The dorsum is dark olive to black, with numerous tiny light spots, ar-

................. 16157$

CH40

02-08-07 10:43:17

PS

PAGE 620

The Monitor Lizards of Papua / 621

Figure 4.7.1. V. indicus swimming in a brackish creek. ranged in more or less distinct broad bands. The tail is blue-turquoise banded (Figure 4.7.2). The slender head of this species is solid slate gray, often with a blue touch. The throat shows a remarkable pinkish, reddish, purple, or orange hue, leading to the vernacular name Peach-throated Monitor Lizard. In juveniles the coloration is more intense, and the bands of light markings on the dorsum are generally more widely separated (Figure 4.7.3). This slender species grows to 120 cm total length. Hatchlings are about 27 cm long. Tail length averages 1.5 to 1.8 times snout-vent length. V. jobiensis is found throughout the lowlands of Papua’s mainland and the offshore islands of Waigeo, Salawati, Biak, and Yapen (Philipp et al. 2004). The Peach-throated Monitor Lizard inhabits mixed alluvial forest and mixed hill forests and prefers the dense microhabitats found there. V. jobiensis is frequently encountered on the forest floor, sun basking or searching for food. If the animals are alarmed they seek refuge by climbing trees and hiding there in holes. This species is often encountered resting on tree trunks, expanding and displaying its strikingly colored throat. In the gloom of the forest interior this is a conspicuous signal that may play an important role in intra- or interspecific communication (Philipp 1999a). V. jobiensis regularly patrols along small brooks, obviously scavenging there. Small fish and shrimps occur at such locations and provide an easy available food source. Stomach content analyses of preserved museum vouchers revealed that V. jobiensis feeds on large spiders, insects, frogs, and reptile eggs (Table 4.7.2). It seems that V. jobiensis is active all year round (Philipp et al. 2004).

................. 16157$

CH40

02-08-07 10:43:29

PS

PAGE 621

622 /

#

.

.

Figure 4.7.2. Adult V. jobiensis resting on a trunk of a rain forest tree.

: Golden Speckled Tree Monitor Lizard Varanus boehmei The Golden Speckled Tree Monitor Lizard has been discovered and described just recently (Jacobs 2003). This tree-dwelling, slender species is characterized by its dorsal body coloration: irregular rows of golden ocelli (eye spots) on a blackish ground color. The largest specimens known today grow to about 90 cm, with the tail comprising two-thirds of its length. Varanus boehmei is endemic to Waigeo Island and has not been observed in the wild until now, thus the knowledge of its ecology is extremely limited. According to reptile traders, this species is mainly found in the eastern parts of Waigeo, inhabiting coastal areas. In captivity, this

................. 16157$

CH40

02-08-07 10:43:45

PS

PAGE 622

The Monitor Lizards of Papua / 623

Figure 4.7.3. Juvenile V. jobiensis exhibiting its striking coloration. species is arboreal and a very adept climber, using its prehensile tail as a fifth leg (Jacobs 2003).

Biak Emerald Monitor Lizard Varanus kordensis The Biak Emerald Monitor Lizard has an intense pattern of olive colored ocelli on a black ground color. In the overall appearance the coloration looks like a black reticulation with wide olive green interspaces. This pattern also covers the limbs and tail. In juveniles the coloration is more intense. With a total length of 80 cm, Varanus kordensis is among the smallest species of the V. prasinus group. The prehensile tail is two times longer than the body and is used as an additional limb when climbing in branches. V. kordensis is endemic to Biak Island where it inhabits different forest types in the lowlands: littoral forest and mixed alluvial forest (N. Ohorella, pers. comm.). This species is highly arboreal, but can be seen regularly on the forest floor searching for food, such as insects or other arthropods. Further ecological data are not available for this species from the wild. In captivity clutch size varies from three to four eggs, which hatch after about 190 days (Jacobs 2004).

Blue Speckled Tree Monitor Lizard Varanus macraei This spectacularly colored monitor has just recently been discovered and described based on specimens originating from the pet trade (Bo¨hme and Jacobs 2001). Blue oblique rows of ocelli on a black background form the diagnostic body coloration of Varanus macraei. The Blue Speckled Tree Monitor is the largest species among

................. 16157$

CH40

02-08-07 10:43:59

PS

PAGE 623

624 /

#

.

.

the V. prasinus species group, growing to about 110 cm. Two-thirds of the total length are made up by the prehensile tail. V. macraei is a diurnal, tree-dwelling monitor lizard endemic to Batanta Island. In captivity this species was observed to lay a single clutch of three eggs, which hatched after 159 days (Jacobs 2004). No field studies have been carried out to learn more about this species in the wild. The sparse information available suggests that V. macraei is rare on Batanta (MacRae, cited in Bo¨hme and Jacobs 2001). Despite this, V. macraei is exported in huge numbers from Indonesia. In 2003, 114 specimens were shipped from Indonesia for the pet trade (UWCTDB 2005). This number may not reflect the actual number of collected specimens, which is undoubtedly higher, since many specimens die before they are shipped. Moreover, the animals traded within Indonesia are not included in these figures. A first field study is needed to investigate the status of V. macraei on Batanta and the possible threats of the reptile trade on this attractive species.

Emerald Monitor Lizard Varanus prasinus prasinus The Emerald Monitor Lizard is a brilliant green tree-dwelling species (Figure 4.7.4). Many specimens show V-shaped, black dorsal stripes. In juveniles this black pattern is more prominent, and fades as the specimens grow. In some adult V. p. prasinus the black pattern almost disappears. V. p. prasinus grows to 90 cm total length, about two-thirds of it are tail. Emerald Monitor Lizards are found in appropriate habitat over much of Papua and its adjacent island Salawati, from sea level to at least 830 m in elevation (Greene 2004). The species inhabits a wide variety of habitats, from monsoon, rain, and palm forests to lagoons (Cogger 1994). Occasionally Emerald Monitor Lizards are seen in remote garden areas climbing on fruiting mango trees and banana plants (Philipp, pers. obs.). V. p. prasinus is a highly specialized tree-dwelling lizard, using its prehensile tail as fifth limb. Furthermore, the tail is used as a counterbalancing steering gear for climbing and jumping. Unlike other arboreal varanid species of Papua that prefer microhabitats along tree trunks and huge branches (e.g., adult V. jobiensis), V. p. prasinus inhabits dense vegetation with predominantly small trees, lots of vines, and leafy vegetation and the periphery of treetops (Greene 1986; Philipp, pers. obs.). Normally V. p. prasinus is a slow-moving lizard, but when alarmed it can flee quickly. V. p. prasinus feeds mainly on arthropods, usually relatively small katydids. Occasionally, rodents and large walking stick insects are taken too. In one case a walking stick evidentially was dismembered before ingestion as none of the insect’s spinous legs were encountered in the stomach, indicating that diverse prey handling tactics occur (Greene 1986). Mertens (1971) reported a captive V. p. prasinus that ate bananas. Vegetarian food items were not encountered in preserved wild caught animals (Greene 1986). Only two of all known monitor lizard species (V. mabitang and V. olivaceus) are known to be frugivorous (Auffenberg 1988, Gaulke 2004). Observations in garden areas (see above) may well indicate that at least

................. 16157$

CH40

02-08-07 10:43:59

PS

PAGE 624

The Monitor Lizards of Papua / 625

Figure 4.7.4. V. p. prasinus climbing a tree in captivity. occasionally V. p. prasinus feeds on ripe fruits or drinks the fruit juice. Greene (1986) reports two neonate Emerald Monitor Lizards that hatched in termite nests, an observation that has not been replicated. In captivity, clutches of up to six eggs are laid and they hatch within 170 days (Jacobs 2004). The second subspecies, V. p. beccarii, inhabits the Aru Islands and is uniformly black.

................. 16157$

CH40

02-08-07 10:44:15

PS

PAGE 625

626 /

#

.

.

Yellow Tree Monitor Lizard Varanus reisingeri The Yellow Tree Monitor Lizard is the latest in a series of discoveries made on specimens obtained through the pet trade. The general appearance is similar to the other members of the V. prasinus group, but the characteristic dorsal color pattern consists of yellow-greenish spots and ocelli, ordered in about 16 cross rows on a black background. The largest known individual grew to 76 cm total length (28 cm head-to-body length, 48 cm tail length). Currently, no field data are available for V. reisingeri. Observations made on captive specimens suggest that this species is arboreal. Available information implies that V. reisingeri is endemic to Misool Island (Eidenmu¨ller and Wicker 2005).

: Spotted Tree Monitor Lizard Varanus similis This pygmy monitor shows a grayish-black ground color with an irregular pattern of lighter colored, ill-defined ocelli or spots. Varanus similis is the smallest species of monitor lizard from Papua, growing to about 60 cm total length. The tail is approximately 1.5 times as long as snout-vent. The prehensile tail is round in section and covered with slightly keeled scales. The Spotted Tree Monitor Lizard is arboreal; it can be seen basking or foraging on trunks or branches of trees. Sometimes it descends to feed on insects and lizards on the ground. This species shelters in hollow limbs, holes, or under loose bark (Cogger 1994). Ritual combat of male V. similis takes place on the ground, and does not show the bipedal clinch phase (Horn 1985; Horn et al. 1994). V. similis is found in the southern, drier parts of Papua. The overall distribution of the V. scalaris species group, to which V. similis belongs, corresponds roughly to the distribution of Eucalyptus tetrodonta (Smith et al. 2004). In captivity the clutch size varies between 9 and 12 eggs. The incubation time is some 130 days. Hatchlings are about 13 cm long (Berghof 2001). The systematics of the V. timorensis group is still under debate, and we expect that changes to the taxonomy of this group will be made in the future (Smith et al. 2004).

Papua Monitor Lizard Varanus salvadorii The Papua Monitor Lizard is perhaps New Guinea’s most remarkable varanid. Often it is referred to as the longest living lizard, growing to 4 m total length or even more (e.g., Allison 1982; MacKay 1992). Such reports have never been substantiated. The largest reliable measurement of a museum voucher is 2.65 m (Bo¨hme and Ziegler 1997a; Schmicking and Horn 1997). Nevertheless, numerous

................. 16157$

CH40

02-08-07 10:44:16

PS

PAGE 626

The Monitor Lizards of Papua / 627

reports (from various reliable sources) of giant individuals of this New Guinea species have been made, suggesting that such giants really exist. Varanus salvadorii is a dark brown to black species with a yellow pattern, consisting of small dots, larger blots, or ocelli (Figure 4.7.5). In juveniles and subadults this coloration is especially striking. The tail is extraordinarily long, up to 2.7 times of the snout-vent length and roundish to triangular in shape (Horn 2004). The snout of V. salvadorii is large and bulbous, the teeth are unusually long and sharp, and several gular folds are present (Mertens 1950). None of these unique characters have been satisfactorily explained, but they are presumably associated with the unique habits of the Papua Monitor Lizard. Despite its large size, V. salvadorii is a mostly arboreal species, able to climb and leap with high speed in the forest canopy (Cann, cited in Horn 2004). The species is assumed to feed on insects, frogs, reptiles, birds and their eggs, and even mammals (Bennett 1995). Mackay (1992) reported that these lizards often lie on branches overhanging trails to ambush birds, reptiles, or small mammals. Knowledge of the geographic distribution of the Papua Monitor Lizard is still very fragmentary. It seems that this species lives in most of Papua’s lowlands and the nearby offshore island of Salawati (Horn 2004; Horn et al. in prep.), at least up to an altitude of 540 m above sea level (Mertens 1942). There, V. salvadorii inhabits rainforest vegetation such as mixed alluvial and mixed hill forest types, as well as riparian forests. V. salvadorii is assumed to be a good swimmer and on several occasions individuals have been observed in forests along rivers. V. salvadorii appears to be restricted to intact undisturbed primary forests and strictly avoids contact with humans (Mackay 1992; Horn 2004). In captivity, clutch size varies between four and twelve eggs, which hatch after

Figure 4.7.5. Tongue-flicking subadult V. salvadorii with the prominent snout.

................. 16157$

CH40

02-08-07 10:44:25

PS

PAGE 627

628 /

#

.

.

about 180 to 250 days, and the hatchlings show remarkable variation in weight: 47 to 68 g (Horn 2004).

Zoogeography At this time 11 species of monitor lizards are recorded for Papua. Whether Varanus cerambonensis—a species widely distributed in the south Moluccas—can also be found in Papua needs further investigation (see Philipp et al. 1999). The same is true for V. finschi. This species is recorded for the Bismarck Archipelago, the adjacent parts of Papua New Guinea, the Kai Islands, and north Australia (Ziegler et al. 1999). The total expected number of taxa inhabiting New Guinea is not clear at the time of this writing. We are aware of at least four more taxa in Papua still waiting to be described: one of the V. prasinus group, one of the V. indicus group, and two of the subgenus Odatria. These are either sister forms of known species or varieties completely unknown to science. Summing up, a total of 17 Varanus taxa inhabiting Papua might be expected. Of the 11 confirmed species of Papua, four (36%) are endemic Papua species (V. boehmei, V. kordensis, V. macraei, V. reisingeri) and seven (64%) are endemic species of New Guinea (V. boehmei, V. panoptes horni, V. jobiensis, V. kordensis, V. macraei, V. reisingeri and V. salvadorii). One species (V. indicus) is shared with Micronesia in the north, seven species (V. doreanus, V. indicus, V. jobiensis, V. panoptes horni, V. prasinus prasinus, V. salvadorii and V. similis) are shared with Papua New Guinea in the east, four species (V. doreanus, V. indicus, V. p. prasinus and V. similis) are shared with Australia (at least the northern Torres Strait islands), and one species (V. indicus) is shared with the Moluccas in the west (Figure 4.7.6). The monotypic subgenus Papusaurus, with its species V. salvadorii, is an endemic subgenus for New Guinea and the offshore island Salawati. New Guinea is the geographical center of distribution of the subgenus Euprepiosaurus. This subgenus comprises 20 taxa. Eight taxa can be found in Papua; 12 taxa in New Guinea and its surrounding offshore islands (including the Aru Islands). These are the eight Papuan taxa plus V. bogerti, V. finschi, V. p. beccarii, and V. telenesetes. The remaining species of the subgenus Euprepiosaurus occur in the Moluccas (V. caerulivirens, V. cerambonensis, V. melinus, V. yuwonoi, V. zugorum), in the Solomon Islands (V. juxtindicus, V. spinulosus), and in Australia (V. keithhorni). Table 4.7.3 summarizes the current published information on the geographic distribution of the different Papuan species of monitor lizards (Pianka et al. 2004; Richards et al. 2002a,b). The highest species diversity is recorded for the districts of Sorong, Merauke, and Jayapura. Six species have small distribution areas, occurring in only a single district of Papua. The remaining five species are more widespread and occur in at least five different districts.

Endemism The four endemic species V. boehmei, V. kordensis, V. macraei, and V. reisingeri are quite rare (Bo¨hme and Jacobs 2001; N. Ohorella, pers. comm.). In combination

................. 16157$

CH40

02-08-07 10:44:27

PS PAGE 628

The Monitor Lizards of Papua / 629

Figure 4.7.6. Number of Papuan Varanus species found in the adjacent regions, Micronesia, Papua New Guinea, Australia and Moluccas. The digits indicate the number of Varanus species inhabiting both neighboring regions (e.g., four species are shared between Papua and Australia). with their limited geographic distribution (Waigeo has a size of 3153.7 km2, Biak 1904.3 km2, Batanta 455.9 km2, and Misool 2033.6 km2), these four species are highly threatened by habitat destruction and exploitation (UNEP 2005). When compared with V. komodoensis, whose distribution area covers about 957 km2 (1950 km2 if transit areas are included) on five different islands, (Auffenberg 1981; Ciofi 2004) it is obvious that further investigations on distribution, population, and ecology of Papua’s four endemic species are essential. All four endemic species—as well as all the other species found on Papua—are listed in CITES Appendix 2 and show up regularly in the pet trade. V. komodoensis reaps the benefits of higher protection (listed in CITES Appendix 1). This level of protection should be considered for V. boehmei, V. kordensis, V. macraei, and V. reisingeri as well, once detailed population and ecological studies are completed. The main threat to these taxa arises from (illegal) logging. In recent years, (illegal) activity by logging companies has grown in Papua. One of the centers of activities is Sorong, and these companies focus on the Raja Ampat Archipelago (TELA 2005). Within Papua, the Raja Ampat Archipelago is an area with high varanid ende-

................. 16157$

CH40

02-08-07 10:44:41

PS PAGE 629

Varanus similis

Varanus salvadorii

Euprepiosaurus

Odatria

Papusaurus

Sorong

................. 16157$

CH40

02-08-07 10:44:43

PS

4

x

x

3

x

x x

Biak-Numfoor 4

x

x x x

Yapen-Waropen 2

x

x

Nabire 2

x

x x

Paniai 0

Mimika 2

x

x

Jayapura 5

x

x

x x x

Mt Jaya 0

Jayawijaya 1

x

7

x

x

x

x x x

x

Merauke

Note: District kabupaten. ‘‘x’’ indicates species is present in that district. For some districts (especially districts of the lowlands) higher total numbers of species might be expected. The low numbers result from a general lack of information for these remote areas. Source: Pianka et al. (2004); Richards et al. (2002a, 2002b).

8

Manokwari x x

1

Total 7

1

6 7 5 1 1 1 7 1

#

x

x x x

x x x x

.

Total

Varanus panoptes horni

Varanus doreanus Varanus indicus Varanus jobiensis Varanus boehmei Varanus kordensis Varanus macraei Varanus prasinus prasinus Varanus reisingeri

Varanus

Species

Subgenus

Fakfak

Table 4.7.3. Distribution of Varanus species in districts of Papua 630 /

.

PAGE 630

The Monitor Lizards of Papua / 631

mism, as it is for many other taxa as well (e.g., Chapters 2.4, 2.5, and 4.9). V. boehmei, V. macraei, V. reisingeri, and at least one undescribed taxa of monitor lizards are endemic to single islands of this archipelago.

Ecology Eight of Papua’s monitor lizard species are inhabitants of intact, undisturbed rainforests. One species (V. indicus) inhabits rainforests as well, but prefers mangroves and other (salt)water-influenced habitats. The remaining two species (V. panoptes horni and V. similis)—restricted to south New Guinea—are inhabitants of drier vegetation types, such as savanna. Two species (V. indicus and V. p. prasinus) are found rarely in the vicinity of human settlements. All other species seem to avoid humans completely. In many areas of Papua five species of monitor lizards are sympatric: V. doreanus, V. indicus, V. jobiensis, V. p. prasinus, and V. salvadorii. The niche segregation strategy of these species is not fully understood, although Philipp (1999a) gave a first insight on this topic for the three species of the V. indicus group. Where all three species can be found sympatrically, V. indicus prefers habitats close to water bodies, V. doreanus prefers rather open habitats within rainforests, and V. jobiensis favors densely vegetated areas (Figurre 4.7.7). Furthermore V. jobiensis is more arboreal than V. doreanus. Within its arboreal habitat V. jobiensis seem to prefer areas close to the trunks or huge branches. In contrast, V. p. prasinus clearly prefers arboreal habitats consisting of fine branches or vines (Greene 1986), as found in the outer area of tree canopies. At present we do not know how V. salvadorii fits into this framework. Beside the described spatial axis, other factors such as food (see Table 4.7.2), temperature, and time might act as further niche partitioning

Figure 4.7.7. Idealized profile diagram of certain vegetation types of mainland Papua and the indwelling Varanus species. ◆: V. doreanus, : V. indicus, : V. jobiensis, O: V. prasinus prasinus, : V. similis, ✪: V. salvadorii, b: V. panoptes horni. a: beach woodland, b: littoral forest, c: mangrove forest, d: mixed alluvium forest, e: early succession phase after windbreak, f: mixed hill forest, g: mixed savannah. Line drawing: Yose Kadrin.

................. 16157$

CH40

02-08-07 10:44:48

PS

PAGE 631

632 / kai m . p h i l i p p & d e v i p . p h i l i p p

axes. Further investigations on the niche segregation strategies of the five sympatric species, based on the competitive exclusion principle, may prove rewarding. Papua lacks any large mammalian predators, except some introduced species, such as Canis familiaris, Felis catus (Flannery 1995; Chapter 4.10). Of the remaining native mammal species, only 6% are aquatic insectivore/carnivores and 5% are carnivores (Flannery 1995). Without doubt, monitor lizards are among the top predators of Papua, sharing this honor with the New Guinea Harpy-Eagle and some of the larger pythons. Therefore the monitor lizards play an important role in the food chain in Papua.

Conservation

live reptile trade Most varanid species are popular in the international pet trade (Figure 4.7.8). This is especially true for the Papuan species (Yuwono 1998). According to Indonesian law (SK Mentan No. 327/Kpts/Um/5/1978, SK Mentan No. 716/Kpts/Um/10/ 1980, SK Menhut No. 301/Kpts–II/1991, Peraturan Pemerintah RI No. 7 Tahun 1999–Lampiran), V. indicus, V. panoptes horni (as V. gouldii), V. prasinus, and V. similis (as V. timorensis) are the only protected Papuan monitor lizard species (SYDI 2004; REIN 2005). These species are not allowed to be traded within or exported from Indonesia without special permits. In order to circumvent these national laws, other names are used. Members of the subgenus Euprepiosaurus show up in the pet trade using the export name V. kalabeck (e.g., Bo¨hme and Jacobs 2001). ‘‘Kalabeck’’ is a nomen nudum and therefore not applicable for any species of monitor lizard. The same is true for the name douarrha (Bo¨hme et al. 1994). Traders use these doubtful names to circumvent national and international quotas for the reptile trade (e.g., V. kordensis was regarded as a junior synonym of V. prasinus until 2002; Jacobs 2002). Nevertheless 1,639 live specimens of Green Tree Monitor Lizards, mainly the protected V. p. prasinus, were exported from Indonesia between 1987 and 1994, using the invalid name V. kordensis (UWCTDB 2005). Similar deceptions are known to occur with the names kalabeck and douarrha for members of the V. indicus group (e.g., Bo¨hme and Ziegler 1997b). These names must not be accepted by any national or international authority, particularly CITES, involved in the trade of wildlife. With all the newly described species and other recent taxonomic changes, an amendment of the relevant Indonesian and international laws seems to be required.

leather industry The skins of Papuan Varanus species seem to play a minor role in the leather industry, compared to some heavily exploited species (i.e., V. salvator and V. bengalensis; Luxmoore and Groombridge 1990). Luxmoore and Grooombridge (1990) report that a minimum of 46,507 V. indicus, 10 V. jobiensis (as V. karlschmidti), 61

................. 16157$

CH40

02-08-07 10:44:49

PS

PAGE 632

The Monitor Lizards of Papua / 633

Figure 4.7.8. V. doreanus and V. jobiensis at a reptile trader in Papua. Note the unsanitary conditions and the numerous injuries of the animals. V. salvadorii, and 12,488 V. timorensis (maybe reflecting a mix of V. timorensis and V. similis) leather items were manufactured from the varanid skins between 1975 and 1986. Skins of various Varanus species are sometimes seen on traditional drums that are sold to tourists (figures on quantity are not available). As all monitor lizards are protected at least under CITES Appendix 2, these drums need appropriate documents to be legally traded.

prospects The monitor lizards of Papua inhabit a wide variety of available habitats. For the well-being of these spectacular species it is essential to protect their habitats. This can be done by establishing a network of nature conservation areas, as has been done in many parts of Papua, and must be followed by strong management of these protected areas. Furthermore, the national and international reptile trade of Papua’s monitor

................. 16157$

CH40

02-08-07 10:45:05

PS

PAGE 633

634 / kai m . p h i l i p p & d e v i p . p h i l i p p

lizards should be monitored more closely. The effect of the trade on local populations should be documented and the quota for each species must be carefully set and adjusted on a regularly basis. Without doubt the most dangerous and ongoing threat to Papua’s wildlife, including the monitor lizards, is the increasing legal and illegal logging activities in Papua.

Acknowledgments We are very grateful to Prof. Dr. W. Bo¨hme and Prof. Dr. H.-G. Horn for numerous inspiring discussions; Prof. Dr. Horn also provided useful comments on a draft of the manuscript. We thank K. Tindige and N. Ohorella for sharing their observations in Papua and to Yose Kadrin for preparing Figure 4.7.7.

Literature Cited Allison, A. 1982. Distribution and ecology of New Guinea lizards. J.L. Gressitt: Monographiae Biologicae 42 (2): 803–813. Auffenberg, W. 1981. The Behavioral Ecology of the Komodo Monitor. University Presses of Florida, Gainesville. Auffenberg, W. 1988. Gray’s Monitor Lizard. University Presses of Florida, Gainesville. Beehler, B.M., W. Crill, B. Jefferies, and M. Jefferies. 1992. New Guinea Harpy-Eagle attempts to capture a monitor lizard. Emu 92: 246–247. Bennett, D. 1995. A Little Book of Monitor Lizards. Viper Press, Aberdeen. Berghof, H.-P. 2001. Die Pflege und Vermehrung des kleinen neuguineischen Baumwaranes Varanus (Odatria) similis Mertens, 1958. Herpetofauna (Weinstadt) 23 (135): 5–13. Bo¨hme, W. 1988. Der Arguswaran (Varanus panoptes Storr, 1980) auf Neuguinea: V. panoptes horni ssp. n. (Sauria: Varanidae). Salamandra 24: 87–101. Bo¨hme, W. 2003. Checklist of the living monitor lizards of the world (family Varanidae). Zool. Verh. Leiden 341: 3–43. Bo¨hme, W., and H.J. Jacobs. 2001. Varanus macraei sp. n., eine neue Waranart der V. prasinus-Gruppe aus West Irian, Indonesien. Herpetofauna (Weinstadt) 23 (133): 5–10. Bo¨hme, W., and H.J. Jacobs. 2004. Varanus macraei. Pp. 212–214 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Bo¨hme, W., K.M. Philipp, and T. Ziegler. 2004. Varanus doreanus. Pp. 168–171 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Bo¨hme, W., and T. Ziegler. 1997a. Großwarane im Museum Koenig, mit Bemerkungen zu Afrikas gro¨ßter Echse. Tier u. Mus. 5 (3): 65–74. Bo¨hme, W., and T. Ziegler. 1997b. Varanus melinus sp. n., ein neuer Waran aus der V. indicus-Gruppe von den Molukken, Indonesien. Herpetofauna (Weinstadt) 19 (111): 26–34. Ciofi, C. 2004. Varanus komodoensis. Pp. 197–204 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Cogger, H. 1994. Reptiles and Amphibians of Australia. Cornell University Press, Ithaca, New York.

................. 16157$

CH40

02-08-07 10:45:05

PS

PAGE 634

The Monitor Lizards of Papua / 635 Dryden, G., and T. Ziegler. 2004. Varanus indicus. Pp. 184–188 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Eidenmu¨ller, B., and R. Wicker. 2005. Eine weitere neue Waranart aus dem Varanus prasinus-Komplex von der Insel Misol, Indonesien. Sauria 27 (1): 3–8. Flannery, T.F. 1995. The Mammals of New Guinea. Reed Books, Chatswood. Gaulke, M. 2004. Varanus mabitang. Pp. 208–211 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Greene, H.W. 1986. Diet and arboreality in the emerald monitor, Varanus prasinus, with comments on the study of adaptation. Fieldiana (Zoology), n.s. 31: 1–12. Greene, H.W. 2004. Varanus prasinus. Pp. 225–229 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Horn, H.-G. 1985. Beitra¨ge zum Verhalten von Waranen: die Ritualka¨mpfe von Varanus komodoensis Ouwens, 1912 und V. semiremex Peters, 1869 sowie die Imponierphasen der Ritualka¨mpfe von V. timorensis timorensis Gray, 1931 und V. t. similis Mertens, 1958 Sauria: Varanidae. Salamandra 21: 169–179. Horn, H.-G. 2004. Varanus salvadorii. Pp. 234–243 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Horn, H.-G., M. Gaulke, and W. Bo¨hme. 1994. New data on ritualized combats in monitor lizards (Sauria: Varanidae) with remarks on their function and phylogenetic implications. Zool. Garten N.F. 64 (5): 265–280. Jacobs, H.J. 2002. Zur morphologischen Variabilita¨t der nominellen Smaragdwaran-Taxa Varanus prasinus (Schlegel, 1839) und Varanus kordensis (A. B. Meyer, 1874) mit Bemerkungen zur Erstzucht des letzteren. Herpetofauna (Weinstadt) 24 (137): 21–34. Jacobs, H.J. 2003. A further new emerald tree monitor lizard of the Varanus prasinus species group from Waigeo, West Irian (Squamata: Sauria: Varanidae). Salamandra 39 (2): 65–74. Jacobs, H.J. 2004. Varanus kordensis. Pp. 205–207 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Luxmoore, R., and B. Groombridge. 1990. Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the CITES Secretariat. World Conservation Monitoring Centre, Cambridge. Mackay, R.D. 1992. Neuguinea. Time-Life, Amsterdam. Mertens, R. 1942. Die Familie der Warane (Varanidae). Dritter Teil: Taxonomie. Abhandlungen Senckenbergischen Naturforschenden Gesellschaft 466: 235–391. Mertens, R. 1950. Notes on some Indo-Australian monitors (Sauria: Varanidae). American Museum Novitates 1456: 1–7. Philipp, K.M. 1999a. Niche partitioning of Varanus doreanus, V. indicus and V. jobiensis in Irian Jaya: preliminary results. Pp. 307–316 in Horn, H.-G., and W. Bo¨hme (eds.) Advances in Monitor Research II. Mertensiella 11: 307–316. ¨ kologie des PazifikwaranPhilipp, K.M. 1999b. Zur Systematik, Zoogeographie und O Artkomplexes (Reptilia: Squamata: Varanidae). M.A. thesis, University of Munich. Philipp, K.M., W. Bo¨hme, and T. Ziegler. 1999. The identity of Varanus indicus: redefinition and description of a sibling species coexisting at the type locality (Sauria: Varanidae, Varanus indicus group). Spixiana 22 (3): 273–287.

................. 16157$

CH40

02-08-07 10:45:06

PS

PAGE 635

636 / kai m . p h i l i p p & d e v i p . p h i l i p p Philipp, K.M., T. Ziegler, and W. Bo¨hme. 2004. Varanus jobiensis. Pp. 189–192 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Pianka, E.R., D.R. King, and R.A. King (eds.). 2004. Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Richards, S., D.T. Iskandar, and B. Tjaturadi. 2002a. Amphibians and reptiles of the Dabra area, Mamberamo River Basin, Papua, Indonesia. In S.J. Richards and S. Suryadi, A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25: 73–75. Richards, S., D.T. Iskandar, B. Tjaturadi, and A. Krishar. 2002b. Amphibians and reptiles of the Yongsu area, Papua, Indonesia. In S.J. Richards and S. Suryadi, Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25: 76–79. Schmicking, T., and H.-G. Horn. 1997. Beobachtungen bei der Pflege und Nachtzucht des Papuawarans, Varanus salvadorii (Peters and Doria 1878). Herpetofauna (Weinstadt) 19 (106): 14–23. Smith, L.A., S.S. Sweet, and D.R. King. 2004. Varanus scalaris. Pp. 451–461 in Pianka, E.R., D.R. King, and R.A. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana. Yuwono, F.B. 1998. The trade of live reptiles in Indonesia. In Erdelen, W. (ed.) Conservation, trade and sustainable use of lizards and snakes in Indonesia. Mertensiella 9: 9–15. Ziegler, T., K.M. Philipp, and W. Bo¨hme. 1999. Zum Artstatus und zur Genitalmorphologie von Varanus finschi Bo¨hme, Horn et Ziegler, 1994, mit neuen Verbreitungsangaben fu¨r V. finschi und V. doreanus (Meyer, 1874) (Reptilia: Sauria: Varanidae). Zool. Abh. Staatl. Mus. f. Tierkd. Dresden 50 (2): 267–279.

Websites Cited REIN 2005: Reptiles of Indonesia: http://www.nature-conservation.or.id/reptiles.html (accessed 19 Feb. 2005). SYDI 2004: Satwa yang Dilindungi: http://www.geocities.com/bksda_jb1/index_files/ lain_files/satwa.html (accessed 19 Dec. 2004). UNEP 2005: UNEP ISLANDS: http://islands.unep.ch/ (accessed 10 Feb. 2005). UWCTDB 2005: UNEP-WCMC CITES Trade Database: http://sea-stour.unep-wcmc.org/ citestrade/ (accessed 4 Sept. 2005). TELA 2005: The Last Frontier: http://www.telapak.org/publikasi/download/ the_last_frontier_en.pdf (accessed 20 Feb. 2005).

................. 16157$

CH40

02-08-07 10:45:06

PS

PAGE 636

4.8. Fishes of Papua gerald r. allen i s he s r e pr e s e nt the largest assemblage of vertebrates in Papua, with a very conservative estimate of 2,650 species. Indonesia forms an integral part of the much heralded ‘‘Coral Triangle,’’ the world’s center for marine biodiversity; fishes and marine invertebrates are abundant in Papuan seas. The majority of species occur on coral reefs, which are well developed in the north at Cenderawasih Bay and around Biak, and in the extreme west in the Raja Ampat Islands. There is also reef development in the vicinity of Jayapura, and along the southwestern coast off the Fakfak Peninsula and between Kaimana Bay and Etna Bay. There still remains a need for basic ichthyological surveys throughout much of Papua, particularly at the little-known southwestern area. Allen and Adrim (2002) listed 1,362 coral reef species from Papuan seas, and it is conservatively estimated that 1,500 species, or about 73% of the entire Indonesian reef fauna, occur around Papua. Most of this chapter is devoted to the largest two components of the fauna: first, the reef fishes, and then the approximately 400 species of freshwater fishes, both of which are relatively well studied. Based mainly on species that have already been recorded from the adjacent seas of Papua New Guinea (Kailola 1987, 1991), it is estimated that at least an additional 750 species can be expected in Papua, including various elasmobranchs (60 species), deep sea fishes (200), pelagics (70), inhabitants of deep reefs below 60–70 m (70), and fishes from soft bottom and miscellaneous inshore, non-reef habitats (350).

F

Reef Fishes

historical background The first collection of marine fishes in New Guinea dates back to the voyage of the French vessel L’Uranie in 1818–1819. About 30 species were collected from the Raja Ampat Islands by the surgeon-naturalists Quoy and Gaimard. Descriptions of these species were published on their return to France in 1824. Ever since this first voyage there has been a more or less steady stream of scientific explorers and collectors visiting the island from France, Great Britain, the Netherlands, Germany, Australia, and America (Chapter 1.2). Australian researcher Ian S. R. Munro published a checklist of the known fishes of New Guinea in 1956, which included approximately 1,350 marine species. More recently, Kailola (1987–1991) published an updated list incorporating all new records appearing since Munro’s work. There has been limited interest in the coral reef fishes of Papua, with the exception of those from the Raja Ampat Islands. Lying off the extreme western end of New Guinea, these islands have attracted the attention of naturalists and scientists Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

637

................. 16157$

CH41

02-08-07 10:43:16

PS

PAGE 637

638 / gerald r . a l l e n

ever since they were first visited by European explorers. Waigeo Island, in particular, was the focus of early French visits by several vessels including L’Uranie, La Coquille (1823), and L’Astrolabe (1826). Consequently, approximately 70 fish species were recorded, and Waigeo is an important type locality for a variety of fishes described mainly by Quoy and Gaimard (1824, 1834), Lesson (1828, 1830–1831), and Cuvier and Valenciennes (1828–1849). Fishes that were originally described from Waigeo by early French researchers include such well-known species as the Black-tipped Shark (Carcharhinus melanopterus), Bluefin Trevally (Caranx melampygus), Bigeye Trevally (Caranx sexfasciatus), Semicircular Angelfish (Pomacanthus semicirculatus), and Sergeant Major (Abudefduf vaigiensis). Following the early French explorations, most ichthyological activity in Papua was conducted by Dutch researchers. The famous surgeon-naturalist Pieter Bleeker periodically received specimens from government agents, and in 1868 he published a collection of Waigeo fishes that included 88 species. He added a further 12 species in subsequent papers. Albert Gu¨nther, the Curator of Fishes at the British Museum, recorded 28 species from the island of Misool during the cruise of the Curac¸ao in 1865 (Gu¨nther 1873). The Dutch ichthyologists Weber and de Beaufort were keenly interested in New Guinean freshwater and marine fishes, and contributed to our knowledge of Raja Ampat fishes during the first half of the past century. The work of de Beaufort (1913) was the most extensive effort on Raja Ampat fishes until recent times, and includes accounts of 117 species based on 748 specimens. These were obtained by de Beaufort during a visit to the East Indies in 1909–1910, and were mainly collected at Waigeo in the vicinity of Saonek Island and Mayalibit Bay. Weber and de Beaufort, with various coauthors including Koumans, Chapman, and Briggs, included an additional 67 records from Waigeo and Misool in The Fishes of the Indo-Australian Archipelago (E.J. Brill, Leiden; 11 volumes published between 1921 and 1962). The Denison-Crockett South Pacific Expedition made small collections at Batanta and Salawati totaling 29 species that were reported by Fowler (1939). The only other fish collection of note was that by Collette (1977), who reported 37 species from mangrove habitat on Misool and Batanta. Most recently, the author made the first comprehensive underwater observations of Raja Ampat fishes during four visits between 1998 and 2002 (Allen 2002, 2003).

origin and composition Papua belongs to the overall Indo-West Pacific faunal community. Its reef fishes are very similar to those inhabiting other areas within this vast region, stretching eastward from East Africa and the Red Sea to the islands of Micronesia and Polynesia. Although most families and many genera and species are consistently present across the region, the species composition varies greatly according to locality. The most abundant families on Papuan reefs, in terms of number of species, are gobies (Gobiidae), wrasses (Labridae), damselfishes (Pomacentridae), cardinalfishes (Apogonidae), groupers (Serranidae), butterflyfishes (Chaetodontidae), surgeonfishes (Acanthuridae), blennies (Blenniidae), parrotfishes (Scaridae), snap-

................. 16157$

CH41

02-08-07 10:43:17

PS

PAGE 638

Fishes of Papua / 639

pers (Lutjanidae), and moray eels (Muraenidae). These 11 families collectively account for about 60–65% of the total reef fauna at a typical location. Papua is part of the Indo-Australian region, the richest faunal province on the globe in terms of marine biodiversity. The nucleus of this region, the Coral Triangle, is composed of Indonesia, Sabah, Philippines, and Papua New Guinea (see Chapter 5.2). Species richness generally declines with increased distance from the Triangle, although the rate of attenuation is generally less in a westerly direction. The damselfish family Pomacentridae is typical in this regard. Indonesia has the world’s highest total of damselfish with 138 species, with the following numbers recorded (Allen 1991) for other areas: Papua New Guinea (109), northern Australia (95), west Thailand (60), Fiji Islands (60), Maldives (43), Red Sea (34), Society Islands (30), and Hawaiian Islands (15). The damselfishes also provide evidence that the Raja Ampat Islands of Papua are very close to the center of marine diversity. Its total of 114 species is the highest recorded for any similar-sized area in the world. Indeed, only Indonesia can match this number at the national level. The Raja Ampat Archipelago is Papua’s richest area for coral reef fishes. The author (Allen, 2001, unpublished data) recorded 1,074 species and predicted at least 1,150 species, utilizing a regression formula based on six indicator families (Chaetodontidae, Pomacanthidae, Pomacentridae, Labridae, Scaridae, and Acanthuridae). The majority of Raja Ampat species (about 60%) have wide-ranging distributions in the Indo-Pacific region. The large number of widely distributed species is not surprising considering that nearly all coral reef fishes have a pelagic (open water) larval stage of variable duration. A further 17% are widely distributed in the tropical west Pacific. Twenty percent have a more restricted regional distribution that is confined to the Indo-Australian archipelago. This latter category includes about 25 species that are either confined to Indonesia or the AustraliaNew Guinea region. These are mainly species that seem to lack efficient dispersal capabilities and are therefore unable to exploit oceanic habitats. Underwater observations by the author throughout Indonesia over the past 30 years indicate that 100–330 species can be recorded during a typical scuba dive of one to two hours duration. This total includes species that are readily observed during daylight hours by a trained observer. There is an additional group of cryptic fishes that is not normally observed in the open, including many eels, nocturnal fishes, and small crevice-dwellers. This unseen component may account for up to 15–20% of the species at a particular dive site. A total of 200 or more species is generally considered to be the benchmark for an excellent fish count at any given site. The Raja Ampat Islands in particular exhibit remarkable site diversity. The 200 species benchmark was obtained at 52% of Raja Ampat sites, more than double the figure for its nearest rival, the Togian and Banggai Islands of Sulawesi (Allen 2001, unpublished). Randall (1998) proposed several key factors for the proliferation of coral reef species in the East Indies region (i.e., Indonesia and surrounding areas). He concluded that one of the most important factors concerns the general physiography of the region in combination with past geological and climatic events. Indonesia’s

................. 16157$

CH41

02-08-07 10:43:19

PS

PAGE 639

640 / gerald r . a l l e n

huge contiguous area, replete with island ‘‘stepping stones,’’ has formed a buffer that discourages local extinctions. The geological history of the region is extremely complex. Indonesia is composed of at least three disparate elements that have merged due to tectonic shifts of continental and oceanic masses (Hall 1996, 1998). The western part of the archipelago (i.e., Java, Sumatra, and southern Borneo) is a remnant of the Asian plate. Much of southern and eastern Indonesia (Nusa Tenggara Islands, Moluccas, and westernmost Papua) originated from the Australian plate. Other portions (e.g., northern Borneo, eastern Sulawesi, Halmahera, and northern Papua) were uplifted from oceanic depths as a result of plate collisions. No doubt each of these separate origins made significant contributions to the present reef fauna. Climate has also played a significant role in the proliferation of reef species in the region. The East Indies region has a long history of relatively stable, warm sea temperatures, which has prevented the mass extinctions that occurred elsewhere. In addition, lower sea levels during past glacial periods apparently formed effective barriers that divided formerly widespread species, thus setting the stage for further evolution. Over the last 700,000 years there have been at least three, and possibly as many as six, glacial periods with concomitant lowering of sea levels which effectively closed much of the connecting sea between the Indian and Pacific Oceans (Allen 1975; Chappell 1981; Potts 1983). Although the actual land barrier (Randall 1998) was incomplete, there were likely other barriers in operation, such as low salinity and high turbidity from river discharge, that discouraged marine dispersal. This barrier effect was probably instrumental in the evolution of numerous geminate or ‘‘twin’’ species pairs that originated from eastern and western isolates of a formerly widespread species. Randall (1998) gave examples of 52 probable ‘‘twin’’ species pairs, a convincing argument for this phenomenon. Most of these ‘‘twin’’ species now exhibit widely sympatric distributions, although a few are narrowly sympatric, mainly in the Java-Bali region. A final factor in Randall’s assessment involves life history modes. The East Indies region is populated by numerous reef fishes that have relatively short larval periods and are thus unable to cross deep-water oceanic barriers. For example, the very rich Indonesian pomacentrid fauna includes numerous species that have larval periods ranging between 7 and 18 days (Thresher et al. 1989). It is likely that many species evolved in peripheral regions and were subsequently transported to the East Indies via ocean currents (Ladd 1960; Woodland 1983; Donaldson 1986; Jokiel and Marinelli 1992), thus further enriching the Indonesian reef fish community.

ecology Coral reef fishes are finely synchronized with their environment. Each species exhibits a precise habitat preference dictated by a combination of factors, including the availability of food and shelter, and various physical parameters such as depth, water clarity, currents, and wave action. The huge number of species found

................. 16157$

CH41

02-08-07 10:43:20

PS

PAGE 640

Fishes of Papua / 641

on coral reefs is a direct reflection of the high number of habitat types afforded by this environment. Coral reef fishes generally demonstrate a higher degree of habitat partitioning than do fishes from cooler seas. A good example of the fine scale on which this principle operates is the Urchin Clingfish (Diademichthys lineatus). It is usually found among the spines of Diadema sea urchins or in nearby branching corals, and feeds primarily on the tube feet of its host urchin or on coral-burrowing mollusks. The coral reef offers numerous examples of fishes that have equally narrow habitat and feeding requirements. Water depth is also an important partitioning factor, and there are numerous examples of coral reef fishes that have well defined depth ranges. There are three major depth categories: shallow (0–4 m), intermediate (5–19 m), and deep (⬎20 m). However, the upper and lower limits of these zones can vary at a particular site, depending on the degree of shelter and general sea conditions. The shallow reef environment is typified by wave action, which in highly protected areas such as coastal bays or lagoons may exert its effect to only a few centimeters depth. In contrast, in exposed outer reef structures the effect of surface waves may extend below 10 m. The intermediate zone harbors the greatest abundance of fishes and live corals. Wave action is minimal, although currents are often strong, and sunlight is optimal for reef-building corals. The deep outer reef slope is characterized by reduced light levels and fewer corals and fishes. Although species numbers are reduced in the deep zone, the species that occur in this habitat are among the most interesting of reef fishes. Many of the new species that were discovered on coral reefs over the past four decades were collected on deep reefs by scuba-diving scientists. This trend continues to this day. Papuan reef environments can be broadly classified in two major categories: sheltered inshore reefs or lagoons, and outer reefs. Under optimal conditions both of these habitats can support extensive beds of nearly 100% coral cover. Inshore or coastal reefs are frequently influenced by freshwater runoff and resultant siltation. Underwater visibility on these reefs is often greatly reduced, particularly during the wet season when rivers are flowing at their maximum. Coastal reefs and lagoons are further characterized by extensive sand or silt bottom areas that may support seagrass beds. In most coastal reef or lagoon situations the maximum depth seldom exceeds 25 m, and due to heavy siltation coral growth is generally sparse below 15 m depth. Outer reefs often have a classical reef structure consisting of a broad shallow reef flat, a raised algal ridge, a reef front zone of surge channels, and a steep outer slope. In a few areas of Papua the bottom plunges steeply from the rocky shore. The cleanest water is found on outer reef slopes and underwater visibility may occasionally exceed 30 m. Coral growth is most abundant between about 5–15 m depth, although in some areas significant growth may extend well below this limit. In shallower waters corals are inhibited by the relentless surge, and in deeper water by the much reduced penetration of sunlight. Although most reef-building corals do not thrive below 30–40 m, a number of reef fishes may penetrate well below

................. 16157$

CH41

02-08-07 10:43:21

PS

PAGE 641

642 / gerald r . a l l e n

these depths. Observations made in research submarines in Hawaii and the Marshall Islands indicate that reef species, including some damselfishes, butterflyfishes, and squirrelfishes, may occur at depths approaching 200 m.

biology The majority of reef fishes are egg layers that employ external fertilization. A few species bear live young that can fend for themselves at birth. Included in this category are sharks, rays, and cusk eels. Generally, two patterns of oviparous or egg-laying reproduction are evident in most reef species. Females of many fishes, including the highly visible wrasses and parrotfishes, scatter relatively large numbers of small, positively buoyant eggs into open water where they are rapidly fertilized by the male. The spawning event is typically preceded by nuptial chasing, temporary color changes, and courtship display in which the fins are erected. This behavior is generally concentrated into a short period, often at sunset or shortly afterwards. This pattern is seen in diverse groups such as lizardfishes, angelfishes, wrasses, parrotfishes, and boxfishes. Typically either pair or group spawning occurs in which the participants make a rapid dash towards the surface, releasing their gonadal products at the apex of the ascent. The fertilized eggs float near the surface and are dispersed by waves, winds, and currents. Hatching occurs within a few days and the young larvae are similarly at the mercy of the elements. Recent studies of the daily growth rings found on the ear bones (otoliths) of reef fishes indicate that the larval stage generally persists for about 1–8 weeks, depending on the species. The extended larval period no doubt accounts for the wide dispersal of many reef species. For example, many fishes that occur on Papuan reefs have geographic ranges that extend from east Africa to Polynesia. The other major reproductive pattern consists of species that lay their eggs on the bottom, frequently in rocky crevices, empty shells, sandy depressions, or on the surface of invertebrates such as sponges, corals, or gorgonians. Among the best known fishes in this category are the damselfishes, blennies, gobies, and triggerfishes. These fishes often prepare the surface prior to egg deposition by cleaning away detritus and algal growth. Bottom spawners also exhibit elaborate courtship rituals that involve aggressive chasing and displaying. This behavior has probably been best studied in the damselfishes. In addition, one or both parents may exhibit a certain degree of nest-guarding behavior in which the eggs are kept free of debris and guarded from potential egg feeders such as wrasses and butterflyfishes. A very specialized mode of parental care is seen in cardinalfishes in which the male broods the egg mass in its mouth. Similarly, male pipefishes and seahorses brood their eggs on a highly vascularized region of the belly or underside of the tail. As a rule, these eggs are less numerous, larger, have a longer incubation period, and hatch at a more advanced developmental stage, than the eggs and larvae of pelagic spawning fishes. Hatching may require up to one week (in anemonefishes, for example) and the larvae then lead a pelagic existence for up to several weeks before settling on the bottom in a suitable reef habitat.

................. 16157$

CH41

02-08-07 10:43:21

PS

PAGE 642

Fishes of Papua / 643

There is scant information on the longevity of most reef fishes. Perhaps one of the longest life spans is that of the Lemon Shark, which may reach 50 years or more. Most of the larger sharks probably live at least to an age of 20–30 years. Larger reef fishes such as groupers (Serranidae), snappers (Lutjanidae), and emperors (Lethrinidae) tend to live longer than smaller species. Otolith (small bones in the inner ear) aging techniques indicate that large groupers may live at least 25 years and some snappers approximately 20 years. Most of our knowledge of smaller reef fishes comes from aquarium studies. The values obtained from captive fishes may exceed the natural longevity due to lack of predation and the protective nature of the artificial environment. Batfishes (Platax) are known to survive for 20 years and even small species such as damselfishes and angelfishes may reach an age of ten years or more in captivity.

conservation Papua contains a wealth of marine ecosystems ranging from extensive coastal mangroves to pristine offshore coral reefs. Lying well off the beaten track and supporting a relatively small human population, the area is still in relatively undisturbed condition. However, there are already disturbing trends evident, particularly the increased use of illegal fishing methods as well as a general depletion of sharks and large reef fishes. Itinerant fishing vessels from other parts of Indonesia are becoming an all too common sight. Sharks and large edible reef fishes are scarce throughout Papua, especially in the Raja Ampat Islands. This situation is typical for Indonesia and is a direct result of overfishing, compounded by the use of illegal fishing methods, especially explosives and potassium cyanide. The general scarcity of large groupers (Cromileptes, Epinephelus, and Plectropomus) and Napoleon Wrasse (Cheilinus undulatus) in particular, is a result of the pervasive influence of the Hong Kong–based live restaurant fish trade. Visiting fishermen seldom respect the fishing rights of local people, which invariably leads to conflict. The situation is particularly bad in relatively remote areas that lack effective police enforcement. Local villagers are literally chased away from their traditional fishing grounds by these armed and dangerous itinerants. The intruders harvest their quarry with impunity, accumulating large numbers of live fishes in floating holding pens. Community programs need to be designed that allow local villages to regain control of their traditional reef areas, with stiff fines and imprisonment facing anyone who uses illegal fishing methods. Also, the practice of outsiders fishing for live groupers and Napoleon Wrasse on traditional reefs needs to halted or at least controlled for the benefit of local communities. Despite this illegal activity, there appears to be less impact from illegal fishing methods in the Raja Ampat Islands than in other parts of Indonesia. A rapid assessment (RAP) by Conservation International in 2001 revealed that most of the 44 dive sites surveyed were in good condition. However, damage from explosives was noted at seven locations. Observations of Napoleon Wrasse, a conspicuous

................. 16157$

CH41

02-08-07 10:43:22

PS

PAGE 643

644 / gerald r . a l l e n

indicator of fishing pressure, showed that it was indeed heavily exploited, a typical situation throughout Indonesia. The species is far more common at Milne Bay Province, Papua New Guinea, where illegal fishing methods are seldom used. Although a number of marine sanctuaries have already been gazetted, they are merely ‘‘paper parks,’’ without any real enforcement. There is an urgent need to establish an effective network of marine reserves in Papua before it is too late. Without question the absence of fishing pressure, or at least much reduced fishing activity, results in robust populations of coral trout and other large edible fishes such as snappers (Lutjanidae) and tuskfishes (Choerodon: Labridae). Two of the best examples are the Montebello and Abrolhos islands in Western Australia where fishing activity is restricted to occasional visits by recreational anglers. Consequently, it is not unusual to see more than 50 coral trout on a single dive. Uncontrolled clear-cut logging also represents a threat to coral reefs, which can be damaged or destroyed by resultant erosion and sediment deposition. Illegal logging in gazetted wildlife reserves continues to be a problem, particularly on Waigeo and Batanta islands. Although still a minor activity, dive tourism has greatly increased in the past few years. There are a number of live-aboard boats operating in the Raja Ampat Islands, and dedicated dive resorts are starting to appear. Reefs need to be protected from anchor damage. It would also be desirable to institute a fee system that benefits local villages situated near popular dive sites. There is convincing evidence that the dive-tour industry and effective conservation of reef ecosystems can comfortably co-exist. A good example is the thriving dive trade and effective resource management at the Bunaken Marine Reserve, Sulawesi.

endemism and hotspots ‘‘Endemism’’ is a term commonly used by biogeographers (scientists who study the geographic distribution of plants and animals) referring to an organism that is restricted to (i.e., endemic to) a particular area. The area may be extremely small, such as a single lake, or extensive, such as the island of New Guinea or the entire Pacific Ocean. Plants and animals that are endemic to exceptionally limited areas are particularly vulnerable to various threats, particularly when close to human population centers. For this reason they are of special interest to conservationists. Regional and local endemism are commonly used to define conservation ‘‘hotspots’’: areas that have a disproportionate number of endemic species. Considering the broad dispersal capabilities via the pelagic larval stage of most reef fishes, it is not surprising that relatively few fish species are endemic to the Papuan region. Allen and Adrim (2003) listed the following five species as Papuan endemics: the Hemiscyllid Shark (Hemiscyllium freycineti; Quoy and Gaimard 1824); two cardinalfishes (Apogonidae) including Apogon leptofasciatus and A. oxygrammus (Allen 2001a); a damselfish (Pomacentridae: Chrysiptera pricei; Allen and Adrim 1992), and the goby (Gobiidae: Eviota raja; Allen 2001b). With the exception of the shark, these species belong to families that exhibit parental care and presumably have brief larval stages. All except Chrysipter pricei from Yapen

................. 16157$

CH41

02-08-07 10:43:22

PS

PAGE 644

Fishes of Papua / 645

Island are currently known only from the Raja Ampat Islands, which appears to be a hotspot for regional endemism within Indonesia (Allen and Adrim 2003).

Freshwater Fishes Although New Guinea has one of the richest marine fish faunas in the world, only about 400 freshwater fishes have been recorded (Allen 1991, unpublished data). By comparison, the Kapuas River system of Kalimantan has nearly 300 species (Roberts 1989) and it is conservatively estimated that there are 2,000 species in the Amazon Basin. In spite of the relatively low numbers, New Guinea’s fishes are highly unusual and deserve special attention.

historical background The first freshwater species from Papua, an ariid catfish (Arius leptaspis), was described in 1862 by the Dutch ichthyologist Pieter Bleeker. This was the beginning of a very significant Dutch contribution, culminating in a series of expeditions between 1903 and 1920. The majority of these collections were summarized by Weber (1913), who was the first author to treat the freshwater fauna comprehensively. Additional species were described in the monumental, 11 volume work Fishes of the Indo-Australian Archipelago by Weber and de Beaufort (1911–1962). Most of the specimens that were collected during the busiest exploration period (1903–1920) are currently deposited at the Zoological Museum of Amsterdam. The early visitors were greatly hampered by limited access to the interior and endemic diseases, particularly malaria. Consequently, their collections were mainly obtained from sites near the coast or inland along the larger, navigable rivers. These inhibiting factors persisted until relatively recent times. Although vast tracts of nearly impenetrable wilderness still remain, a reasonably good network of commercial and private air routes has developed. The increased use of helicopters associated with mining and logging ventures has also facilitated biological exploration in remote areas. There was very little collecting activity in Papua between 1921 and 1982, the notable exception being the Dutch expedition of 1954–1955 (Boeseman 1963). Fishes were collected on the latter expedition from widely scattered areas including Lake Sentani and the nearby Grime and Digul rivers, Paniai Lakes, Lake Yamur, and the central Vogelkop Peninsula in the vicinity of the Ayamaru Lakes and nearby Lake Aitinyo. These collections were eventually reported by Allen and Boeseman (1982) and the specimens were lodged at the National Museum of Natural History (Leiden) and the Western Australian Museum (Perth). Between 1983 and 2002, the author made nine visits to Papua, resulting in valuable collections from the following localities: Jayapura vicinity (Lake Sentani, Grime River, Bewani River, and Cyclops Coast), Mamberamo River, Danau Bira, Nabire, Timika region, Paniai Lake, Wapoga River, Yapen Island, Biak, Lake Yamur, Etna Bay, Triton Lakes, Bomberai Peninsula, Vogelkop Peninsula (Ayamaru Lakes, Lake Aitinyo, Bintuni region, Teminabuan, Senopi, Manokwari, and

................. 16157$

CH41

02-08-07 10:43:22

PS

PAGE 645

646 / gerald r . a l l e n

Sorong), and the Raja Ampat Islands (Waigeo, Batanta, Salawati, Misool). These collections are deposited at the Museum Zoologense Bogoriense (Bogor, Indonesia) and the Western Australian Museum.

origin, composition, and distribution New Guinea and Australia have been connected by dry land in the region that is now the Torres Strait and the Arafura Sea for a significant part of their geological past (Loeffler 1977). Nearly all families, most genera, and about 34 species are shared by these two areas. Two closely related families, rainbowfishes (Melanotaeniidae), and blue-eyes (Pseudomugilidae) are endemic to this combined region. The fish fauna of New Guinea and Australia is very different from that of other continental tropical regions such as Southeast Asia, Africa, and South America. The latter areas tend to be dominated by cichlids and primary division ostariophysan fishes such as carps, barbs, loaches, characins, and catfishes. Cichlids were recently introduced to the New Guinea region and ostariophysans are represented only by plotosid and ariid catfishes, but in contrast to the primary division species that evolved entirely in fresh water, they are considered to be secondary division fishes of marine derivation. In fact, all the fishes of the region, with the exception of the Australian lungfish (Neoceratodus), bony tongues (Scleropages), and possibly galaxiids (a southern Australian family), were derived from marine ancestors. Most of Indonesia to the west of Papua, with the possible exception of Timor Island and western Sulawesi, is part of the Asian continental plate, which until about 20 million years ago was well separated from the New Guinea-Australian plate. Even today a barrier of deep oceanic water remains that effectively prevents the dispersal of fresh water fishes. The Dutch ichthyologist Max Weber delineated this zoological boundary in 1919. Accordingly, it is often referred to as ‘‘Weber’s Line,’’ lying just to the west of the Raja Ampat Islands (westernmost Papua) and well to the east of the more famous Wallace’s Line. There is little information about the evolutionary details of the present freshwater fish fauna. No fossils have been found so far that would help to elucidate this fauna’s origins. Certainly, for much of its history, the southern platform of New Guinea must have been the haunt of ceratodid lungfishes. As many as seven species once inhabited northern Australia and the modern Neoceratodus of eastern Australia has remained virtually unchanged for over 100 million years (Kemp and Molnar 1981). However, lungfishes did not survive in New Guinea. Perhaps the most ancient fish still found there is the Saratoga (Scleropages jardinii), a member of the primitive bony tongue family Osteoglossidae. The family apparently evolved in fresh waters of the Southern Hemisphere and although it is considerably older, fossil records date back to 38–55 million years ago. The family is also represented today in southeastern Asia, South America, and Africa. Marine families such as ariid and plotosid catfishes, atherinids, gudgeons, and gobies provided the evolutionary nucleus of the present fauna. They probably reached the northward drifting continent (Australia and the attached southern

................. 16157$

CH41

02-08-07 10:43:23

PS

PAGE 646

Fishes of Papua / 647

New Guinea) after it reached its present position adjacent to the Indonesian Archipelago. Some of the original colonizers no doubt arrived as drifting pelagic eggs or larvae, a mode of dispersal that is common among modern inshore fishes. It is not difficult to imagine that certain marine fishes, particularly euryhaline estuarine dwellers, were able to penetrate inland. Once established in this extensive habitat, the stage was set for further evolution, which occurred when populations became fragmented due to the formation of natural barriers resulting from catastrophic geological events. The most prominent groups of freshwater fishes in Papua include ariid and plotosid catfishes (combined total of 32 species), rainbowfishes and allied atherinoid fishes (53 species), and gobioid fishes (90 species). These major groupings represent 56% of the total species. The distribution of purely freshwater species is strongly correlated with the island’s geology. There are two major faunal provinces, north and south of the lofty Central Dividing Range. The southern province is by far the most speciose, reflecting its long, relatively stable history. As would be expected due to the recent land connection, this part of Papua exhibits the strongest affinity with Australia. About 30 species are common to both regions. Also, the presence of several closely related rainbowfishes and gudgeons, living on either side of the Arafura Sea, indicates that their speciation was a relatively recent event, perhaps having occurred within the past 6,000–8,000 years. Northern New Guinea was formed in relatively recent geological times, within the past 10 million years, as a result of upthrusting in the zone of collision between the Australian and Pacific plates (Loeffler 1977). It is evident from their relationships that components of the northern fauna were derived from the older southern fauna. It is likely that the northern forms evolved from ancestral species that became isolated as a result of the uplifting of the central mountains, which occurred 5–6 million years ago. No indigenous fishes have been found above an elevation of 1,800 m. Highland fishes in general are restricted to broad, low elevation valleys, the lower moderategradient sections of headwater streams, and lake habitats. Native fishes are conspicuously absent in many upland areas where suitable habitat exists, such as the extensive Baliem Valley. Their absence probably stems from the violent geological past of the central mountains. There was concurrent volcanism and glaciation as recently as 300,000 years ago. Moreover, widespread glaciation and greatly lowered temperatures persisted until 15,000 years ago. The fishes that have been most successful in penetrating the mountains include plotosid catfishes, gudgeons, and gobies. Although the island is divisible into major northern and southern faunal regions, at least four distinct provinces are evident (Allen 1991). Their recognition is based on the presence of a substantial number of regional endemics. Three of these provinces are partially or entirely within Papua and are described briefly in the following sections.

................. 16157$

CH41

02-08-07 10:43:23

PS

PAGE 647

648 / gerald r . a l l e n

Great Southern or Trans-Fly Province This region includes most of the southern half of central New Guinea. It stretches eastward from the narrow isthmus at the base of the Vogelkop Peninsula in Papua to the Purari River of Papua New Guinea, or a linear distance of about 1,200 km. The area contains the most extensive alluvial lowland plains on the island, largely composed of deposits from the Digul and Fly river systems. Leading families for endemism include Ariidae (10 species), Eleotridae (6), and Plostosidae (5). In addition to 35 endemic species, the region is characterized by 34 species that are shared with northern Australia. There are also four endemic genera of ariid catfishes (Cochlefelis, Doiichthys, Nedystoma, and Tetranesodon). The Papuan portion of this province contains about 70 species of purely freshwater fishes. The Aru Islands, lying approximately 120 km south of Papua, is provisionally included in this province. The island group was part of the former land connection between Australia and Papua, hence is of special interest to biologists. Freshwater fishes are poorly known; no collections have been made since Dutch expeditions visited during the early 1900s. The fishes are currently considered to be the same species that occur on the mainland, but a modern survey is urgently required to re-evaluate their status. It is possible that a high level of endemism exists.

Great Northern or Mamberamo-Sepik Province This is the largest province and includes most of the northern half of central New Guinea. It contains several of the island’s largest rivers including the Mamberamo, Sepik, Ramu, and Markham. Associated with these river basins are extensive lowland alluvial plains with meandering and braided channels that undergo periodic flooding. The major rivers also have vast foothill and low mountain tributaries. Barriers between the four major systems are relatively insignificant and appear to be very recent. Consequently, there is considerable faunal uniformity within the region. The total fauna consists of about 145 species, of which 57 are pure freshwater forms (including 7 introductions), and the remainder have marine life-cycle stages. There are many endemic species of ariids (5), melanotaeniids (18), gobies (6), and gudgeons (8), as well as two endemic melanotaeniid genera (Chilatherina and Glossolepis).

Vogelkop Peninsula and Western Islands Province The Vogelkop comprises the extreme western end of the mainland and is essentially isolated by a narrow mountainous isthmus. It includes the main Vogelkop Peninsula, the smaller Bomberai Peninsula (lying north and south respectively of the intervening Bintuni Bay), and also the mountainous region west of Etna Bay. The region consists mainly of lowland rainforest with large tracts of heavily corrugated karst topography. There are also relatively high mountains along the northern coast of the Vogelkop. The province also includes the Raja Ampat Islands, lying offshore immediately to the west. The four largest islands, Waigeo, Batanta, Salawati, and Misool, were connected to the mainland as recently as 10,000 years ago when sea levels were 100 m lower. About 90 species are currently known from

................. 16157$

CH41

02-08-07 10:43:24

PS

PAGE 648

Fishes of Papua / 649

the province, including 30 pure freshwater forms and five introduced species. The largest groups of endemics include rainbowfishes (15 species) and gudgeons (6). The narrow isthmus connecting the broad central portion of the island (eastern Papua) with the Bomberai and Vogelkop peninsulas is characterized by a high number of endemic species and at least merits subprovincial recognition. Especially interesting is the rugged karst area between Arguni and Etna bays. Although relatively few species are documented, most are endemic to this small region. A high level of endemism is particularly evident in the Triton Lakes, Mbutu Basin, and Lake Kaifayama, which lie immediately west of Etna Bay. There is also an endemic melanotaeniid genus (Pelangia) known only from the Mbutu Basin.

ecology Freshwater habitats can be broadly classified as either ‘‘lotic’’ (flowing) or ‘‘lentic’’ (still). Lotic ecosystems include perennial and intermittent streams, flowing springs or seepages, artificial ditches, and flumes. Lentic systems contain two types of standing waters which ecologists refer to as palustrine (bogs, marshes, and swamps) or lacustrine (lakes, ponds, and reservoirs). Both of these major habitat types are well represented in Papua, and provide a home for fishes. Within each of the main ecosystems there is considerable variation, depending on such factors as water clarity, flow rate, temperature, substrate type, availability of shelter, exposure to sunlight, and so on. Freshwater sites throughout Papua are populated by relatively few species, usually ranging between 2–14 species per site. Fishes inhabiting a particular stretch of stream or section of pond are finely tuned to their surroundings. Their continued survival is dependent on the availability of adequate food resources and shelter, as well as the correct conditions for spawning. Food supplies fluctuate greatly according to the seasons, and the life cycles of resident fishes are often geared to make maximum use of these resources for the production of gametes and the growth of their young. Many Papuan fishes, including plotosid catfishes, glassfishes, hardyheads, and rainbowfishes exhibit spawning peaks during the rainy season. This strategy offers young fish an improved chance of survival as flood conditions provide shelter in the form of inundated streamside vegetation. In addition, the increased turbidity offers further protection from bird and fish predators. The aquatic food chain consists of complex interactions of producer and consumer organisms. Fishes play a major role, utilizing a wide range of food resources that include organic detritus, microscopic algae (diatoms), encrusting or filamentous algae, bottom-rooted or floating plants (macrophytes), planktonic animals (especially crustaceans), aquatic insects and their larvae, and a range of terrestrial insects and other small animals that live near the water’s edge. Fishes, in turn, are keenly sought by a range of predators, of which birds and humans are the most prominent. Birds in particular feed on huge numbers of fishes, especially during drought periods.

................. 16157$

CH41

02-08-07 10:43:24

PS

PAGE 649

650 / gerald r . a l l e n

biology Several major spawning modes are exhibited by Papuan freshwater fishes. By far the most common type is external fertilization in which eggs and sperm are simultaneously released, or nearly so, by pairs or aggregations. The majority of species are demersal (bottom) spawners. Eggs are negatively buoyant and sink to the bottom or adhere to rocks or vegetation. The bottom, whether it consists of rocks, gravel, plants, or debris, hides the eggs from potential predators (especially surface and mid-water feeding fishes), and protects them from overheating or heavy buffeting which could occur if they floated at the surface. This demersal mode contrasts sharply with the floating pelagic eggs and larvae of most marine fishes. Many fishes simply scatter their eggs randomly over the bottom, but others such as the eel-tailed catfishes (Plotosidae), gobies (Gobiidae), and gudgeons (Eleotridae) construct nests and exhibit various degrees of parental care. Frequently one or both parents drive away intruding fishes, and methodically fan the eggs with their pectoral fins to remove any silt or debris. This behavior usually only lasts until hatching, at which time the tiny fry must fend for themselves. However, the introduced cichlid (Sarotherodon mossambica) may guard their brood for several weeks. Freshwater eels (Anguillidae) and the Barramundi (Lates calcarifer) exhibit what is known as a catadramous life cycles. They spend much of the adult stage in freshwater, but undergo a spawning migration to the sea. Perhaps the most unusual reproductive habits are displayed by a small number of fishes that brood their eggs (and sometimes also the young) in either the mouth or some other part of the body. Mouth brooding species in Papuan fresh waters include the Saratoga (Scleropages jardinii), fork-tailed catfishes (Ariidae), cardinalfishes (Apogonidae), and the introduced cichlid (Sarotherodon mossambica). Usually the male parent takes the egg mass into his mouth immediately after spawning. However, this task is assumed by the female in the Saratoga and also the cichlid. The incubation period is variable, depending on species, but generally ranges from a few days up to three weeks. Parental duties terminate at hatching for catfishes and cardinalfishes, but recently hatched saratogas and cichlids may be kept in the mouth or close by the parent for several weeks. Pipefishes (Syngnathidae) are pouch brooders. The female deposits her eggs on the underside of the male’s tail on a special vascularized area that is protected by a pair of skin flaps. The eggs are carried until hatching, at which time the young are ready to care for themselves. Similarly, the male Nurseryfish (Kurtus gulliveri) carries a cluster of eggs on a peculiar hook-like process protruding from his forehead.

conservation There remains an urgent need to conduct floral and faunal surveys throughout much of Papua to identify areas of special conservation value. Many freshwater fishes, particularly the rainbowfishes (Melanotaeniidae), appear to be restricted to an isolated lake or small part of a single river system. These species are highly vulnerable to environmental disturbances such as clearcut logging, mining, and

................. 16157$

CH41

02-08-07 10:43:24

PS

PAGE 650

Fishes of Papua / 651

dam construction. Until just a few years ago these threats were minimal, but the situation is rapidly changing. Both Papua and Papua New Guinea have government conservation agencies staffed by skilled personnel, but they are frequently frustrated when economic concerns are given a higher priority than protection of native wildlife. The opening of a mining or logging operation usually injects much needed capital into the local economy. Consequently, key conservation issues are often completely ignored. Local people need to be informed of the uniqueness of their plants and animals and the need to protect them. Unfortunately, this is usually difficult to implement. Of all the special freshwater habitats in Papua, Lake Sentani is probably the most highly threatened, mainly due to its proximity to a major population center (Jayapura/Sentani) and resultant overfishing and pollution. This lake also suffers the fate of many other places in Papua that support immigrant populations: the introduction of undesirable exotic fishes. Species such as Carp (Cyprinus carpio), Walking Catfish (Clarias batrachus), Tilapia (Oreochromis mossambica), Climbing Perch (Anabas testudineus), and Snakehead (Channa striata) are particularly damaging to the native fauna. Unfortunately, the seemingly remote Mamberamo River system is also plagued by undesirable exotics. Nearly 18% of the fauna consists of exotic species that were mostly introduced in the 1970s and 1980s and are now well established. No other major river system in New Guinea has such a high percentage of introduced fishes. Virtually all of the Mamberamo introductions have been shown to negatively impact native fish populations wherever they have been released. Exotics compete for living space and available food resources, or feed directly on native species. Tilapia (S. mossambicus) and Carp (C. carpio) are also notorious for adversely affecting the environment by creating turbid conditions in formerly clear lakes, and badly crowding native fishes due to prolific breeding. It is too late to save the Mamberamo system because once established the introductions are virtually impossible to eradicate. Every effort should be made to prevent further introductions in other systems of Papua, but there does not appear to be any practical way to control this activity, nor has any concern been expressed by government agencies.

endemism and hotspots The Papuan freshwater fish fauna is strongly endemic. Excluding the 128 species that have marine life-cycle stages, the only nonendemics are the 34 species shared with Australia. Therefore, 84% of the freshwater dependent species are found exclusively on the island of New Guinea. The geographic ranges of many of these species includes both Papua and Papua New Guinea, but 64 species are restricted to Papua. Rainbowfishes of the family Melanotaeniidae are the leading family of Papuan endemics, with 30 species represented. Analysis of regional endemism within Papua reveals that there are six major centers, of which the Great Northern (21 species) and Bird’s Neck Isthmus (17 species), are the most important. The Bird’s Neck area has a particularly impres-

................. 16157$

CH41

02-08-07 10:43:25

PS

PAGE 651

652 / gerald r . a l l e n

sive total considering its relatively small area (approximately 15,000 km2) compared to other regions. In addition to these regional areas of endemism there are several highly localized hotspots, of which the Triton Lakes in the Bird’s Neck is unsurpassed, with nine endemic species. Other significant hotspots include Lake Sentani and vicinity (5 endemics), Raja Ampat Islands (5 endemics), Wapoga River system (4 endemics), and the vicinity of Timika (4 endemics).

Acknowledgments The author’s studies of New Guinea fishes were greatly aided by support from the Western Australian Museum, Indonesian Institute of Sciences (LIPI), Conservation International, The Nature Conservancy, University of Cenderawasih, National Geographic Society, and PT Freeport Indonesia.

Literature Cited Allen, G.R. 1975. The Anemonefishes, Their Classification and Biology, 2nd ed. Tropical Fish Hobbyist Publications, Neptune City, New Jersey. Allen, G.R. 1991a. Damselfishes of the World. Aquarium Systems, Mentor, Ohio. Allen, G.R. 1991b. Field Guide to the Freshwater Fishes of New Guinea. Publication No. 9. Christensen Research Institute, Madang, Papua New Guinea. Allen, G.R. 2001a. Two new species of cardinalfishes (Apogonidae) from the Raja Ampat Islands, Indonesia. Aqua, J. Ichthy. Aquat. Biol. 4 (4): 143–149. Allen, G.R. 2001b. Description of two new gobies (Eviota, Gobiidae) from Indonesian seas. Aqua, J. Ichthy. Aquat. Biol. 4 (4): 125–130. Allen, G.R. 2002. Reef fishes of the Raja Ampat Islands, Papua Province, Indonesia. Pp. 46–57 in McKenna, S., G.R. Allen, and S. Suryadi (eds.) A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, D.C. Allen, G.R. 2003. Coral reef fishes of the Raja Ampat Islands. Pp. 42–58 in Donnelly, R., D. Neville, and P.J. Mous (eds.) Report on a rapid ecological assessment of the Raja Ampat Islands, Papua, Eastern Indonesia. The Nature Conservancy–Southeast Asia Center for Marine Protected Areas, Bali, Indonesia. Allen, G.R., and M. Adrim. 1992. A new species of damselfish (Chrysiptera: Pomacentridae) from Irian Jaya, Indonesia. Rec. West. Aust. Mus. 16 (1): 103–108. Allen, G.R., and M. Adrim. 2003. Coral reef fishes of Indonesia. Zool. Stud. 42 (1): 1–72. Allen, G.R., and M. Boeseman. 1982. A collection of fishes from western New Guinea with descriptions of two new species (Gobiidae and Eleotridae). Rec. West. Aust. Mus. 10: 67–103. Bleeker, P. 1868. Notice sur la faune ichthyologique de l’ıˆle de Waigiou. Versl. Akad. Amsterdam (2) II: 295–301. Chappell, J. 1981. Relative and average sea level changes and endo-, epi-, and esogenic processes on the earth. Pp. 411–430 in Sea Level, Ice, and Climatic Change. Int. Assoc. Hydrol. Sci. Publ. 1. Collette, B.B. 1977. Mangrove fishes of New Guinea. Pp. 91–102 in Teas, H.J. (ed.) Tasks for Vegetation Science. W. Junk, The Hague. Cuvier, G., and A. Valenciennes. 1828–1849. Histoire naturelle des poissons. 22 vols. Paris.

................. 16157$

CH41

02-08-07 10:43:25

PS

PAGE 652

Fishes of Papua / 653 de Beaufort, L.F. 1913. Fishes of the eastern part of the Indo-Australian Archipelago with remarks on its zoogeography. Bijd. Neder. Dierk., Amsterdam 19: 95–163. Donaldson, T.J. 1986. Distribution and species richness patterns of Indo-West Pacific Cirrhitidae: support for Woodland’s hypothesis. Pp. 623–628 in Uyeno, T., R. Arai, T. Taniuchi, and K. Matsuura (eds.) Indo-Pacific Fish Biology: Proceedings of the Second International Conference on Indo-Pacific Fishes. Ichthyol. Soc. Japan, Tokyo. Fowler, H.W. 1939. Zoological results of the Denison-Crockett South Pacific Expedition for the Academy of Natural Sciences of Philadelphia 1937–1938. Part III.–Fishes. Proc. Acad. Nat. Sci. Philadelphia 91: 77–96. Gu¨nther, A. 1873. Reptiles and fishes of the South Sea islands. In Brenchley, J.L., Jottings during the cruise of H.M.S. Curac¸ao among the South Sea Islands in 1865. Cruise Curac¸ao: 1–487, Pls. 1–59. Hall, R. 1996. Reconstructing Cenozoic SE Asia. Pp. 153–184 in Hall, R., and D.J. Blundell (eds.) Tectonic Evolution of SE Asia. Geol. Soc. London Spec. Publ. 106. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. Pp. 99–131 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of South-East Asia. Backhuys, Leiden. Jokiel, P., and F.J. Martinelli. 1992. The vortex model of coral reef biogeography. J. Biogeogr. 19: 449–458. Kemp, A., and R.E. Molnar. 1981. Neoceratodus forsteri from the lower Cretaceous of New South Wales. Aus. J. Paleo. 55: 211–217. Ladd, H.S. 1960. Origin of the Pacific island molluscan fauna. Amer. J. Sci. 258A: 137–150. Lesson, R.P. 1828. Description du noveau genre Ichthyophis et des plusieurs espe`ces ine´dites ou peu connues de poissons, recueillis dans le voyage autour du monde de la Corvette ‘‘La Coquille.’’ Mem. Soc. Nat. Hist. Paris 4: 397–412. Lesson, R.P. 1830–31. Poissons. Pp. 66–238 in Duperrey, L. (ed.) Voyage autour du monde, . . . , sur la corvette de sa Majeste´ La Coquille, pendant les anne´es 1822, 1823, 1824 et 1825, Zoologie 2 (part 1). Paris. Loeffler, E. 1977. Geomorphology of Papua New Guinea. Australian National Univ. Press, Canberra. Potts, D.C. 1983. Disequilibrium among Indo-Pacific corals. Bull. Mar. Sci. 33: 619–632. Quoy, J.R.C., and J.P. Gaimard. 1824. Poissons. Pp. 183–401 in de Freycinet, L.M. (ed.) Voyage autour du monde, Enterpris par ordre du Roi exe´cute´ sur les corvettes de S.M. L’Uranie et La Physicienne pendant les anne´es 1817 1818 1819, et 1820. Paris. Quoy, J.R.C., and J.P. Gaimard. 1834. Voyage de de´couvertes de ‘‘L’Astrolabe’’ exe´cute´ par ordre du Roi, pendant les anne´es1826–1829, sous le commandement de M.J. Dumont d’Urville. Poissons III: 647–720. Randall, J.E. 1998. Zoogeography of shore fishes of the Indo-Pacific region. Zool. Stud. 37: 227–268. Roberts, T.R. 1989. The freshwater fishes of western Borneo (Kalimantan Barat, Indonesia). Mem. Cal. Acad. Sci. 14: 1–210. Thresher, R.E., P.L. Colin, and L.J. Bell. 1989. Planktonic duration, distribution, and population structure of western and central Pacific damselfishes (Pomacentridae). Copeia: 420–434. Weber, M. 1913. Susswasserfische aus Niederlandisch Sud- und Nord-Neu Guinea. Nova Guinea (Leiden) 9: 513–613. Weber, M., and L.F. de Beaufort. 1911–1962. The Fishes of the Indo-Australian Archipelago. Vols I–X1. E.J. Brill, Leiden. Woodland, D.J. 1983. Zoogeography of the Siganidae (Pisces): an interpretation of distribution and richness patterns. Bull. Mar. Sci. 33: 713–717.

................. 16157$

CH41

02-08-07 10:43:25

PS

PAGE 653

4.9. Birds of Papua andrew mack and jack dumbacher p p ro x i m at e l y 831 bird species have been recorded from the New Guinea region, including Papua and Papua New Guinea (Beehler et al. 1986; Coates 1985, 1990; Rand and Gilliard 1967; Sibley and Monroe 1990). This represents approximately 8.6% of the extant global avian alpha diversity (Table 4.9.1) and is roughly equivalent to the number of species recorded on the continent of Australia. Approximately 657 species are recorded from Papua (Appendix 8.2). This list is tentative because a full systematic review of the literature, existing collections, and sight records has not yet been completed. Although most species are shared with Papua New Guinea, 67 species recorded from Papua have not been recorded in PNG (Appendix 8.2). Of these 38 are also unknown from surrounding Indonesia, Australia, or Melanesia and can be considered endemic to Papua. Of the nonendemic taxa, 285 occur only in PNG and Melanesia (thus comprising endemics to the New Guinea/Melanesia region), 141 species are fairly widespread globally, 110 are shared with Australia, and the remaining 83 have distributions overlapping with Australia or the remainder of Indonesia. Seventy-six species on the list can be classified as migrants, with at least some of the population moving in and out of Papua, either to Australia or Asia; 21 can be considered vagrants; one species is introduced; and approximately 558 are presumably breeding permanent residents (Appendix 8.2).

A

Table 4.9.1. Distribution of bird species of Papua and neighboring regions Distribution

No. of species

Percent of avifauna

World

9,702

100*

New Guinea Region

831

8.6*

Papua

657

6.8*

Land and freshwater species breeding in Papua

552

5.7*

Endemic to Papua

38

5.8•

Shared with PNG and Melanesia

285

43.4•

Shared with Australia

110

16.7•

83

12.6•

141

21.5•

Shared with eastern Indonesia and Australia Widespread

Note: * indicates percent of the world’s avifauna (bird species), • indicates percent of Papua’s avifauna. Source: Classification follows Sibley and Monroe (1990). Distribution data compiled from various sources, including Beehler et al. 1986; Beehler and Finch 1985; Coates 1985, 1990; Mayr 1941; Rand and Gilliard 1967; Sibley and Monroe 1990.

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

654

................. 16157$

CH42

02-08-07 10:43:32

PS

PAGE 654

Birds of Papua / 655

Several bird families have important radiations in Papua. One of the most notable avian groups is the birds of paradise (Paradisaeidae) with 24 species in Papua, excluding the Cnemophilines and Macgregoria (Cracraft and Feinstein 2000). Birds of paradise have been the focus of more research than any other avian family in New Guinea because of their unusual mating systems, behaviors, and plumage (Frith and Beehler 1998). However, other families are comparably diverse and dominant in the New Guinea avifauna, such as honeyeaters (Meliphagidae) and bowerbirds (Ptilonorhynchidae). Twelve species of berrypeckers are placed in two families, Melanocharitidae and Paramythiidae (Sibley and Monroe 1990). These are the only widely recognized bird families endemic to New Guinea. No berrypeckers have been studied in the field, so our knowledge of this unique radiation is limited to what can be gleaned from museum data and a few published observations. Twenty-three species of kingfisher occur in Papua (four species of Alcedinidae and 19 species of Dacelonidae), more than four times the number of kingfisher species found in North and South America combined. Other notably species-rich families include Columbidae (42 species of about 300 species worldwide), Meliphagidae (55 species of about 178 species worldwide), Megapodiidae (9 of the world’s 19 species), Acanthizidae (20 of the world’s 65 species), and Psittacidae (46 species of about 340 worldwide).

History of Ornithological Exploration Since humans first arrived in Papua, they have studied and used birds. The first humans arrived in New Guinea sometime between 26,000 and 50,000 years ago (Bulmer 1982). Traditional societies of New Guinea have used birds for food, clothing, decoration, rituals, clan totems, and as wealth to be traded both within New Guinea and to outside countries (Bulmer 1982; Majnep and Bulmer 1977). Over thousands of years, native New Guineans have accumulated great stores of taxonomic (Diamond 1966) and natural history information about birds (e.g., Feld 1990; Kocher-Schmid 1991, 1993; Majnep and Bulmer 1977). While most of this anthropological information remains unrecorded, some groups of anthropologists and ethnographers (especially Bulmer, Diamond, and others) have gone to great lengths to record and preserve this information. Much work remains to be done in this field. Birds have been important for New Guinea’s trade and exploration for millennia. Birds and bird of paradise plumes were traded with Asia as long as 5,000 years ago (Frith 1971; Swadling 1995), and birds, as well as other aspects of New Guinea’s unique biota, attracted European naturalists such as Alfred Russel Wallace to spend months or years exploring mainland New Guinea. The birds of New Guinea and the ornithologists who studied them have played pivotal roles in broader arenas of science, exploration, and modern biological thought. A. R. Wallace’s treatise on the Malay Archipelago (Wallace 1869) has become important reading for any student of biogeography. His time in Papua helped him to postulate independently that natural selection is the process that

................. 16157$

CH42

02-08-07 10:43:33

PS

PAGE 655

656 / a n d rew m a c k & ja c k d um b ac her

explains much of evolution (a proposal that greatly affected the entire world when Charles Darwin published The Origin of Species.) And New Guinea’s birds motivated many of the early expeditions into the rugged interior of Papua (e.g., OgilvieGrant 1915). New Guinea’s birds also affected Ernst Mayr, who later formulated ideas about evolution and speciation that form the foundation of modern evolutionary biology. The history of academic ornithological studies date back to the early and mid1800s when a number of seaboard naturalists surveyed natural resources on Pacific islands (e.g., Lesson on the Coquille, Quoy and Gaimard on the Astrolabe, and other ships including the Triton, Basilisk, and Rattlesnake). These early collectors were followed by explorer-naturalists who probed deeper into the hostile terrain of New Guinea in the mid- and late 1800s. Odoardo Beccari explored the Arfak Mountains of the Vogelkop region; Wallace explored the Raja Ampat Islands of western Papua and the Aru Islands; Luigi D’Albertis sailed up the Fly River on three different expeditions; Carl Hunstein explored the Sepik; William Macgregor penetrated the mountains of southeast Papua New Guinea, and numerous government patrols explored Dutch New Guinea. The first significant synthesis of Papuan birds was Count Tommaso Salvadori’s Ornitologia della Papuasia e delle Molucche that appeared in 1880–1882 in three volumes. Following Salvadori, ornithological exploration flowered, and several private collectors funded expeditions to document avian species and collect bird specimens. These include Lord Walter Rothschild of England who sponsored collectors and volumes of ornithological studies based upon the collections. Much of this work was published in Rothschild’s own journal, Novitates Zoologicae, by Ernst Hartert, Erwin Stresemann, William Ogilvie-Grant, Ernst Mayr, and others in the late 1890s through the early 1900s. The work of professional, independent collectors (e.g., A. S. Meek, A. E. Pratt and sons, Andrew Goldie, and others) came to be overshadowed by the large expeditions, such as the Dutch expedition led by Hendrik A. Lorentz in 1909, the British Ornithologists’ Union expedition led by Goodfellow in 1909–1910, and a second BOU expedition led by Wollaston in 1912 (Ogilvie-Grant 1915; Wollaston 1912). After a lull in exploration during World War I (1914–1918), ornithological work resumed in the 1920s, notably by Ernst Mayr, the Pratts, Shaw Mayer, and Stein. Building on the growing museum collections, a major jump forward began with Richard Archbold’s three New Guinea Expeditions between 1933 and 1939. These expeditions brought intensive, multi-taxa collections out of key areas across the island of New Guinea, using amphibious aircraft for support. Perhaps most notable was the Archbold Snow Mountains Expedition of 1938–1939, which procured nearly 4,000 specimens of birds from Papua. Other similar museum expeditions included the Dennison-Crockett and Whitney South Seas expeditions that collected widely across the region. The Archbold Expeditions continued with a brief respite during the Second World War, ending with the seventh in 1964. Biological study matured in New Guinea during the mid-20th century, with numerous publications in the American Museum of Natural History’s Bulletin and Novitates. A

................. 16157$

CH42

02-08-07 10:43:33

PS

PAGE 656

Birds of Papua / 657

journal Novae Guinea appeared and carried many important scientific papers about the region, including on ornithology. After the Dutch departed Papua, ornithological exploration diminished. In the second half of the 20th century most work has been done on short visits by a small number of individuals and survey teams (e.g., Beehler et al. 1995; Diamond 1982b, 1987; Mack and Alonso 2000; Mees 1980, 1982). The need for continued exploration in Papua is greater than ever. Because of its rugged terrain and poor infrastructure, many parts are still relatively unexplored and others have not been visited by ornithologists in more than half a century. Given the growing threats to Papua’s forests from mining, logging, and human expansion (Chapter 7.1), it is important that exploration improve (see Supriatna 1999) and that basic surveys be conducted and specimens collected. Moreover, virtually no long-term studies of Papua’s birds have been undertaken. Our knowledge of the ecology and behavior of Papuan birds is almost entirely derived from field studies in neighboring Papua New Guinea, which has more active and developed research programs. Historical collections are very important for understanding the ornithology of Papua, and the work of these earlier ornithologists has left a legacy of avian taxonomy and systematics that far exceeds that of any other vertebrate or invertebrate group.

Biogeography of New Guinea Birds Biogeography is a science that focuses primarily on understanding and explaining the geographical distribution of organisms. Especially in places as geologically complex as Papua, biogeographical distributions may help explain the origin of bird families, understand the processes of evolution and speciation, and it may help with practical problems such as identifying priority areas for nature conservation. For Papua, we will split the biogeography section into two major sections: the first involves understanding the composition of Papuan bird fauna and its origin in the South Pacific, and the second involves understanding the distribution and ranges of bird species within the island of New Guinea.

origin and composition of papuan avifauna Understanding the Papuan avifauna and its origin has been informed by several recent developments. With recent advancements in geology such as plate tectonics, island-arc dynamics, and sophisticated GIS and dating techniques, we now understand that New Guinea is a composite landmass consisting of the northern margin of the Australian continental plate and at least 32 distinct tectonostratigraphic terranes (Chapter 2.1; Davies et al. 1996; Pigram and Davies 1987; Polhemus 1996). At the risk of repeating concepts treated in greater detail elsewhere in this volume, we shall summarize some geological events and their effects on the current Papuan avifauna. The Australian continent broke away from Antarctica roughly 50 million years

................. 16157$

CH42

02-08-07 10:43:34

PS

PAGE 657

658 / a n d rew m a c k & ja c k d um b ac her

ago (mya) and has been drifting northward to its present position. Because New Guinea is located at the northern fringe of the Australian continent, the avifauna of Papua is inextricably tied to that of Australia. Thus, Australia and New Guinea’s combined avifauna has been relatively isolated from other continents for the last 50 million years or more. Approximately 92% of New Guinea’s bird families are shared with Australia (numbers cited are based on Beehler et al. 1986 and Pratt 1982, with modifications from more recent molecular phylogenetics findings). Prior to the Miocene, much of what is now New Guinea may have been underwater or have formed island archipelagos (Dow 1977; Dow and Sukamto 1984); geologists disagree about how much land was present and how continuous it was with other nearby islands and the continent. Regardless of the details, the Australian continent was certainly the primary source of the present avifauna at least of southern Papua, which is part of the Australian plate. The northern basins of Papua, the Vogelkop peninsula, and the far eastern portion of Papua New Guinea are composed of terranes (tectonic plates) that originated or wandered elsewhere before colliding with New Guinea over the last 25 million years (Pigram and Davies 1987; Pigram and Symonds 1991). Tremendous advances in theory and empirical measurement (through GIS, magnetic positioning, and radio-isotope dating) have allowed geologists to reconstruct the historical movements of many of these terranes (Hall 2001, 2002). It now appears that many of these terranes originated or traveled far out in the Pacific Ocean, and may have had very distinct avifaunas. If a bird family originated in New Guinea or on an oceanic plate, then one would expect that it would have a center of distribution in New Guinea and would be poorly represented elsewhere. Several endemic or nearendemic families show a distribution that suggests that they evolved primarily in New Guinea or Pacific Islands rather than primarily on the Australian mainland. These families include owlet-nightjars (Aegothelidae), birds of paradise (Paradisaeidae), cnemophilines (Cnemophilinae), and berrypeckers (Melanocharitidae and Paramythiidae). An alternative hypothesis would explain the same observed distribution. Because these families are rainforest-adapted birds, they may have evolved in Australian humid tropical forests that were absent or uncommon in other parts of the continent. As the Australian climate became hotter and more arid, forest bird species would have retreated north and east, where rainforest remnants still occur. Because rainforest is now rare in Australia, many of these bird families would now have species or family distributions that are centered on New Guinea. Finally, although Papua is currently located at the eastern end of the Malay Archipelago, Papuan avifauna only weakly reflects its proximity to other Indonesian provinces and to Asian avifauna. Several Asian or Indonesian bird families are noticeably absent from Papua, including woodpeckers (Picidae), babblers (Timaliidae), broadbills (Eurylaimidae), bulbuls (Pycnonotidae), and several other smaller groups. Likewise, many Papuan or Australian groups do not extend into Asia or Borneo, including the birds of paradise (Paradisaeidae), bowerbirds (Ptilonorhynchidae), cassowaries (Casuariidae), owlet-nightjars (Aegothelidae), honey-

................. 16157$

CH42

02-08-07 10:43:34

PS

PAGE 658

Birds of Papua / 659

eaters (Meliphagidae), jewel-babblers (Cinclosomatidae), and others. Furthermore, many of the Asian avian families that do appear in Papua are only sparsely represented, such as thrushes (Turdidae), shrikes (Laniidae), and hornbills (Bucerotidae). Similarly, many primarily Australian lineages are poorly represented in Borneo or Southeast Asia. Alfred Russel Wallace was the first naturalist to understand and appreciate that New Guinea’s avifauna was more allied with that of Australia than with that of most other Indonesian provinces. He identified a line, now known as Wallace’s Line, which demarcates the boundary between primarily Asian fauna and the primarily Australian fauna. The line falls just east of Java and Borneo, and west of the Moluccas, and the line is known to coincide with stretches of deep water ocean that have persisted for millions of years (Wallace 1869, 1876). These oceanic stretches have hindered dispersal of mammals and birds, and thus the two avifaunas (Asian and Australasian) have only weakly mixed.

distribution of the avifauna within new guinea The island of New Guinea is rugged, so populations of bird species can be isolated by a number of geographic barriers. For montane species, mountaintops may be separated by deep valleys or uninhabitable expanses of lowland forest. Lowland populations may be separated by wide rivers, high mountain ranges, or even inland ocean bays. Consequently, populations are often broken into multiple avian subspecies or even separate species, and many species have broken ‘‘checkerboard’’ distributions, separated by regions where the species is absent. The following sections describe biogeographic regions that are important for understanding bird distributions.

Montane Regions Many bird species are limited to high montane regions, and several mountain ranges have endemic species or subspecies. Mountains have long been considered by biologists to be ‘‘islands in the sky’’ separated from other mountains by ‘‘seas’’ of unsuitable lowland habitat. Thus when a bird population on one mountain can be morphologically distinguished from bird populations on other mountains, the populations are often described as distinct species. If variation is weak or poorly understood, then the populations are usually at least given the designation of subspecies. In Papua montane habitats are found in several regions: the Raja Ampat Island highlands (Batanta, Salawati, and Waigeo islands), the Vogelkop Mountains (Tamrau and Arfak), the Bomberai highlands (Fakfak and Kumawa mountains), the Wandammen Range, the Van Rees Mountains, the Foja Mountains, the Cyclops Mountains, the mountains of Yapen Island, and the mountains of the Central Range (Diamond 1985). Although the Papuan land west of Cenderawasih Bay is mostly composed of Gondwanan continental terranes, these plates probably did not arrive in their present position until about 10 mya (Pigram and Davies 1987; Pigram and Symonds 1991). These terranes likely each had a different history, and are home to several

................. 16157$

CH42

02-08-07 10:43:34

PS

PAGE 659

660 / a n d rew m a c k & ja c k d um b ac her

endemic birds. Nine upland species are endemic to these highlands west of Cenderawasih, including White-striped Forest Rail (Rallina leucospila), Vogelkop Scrubwren (Sericornis rufescens), Vogelkop Whistler (Pachycephala meyeri), Vogelkop Melidectes (Melidectes leucostephes), Arfak Honeyeater (Melipotes gymnops), Grey-banded Mannikin (Lonchura vana), Vogelkop Bowerbird (Amblyornis inornatus), Western Parotia (Parotia sefilata), and Arfak Astrapia (Astrapia nigra) (Stattersfield et al. 1998). Eleven additional species have ranges that extend into other mountain ranges, but many of these have distinct subspecies in the western Papuan highlands. The Foja and Cyclops mountains, and the North Coastal ranges (Bewani, Torricelli, and Prince Alexander ranges) form a biogeographically distinct region called the north Papuan mountains (Stattersfield et al. 1998). These are relatively low mountains, but they are separated from the Central Ranges and other mountains by vast stretches of lowland forest in the Sepik and Mamberamo river basins. There are three bird species that are endemic to the north Papuan mountains, including Mayr’s Forest Rail (Rallina mayri), Mayr’s Honeyeater (Ptiloprora mayri), and the Golden-fronted Bowerbird (Amblyornis flavifrons). There are an additional three restricted-range species that occur here and west Papuan highlands and parts of the Central Ranges (see Stattersfield et al. 1998). The Adelbert Mountains and the Huon ranges (Finisterre, Saruwaged, Rawlinson mountains) are primarily oceanic terranes. Although these mountains can be very high (up to 4,100 m in the Finisterre), they are relatively young. The Huon terranes joined New Guinea as early as 10 mya or more (Pigram and Davies 1987; Pigram and Symonds 1991), but the mountains only began forming as recently as 3 mya or later (Abbott et al. 1997). There are only about six endemic bird species found on these mountains, including Huon Melidectes (Melidectes foersteri), Spangled Honeyeater (Melipotes ater), Fire-maned Bowerbird (Sericulus bakeri), Wahnes’ Parotia (Parotia wahnesi), Huon Astrapia (Astrapia rothschildi), and the Emperor Bird of Paradise (Paradisaea guilielmi). Another five restricted-range species have ranges that extend into the Central Ranges. The Central Cordillera extends from just south of Cenderawasih Bay east to Milne Bay in Papua New Guinea, and include the Snow (Maoke, Jayawijaya) Mountains, Star Mountains, Central Highlands, Eastern Highlands, and Southeast Highlands. There are 39 bird species endemic to the Central Cordillera. These include several species that occur only in high alpine grassland areas above ca 3,000 m elevation—a habitat that is absent from nearby islands outside New Guinea. These endemics include the Snow Mountain Quail (Anurophasis monorthonyx), Alpine Pipit (Anthus gutturalis), the Snow Mountain Robin (Petroica archboldi), the Snow Mountain Mannikin (Lonchura montana), and the Alpine Mannikin (Lonchura monticola).

Lowland Regions Several distinct lowland regions also have isolated endemics and restricted-range species. These are often broken into only four major lowland areas: the Bintuni

................. 16157$

CH42

02-08-07 10:43:35

PS

PAGE 660

Birds of Papua / 661

lowlands, the north Papuan lowlands (comprising the Mamberamo and SepikRamu basins), the south Papuan basin, and the Trans-Fly lowlands. The first three are ecologically similar but are separated by high mountains or large bays, and the Trans-Fly is ecologically separated from the adjacent south Papuan lowlands by its reduced and more strongly seasonal rainfall. The Bintuni basin includes the islands of Misool and Salawati as well as the vast lowlands of the Vogelkop and Bomberai Peninsula. These support four endemic bird species, including the Red-billed Brush Turkey (Talegalla cuvieri), the Western Crowned Pigeon (Goura cristata), the Black Lory (Chalcopsitta atra), and the Olive-crowned Flowerpecker (Dicaeum pectorale). The Central Cordillera clearly restricts avian dispersal between the northern and southern watersheds of New Guinea, acting as a formidable barrier to bird movement. Thus the mountains define the northern and southern lowland basins. The north Papuan lowlands include the large Mamberamo River drainage, the Sepik River drainage, and the Ramu River drainage. There are only five species endemic to the north Papuan lowlands: the Brown Lory (Chalcopsitta duivenbodei), Edward’s Fig-parrot (Psittaculirostris edwardsii), Salvadori’s Fig-parrot (Psittaculirostris salvadorii), Brass’s Friarbird (Philemon brassi), and the Pale-billed Sicklebill (Epimachus bruijnii). The southern lowlands are divided by ecological region into the south Papuan basin that is primarily rainforest, and the Trans-Fly that has a strongly seasonal climate with savannah and monsoon forest. Together, these two regions have only six endemic species. Species endemic to the south Papuan basin include the Striated Lorikeet (Charmosyna multistriata), the White-bellied Pitohui (Pitohui incertus), and the Greater Bird of Paradise (Paradisaea apoda); the species endemic to Trans-Fly include the Fly River Grassbird (Megalurus albolimbatus), the Greycrowned Mannikin (Lonchura nevermanni), and the Black Mannikin (Lonchura stygia). We believe, however, that the real diversity of lowland taxa may be underappreciated by this species-level analyses. Many lowland species show a surprising level of intraspecific morphological variation, and some lowland species have as many as 20 or more described subspecies. For example, the Variable Pitohui (Pitohui kirhocephalus) is tremendously variable in coloration and size, and has 20 described subspecies in New Guinea and the nearby continental islands; the Little Shrike-thrush (Colluricincla megarhyncha) has up to 23 subspecies by some accounts, although the morphological variation is less pronounced (Rand and Gilliard 1967). We believe that this large number of subspecies may be due, in part, to biases of taxonomists working in Papua, because despite significant morphological variation among lowland populations, the populations have traditionally been lumped into single species rather than being split into multiple species. This has been done for two reasons. First, it is assumed that lowland populations are not isolated from other nearby lowland forests. Lowland forests form a continuous ring around the island, and even the most isolated valleys have corridors of lowlands connecting them to larger adjacent populations. Therefore, it was assumed that there was substantial interbreeding among the different subspecies or that their records were just points along a

................. 16157$

CH42

02-08-07 10:43:35

PS

PAGE 661

662 / a n d rew m a c k & ja c k d um b ac her

continuous species cline. Second, at the time when many large taxonomic revisions were done, sampling was relatively sparse. Thus it was assumed that as more samples were made, more intergrading populations would be found in the forest corridors between the major basins. Splitting such taxa is common for isolated montane species, but has not been done in lowland taxa in part because there is a presumption that there must be some gene flow, and that intergrading populations would be found if there were greater sampling. We surveyed the putative ranges of 120 subspecies from 28 avian polytypic species (based on Rand and Gilliard 1967), and plotted the ranges of the subspecies on a map of New Guinea (see Figure 4.9.1). Many subspecific boundaries appeared to coincide. For example, nearly 30 subspecies’ ranges bordered on the Huon mountains, suggesting that Huon Peninsula is an important vicariance barrier (a barrier that prevents gene flow), and many more ranges were divided at the Vogelkop isthmus. Other discontinuities were more surprising: five subspecies’ boundaries coincided with the border mountains in the north Papuan lowlands, and 15 subspecies’ boundaries coincided with a hypothesized inland bay called the ‘‘Aure trough’’ (Pigram and Davies 1987). Many of these same natural boundaries were appreciated much earlier (Hartert et al. 1936) but have not received a great deal of comment in recent works. We are actively examining genetic differentiation among these lowland basins. Our preliminary data suggest a significant split between the north and south coasts in several species, including Colluricincla megarhyncha, Pitohui kirhocephalus, the lowland owlet-nightjar, Aegotheles bennettii (Dumbacher and Fleischer 2001; Dumbacher et al. 2003), and certainly others. The genetic divergence among the Pitohui kirhocephalus groups is quite marked, and probably warrants splitting them into separate species: a western Papuan Island group, a south coast group, and a north coast group. Interestingly, members of both the north coast and south coast groups are present in the Bintuni Basin, suggesting that both of these groups may have invaded after the Bintuni terranes joined the main island of New Guinea. It is easy to imagine the impact of a mountain range that forms and splits a population in two, or an inland bay that separates coastal populations. But more complex biogeographic processes may be taking place on the north coast of New Guinea, as a consequence of the Australian plate moving northward. We now believe that many of the northern islands (e.g., New Britain and New Ireland) are on course to collide with New Guinea in the future. Because birds are good overocean dispersers relative to many other vertebrates, they are capable of moving to offshore islands or volcanic island arcs as they become closer to the mainland (Diamond 1973). Once on these islands, birds can evolve independently and potentially become different enough to become new species. Later, when the island collides with the mainland, the island species will come into contact with the mainland species. Some will merge into one gene pool, some may go extinct, and

................. 16157$

CH42

02-08-07 10:43:36

PS

PAGE 662

Birds of Papua / 663

Figure 4.9.1. Map of New Guinea lowlands. Areas over 1,000 m elevation are shown in gray. BB: Bintuni Basin; MB: Meervlakte Basin; SB: Sepik Basin; RB: Ramu Basin; CVB: Cape Vogel Basin; SPP: Southeast Papuan Peninsula; SPB: South Papuan Basin; TF: Trans-Fly lowlands. We surveyed the ranges of 120 subspecies from 28 avian polytypic species, and the numbers on this figure correspond to the number of avian subspecies whose ranges terminate at this locality. Lowercase letters following the numbers denote hypothesized geologic barriers or dispersal routes at these localities. at: Aure Trough; mb: Milne Bay Break; hp: Huon Peninsula; bm: border mountains; wm: Wandammen Mts; ab: Aetna Bay. Dashed gray line depicts extent of the Trans-Fly (tf) lowlands. The vertical line down the center is the political boundary between Papua, Indonesia (to the west), and Papua New Guinea (to the east). some may persist as different species for long periods or even spread across the mainland. We believe that the processes of island formation, island colonization, and island accretion can account for some of the rich biodiversity of Papua, especially in the northern basins and northern mountain ranges, such as the Huon, Adelbert, Torricelli, and others. Likewise, New Guinea has served as a bird colonization source for thousands of islands throughout the tropical Pacific. Detecting such histories may be difficult, because for any single case, there will be alternative historical explanations. Several species or groups have strong ecological preferences that limit their geographic distribution. One species, the Greater Melampitta, Melampitta gigantea, inhabits limestone karst habitat that is limited but widespread. These melampittas prefer to build nests in cave tunnels at the bottom or sides of limestone sinkholes,

................. 16157$

CH42

02-08-07 10:43:52

PS

PAGE 663

664 / a n d rew m a c k & ja c k d um b ac her

from which they emerge during the day to forage (Diamond 1983; J. Dumbacher, pers. obs.). Melampitta’s distribution is likely limited somewhat by the availability of suitable habitat, although it has been recorded in non-karst regions in the Bewani Mountains. Similarly, the finch Lonchura montana occurs in montane grasslands and is generally restricted to those habitats. But often in tropical environments, bird species have patchy distributions even with respect to their preferred habitat, and this is especially true in Papua. The causes of the patchiness may be due to historical vicariance or barriers to gene flow (Heads 2001a,b,c), ecological exclusion of similar competitive species in different areas (Diamond 1973), or simply to the natural process of local extinctions and recolonization (Diamond 1973). The significance of patchiness is discussed in detail in Diamond (1973).

Ecology Most Papuan bird species live in just one of the main habitat categories (described in detail in other chapters): marine, mangrove, aquatic, savanna, wet tropical forest (lowland through upper montane), and alpine habitats (Table 4.9.2). Some species may occur across habitat types, particularly aerial hunters such as swifts and raptors, but most species have fairly specific habitat requirements. To some extent, the overall diversity of birds in a region depends on the diversity and extent of available habitats and the niches contained therein. Many neotropical sites have greater niche diversity than New Guinea (Pearson 1977; Ridgely et al. 2005).

Table 4.9.2. Primary habitat preferences of Papuan birds Number of species Preferred habitat

Percent of Papuan species

Pelagic

16

2.4

Aquatic (incl. lowland and montane freshwater)

64

9.7

Coastal (incl. nearshore, beach, mangrove, coastal forest, estuary)

65

9.9

Savannas

59

9

Grasslands (incl. lowland and montane grasslands)

47

7.2

212

32.3

Lowland rainforest Hill forest

55

8.4

130

19.8

Alpine

6

0.9

Towns/other

3

0.5

Montane forest

................. 16157$

CH42

02-08-07 10:43:54

PS

PAGE 664

Birds of Papua / 665

elevation It is well known from studies around the world (e.g., Kitayama 1992a,b) that forest composition and habitat change with elevation, though the proximate causes of these changes are not always apparent (Ashton 2003). The rugged Central Cordillera running the length of New Guinea creates a fairly complex mix of forest vegetation types within relatively small areas (Hyndman and Menzies 1990; Johns 1976; Paijmans 1970, 1976). Elevation generally affects distribution of birds (Lomolino 2001; Patterson et al. 1998; Rahbek 1997; Sanchez-Cordero 2001), and this is true for New Guinea as well (Beehler 1982; Diamond 1972, 1973). Diamond (1972) describes in detail how elevation helps partition bird communities along elevational gradients on Mt Karimui, PNG. He found that many genera contain species that segregate by elevation; good examples include the species of Ptilorrhoa, Rhipidura, Sericornis, and Crateroscelis. It is logical that closely related congeners would share similar niches and ecological requirements and thus would compete (Diamond 1978); the principal of competitive exclusion (Hardin 1960) would lead them to segregate.

diet and guilds The morphology, physiology, and behaviors of birds are strongly influenced by their diets (Brandl et al. 1994; Karasov and Levey 1990; McNab 1988). In New Guinea, relatively few studies have examined relationships between diet and morphology, behavior, and physiology (McNab 2005; Wooller and Richardson 1988; Wooller et al. 1990), but these suggest that adaptations for different diets are similar to those found in other birds worldwide. Although it is difficult to unambiguously break the continuum of avian feeding preferences into discrete categories, a rough breakdown reveals some general trends (Table 4.9.3). The species that are strongly insectivorous (listed as insectivores under heading of carnivore in Table 4.9.3) comprise over 42% of the avifauna of Papua. Those terrestrial species that would be considered typical carnivores, specializing on vertebrate prey, comprise just over 7% of the Papuan avifauna. The following sections address the major trophic groups.

Carnivores and Predation Because of the absence of large mammalian predators in New Guinea, large raptors probably play an unusually important role on the island. There are 40 species of raptors (including owls) in Papua. Most notably, the New Guinea Harpy Eagle (Harpyopsis novaeguineae) is one of the largest terrestrial predators on the island of New Guinea; only some varanids and snakes are larger. It occurs from near sea level to the tree line at 3,200 m, and feeds predominantly on ringtails, cuscus, and forest wallabies, though it preys on other mammals, megapodes, reptiles, and juvenile cassowaries as well (Beehler et al. 1992; Watson and Asoyama 2001). The status of raptors in Papua is not well known, but presumably most are not as threatened as raptors from the more densely populated parts of Indonesia (van Balen 1998).

................. 16157$

CH42

02-08-07 10:43:55

PS

PAGE 665

666 / a n d rew m a c k & ja c k d um b ac her

Table 4.9.3. Trophic specializations of Papuan birds Primary dietary category

Number of species

Carnivores (n 410) Insectivore (and some other invertebrates) Insectivore (and some vertebrates) Insectivore (and some fruit and seeds) Insectivore (and some nectar) Aquatic and marine invertebrates Terrestrial vertebrates (and some insects) Fish (and some invertebrates)

214 26 24 10 44 47 45

Frugivores (n 121) Fruits Fruit and insects (and some vertebrates) Fruit and seeds (and some insects)

39 54 28

Granivores (n 37) Seeds Seeds and insects (and some fruit)

22 15

Herbivores (n 17) Plant matter and insects (incl. aquatic)

17

Nectarivores (n 49) Nectar Nectar and fruit Nectar and insects (and some fruit)

6 19 24

Omnivores (n 18) Unclassifiable*

8

Note: Bird species of Papua for which we have data (see Appendix 8.2) assigned broad dietary categories. * Combines three or more of the above categories, without an obvious strong emphasis on one Source: summarized in Coates 1985, 1990; personal observations.

Among the carnivores (Table 4.9.2) we have included the piscivores (fish eaters). Papua has at least 15 species of herons and egrets, most of which eat significant amounts of fish. Other piscivores include grebes, a pelican, a darter, cormorants, osprey, terns, and some kingfishers. All of these are vulnerable to transformation of wetlands for agricultural purposes, sedimentation from mining, or the damming of rivers. Transformation of wetlands to rice production does not provide good habitat for piscivorous ardeids, though paddies can be suitable for the introduced Cattle Egret (Richardson and Taylor 2003). As top predators in the aquatic food chain, piscivorous birds are known to accumulate certain toxins that humans release into the environment, particularly organochlorides. During the period of unregulated DDT use in the United States, populations of piscivores such as pelicans, Bald Eagles, and cormorants plummeted in numbers due to the egg shell thinning caused by toxic accumulations of organochlorides in the laying females (Hickey and Anderson 1968). Use of these

................. 16157$

CH42

02-08-07 10:43:55

PS

PAGE 666

Birds of Papua / 667

pesticides was banned in the United States, but they are still used in many parts of the world. When properly applied these pesticides can be a potent weapon in the fight against malaria and other mosquito-borne diseases. However, without proper application, damage to birds can be unnecessarily high. In some parts of the world, populations of piscivorous birds are still suffering from the release of organochlorides to the aquatic environment (Connell et al. 2002, 2003; De LucaAbbott et al. 2001). There are few data or tests from Papua, but given the push to increase industrial agricultural outputs of rice and oil palm, it seems prudent to voice concern over how pesticides will be applied.

Pelagic and Migrant Species Very little is known of the pelagic birds (species that live in the open ocean) that occur near Papua. The most relevant data come from the seas to the west of Papua (e.g., Cadee 1985). As far as we know, there are no breeding colonies of Procellariform species in Papua, but several pelagic terns and boobies are known to breed in eastern Indonesian waters. Because many of the pelagic species found in the oceans off Papua are widespread and better known from other parts of the Indian and Pacific oceans, relatively little attention has been devoted to them, in favor of attention to the endemic and unique terrestrial avifauna. Seventy-five species of migratory birds are listed under treaties to protect migratory birds between Australia and Japan (Japan-Australia Migratory Bird Agreement) and Australia and China (China-Australia Migratory Bird Agreement). All of these migratory species are known to, or could be expected to, also pass through or stay in Papua for at least some short period. Additionally, there are roughly an additional 65 species of terrestrial and aquatic species that are known to, or could be reasonably expected to, move between New Guinea and Australia (Dingle 2004) (Appendix 8.2). The number of birds moving between Australia and New Guinea is large enough that conservation of some ‘‘Australian’’ species requires more information about and management in the New Guinea ‘‘wintering’’ grounds (Legge et al. 2004).

Frugivores and Seed Dispersal The avifauna of the Papua, and New Guinea in general, has an exceptionally high number of frugivorous species (Pearson 1977) that play crucial roles as dispersers of seeds. Some scientists have speculated that in the absence of many potential mammalian competitors (e.g., primates, squirrels) frugivorous birds have diversified more here than in other tropical regions. Roughly 90% of the woody plants in the New Guinea rainforests produce fleshy diaspores that appear to be adapted for dispersal by birds and Pteropodid fruit bats (Mack and Wright 2005). In the absence of dispersal by birds, many plant species would not be able to sustain their populations (Mack et al. 1999; Mack and Wright 2005). For a combination of reasons, many trees from Papuan rainforests have unusually large diaspores (Mack 1993). Thus a relatively small number of bird species—the three cassowaries C. casuarius, C. unappendiculatus, and C. bennetti and the hornbill Rhyticeros plica-

................. 16157$

CH42

02-08-07 10:43:56

PS

PAGE 667

668 / a n d rew m a c k & ja c k d um b ac her

tus—are inordinately important for the maintenance of diverse floras in Papuan forests (Mack and Wright 2005). If the birds become extirpated, one could expect a gradual loss of plant diversity. Where these four bird species are heavily hunted there is the potential for extirpation. Fortunately, at least some of these species appear to have large ranges and seasonal movements that might reduce the risk of local extirpation. Some tropical forest trees, particularly in the Myristicaceae and Meliaceae, have fairly narrow sets of frugivorous birds that can extract seeds from their capsules, so maintenance of these tree species populations depends on the conservation of a small number of frugivorous bird species (Beehler and Dumbacher 1996). The frugivorous avifauna includes a large number of pigeons and doves. At some localities 20 or more species of columbids can be sympatric (Beehler et al. 1995; Bell 1982b; Mack and Wright 1996). This presents a classic example of niche partitioning, with species in closely related taxa (e.g., Ptilinopus spp.) dividing up the fruit resource based on where they forage and their body size (Bell 1983a; Diamond 1973, 1978; Frith et al. 1976; Pratt 1984; Pratt and Stiles 1983; Terborgh and Diamond 1970). Since these seminal papers, little research had been conducted with columbids or the issues of niche partitioning by frugivores in New Guinea. Intensive study in this region of the ecology of pigeons, and especially their competitive interactions and resource partitioning, would be highly rewarding (Pratt 1984; Pratt and Stiles 1983). Figs (Ficus spp.: Moraceae) present a special fruit resource for tropical frugivores (Shanahan et al. 2001). In Papua several bird species specialize in consuming figs (Beehler 1982, 1989; Beehler and Dumbacher 1996), including the Vulturine Parrot (Mack and Wright 1998), the Manucodes (Beehler 1985; Frith and Beehler 1998), and some cuckoo-shrikes. Figs provide a nutritious resource that is often reliable due to its asynchronous fruiting phenology (Beehler 1985; O’Brien et al. 1998), and the movements of birds tracking figs or other fruit resources can play an important role in forest generation (Kinnaird et al. 1996). Protection of keystone fruit resources and other important fruiting species could be important to help promote forest regrowth and recovery after logging (Hamann and Curio 1999; Holbrook et al. 2002).

Nectarivores and Pollination An important component of the avifauna of Papua consists of nectarivorous birds, particularly the large radiations of honeyeaters (Meliphagidae: 55 spp.) and lories (Loriinae: 19 spp.). Based on what is known from other tropical forests and the large number of ornithophilous flowers in Papuan forests, it is safe to assume nectarivorous birds play a crucial, but not well understood, role. In one of the few studies of nectar feeding birds in PNG, roughly 20% of the flowering trees in a plot were visited by nectar feeding birds (Brown and Hopkins 1996) and 13% of the bird species at the site fed upon nectar (Brown and Hopkins 1995). The proportion of rainforest trees that are visited by nectarivorous birds and are probably pollinated by them is higher in New Guinea than in most tropical forests. This

................. 16157$

CH42

02-08-07 10:43:56

PS

PAGE 668

Birds of Papua / 669

difference is probably due to the taxonomic affinities of nectarivores in different regions. In Papua, most nectarivorous birds are honeyeaters and lories that feed mostly on canopy trees, whereas in Latin America, most are hummingbirds that feed mostly on herbs, vines, and epiphytes (Brown and Hopkins 1995). This difference in the ecology of New Guinea forests and neotropical forests is quickly apparent to anyone who has visited both. The large number of nectarivorous birds that congregate in flowering trees in New Guinea are a diverse mixture of many genera with dramatically different morphologies, from tiny myzomelids to large lories. In the cases of neotropical ornithophilous trees, most avian visitors are small hummingbirds and other small passerines (e.g., Chlorospiza, Dacnis). In New Guinea competition may be reduced by honeyeater species using different strategies within the same tree (Collins and Paton 1989) and by the birds’ different morphologies (Paton and Collins 1989). But, as evinced by the fighting and displacements by different species in such aggregations, competition is strong. Indeed, two Myzomela species that have only recently (300 years) come into contact exhibit evidence of character displacement, probably as a result of competition for nectar resources (Diamond et al. 1989). Competition by birds affects pollen movement, seed set, and outcrossing (Ramsey 1988, 1989), but is unexamined in New Guinea where it could have important implications for economically important tree species (Brown and Hopkins 1995). In New Guinea there is little evidence of fairly tight mutualisms between any plant and a bird species or genus (Brown and Hopkins 1995). Most New Guinea nectarivores probably feed upon a diversity of plant taxa and flower types, which is more typical of plant-pollinator systems (Waser et al. 1996). However, this does not imply birds are not important as pollinators, and it will be important to conserve the full suite of nectarivores in a rainforest in order to ensure plant reproduction continues successfully (Bawa and Krugman 1991; Bond 1994; Kearns et al. 1998). It is difficult to assess the importance of birds as pollinators because virtually no studies have examined birds and pollination in the region. We are unaware of any study in the New Guinea region that explicitly quantifies how seed sets are affected by avian pollinators. Nectar-feeding birds can become abundant locally when key plants are in flower, then emigrate when flowering terminates (McGoldrick and MacNally 1998). The scale of local movements of such birds tracking resource availability in New Guinea is unknown, but eruptions of nomadic or migratory nectarivorous birds seem timed with large flower crops that were often asynchronous with less copiously-flowering species (Bell 1982a; Brown and Hopkins 1996). Given the right circumstances of nectar production and density of flowers, nectar resources can be temporarily defended by aggressive nectarivores (Beehler 1994), as sometimes also happens with fruit resources (Pratt 1984). Detailed long-term studies of bird populations, coupled with phenological studies, are needed to elucidate what appears to be complex patterns of movements by nectar-feeding birds, particularly lorikeets.

................. 16157$

CH42

02-08-07 10:43:57

PS

PAGE 669

670 / a n d rew m a c k & ja c k d um b ac her

Insectivores The large number of insectivorous species at any one site would lead one to suspect competition could be intense. However, insects and the places they occur are extremely diverse and thus provide many distinct foraging niches that different species of insect feeding birds could specialize upon. In New Guinea this seems to be the case, with species sorting by size, substrate, and vertical distribution (Bell 1982c, 1983a; Croxall 1977). Birds have a variety of hunting styles, from aerial hawking species (e.g., swifts), to ground-dwelling species that search the forest litter for insects (e.g., Pittas). Just as frugivores and nectarivores provide economically valuable ecosystem services as seed dispersers and pollinators, insectivorous birds provide a vital service in control of insect pests. Birds can play a role in pest management of simple temperate agricultural systems (Mols and Visser 2002; Tremblay et al. 2001). More recently there has been evidence that insectivorous birds are important for regulating insect pests in tropical agroforestry, cacao, and coffee plantations (Perfecto et al. 2004, Philpott et al. 2004). However, only recently has evidence emerged that birds can be important in reducing insect pests in tropical forests (van Bael et al. 2003; van Bael and Brawn 2005). Although data from New Guinea are not available, given the great density and diversity of insectivorous birds in the region (Table 3.9.2) and at specific sites (Bell 1982b; Mack and Wright 1996), insectivorous birds are probably economically important in Papua for the role they play controlling insect pests in agricultural and agroforestry systems.

birds of paradise (paradisaeidae) The study of avian behavior in the New Guinea region is dominated by studies of the birds of paradise. This short chapter cannot cover the detailed studies of birds of paradise by B. Beehler, C. Frith, D. Frith, T. Gilliard, M. LeCroy, J. Diamond, T. Pratt, M. Pruett-Jones, and S. Pruett-Jones, but all the work is comprehensively summarized and synthesized in Frith and Beehler (1998). The birds of paradise have been the focus of so much study because they are models for studies of strong sexual selection (Diamond 1981), hybridization (Fuller 1995), courtship and lek display behaviors (Beehler 1988; Frith and Frith 1988; Pruett-Jones and PruettJones 1988), sexual dimorphism and extravagant male plumages (Frith 1981), frugivory (Beehler 1989), and polygamy (Beehler 1987). There is much traditional lore about the birds (Healey 1993) and local people use their feathers as adornment (Swadling 1995). Within the birds of paradise there is great diversity. Taxa range from primarily monogamous (e.g., Manucodia spp.) to promiscuous polygamous species (e.g., Paradisaea spp.). There are highly frugivorous taxa (e.g., Paradisaea spp.) and others that are nearly wholly insectivorous (e.g., Drepanornis albertisi). A few are territorial while most do not appear to defend territories. Males may display as isolated individuals, or in collective display areas called leks, in which many males display within close proximity with interactive dances and vocal behavior. The stronger the sexual selection in a species (i.e., females selecting a small proportion

................. 16157$

CH42

02-08-07 10:43:58

PS

PAGE 670

Birds of Papua / 671

of the males for the majority of copulations), the stronger the selection for specialized plumage, dances, and vocalizations (Christidis and Schodde 1993; Diamond 1981; Emlen and Oring 1977). This selection appears to have led to a fairly rapid evolution of diverse morphologies, but with relatively little genetic divergence. Thus the phylogenetic branches between quite different-looking taxa are short and poorly resolved (Nunn and Cracraft 1996), suggesting that some taxa have evolved more rapidly in morphological and behavioral characters than in other characters. The large number of recorded hybrids (Frith and Beehler 1998; Fuller 1995), even among different genera of birds of paradise (e.g., Frith and Frith 1996), supports this conclusion.

bowerbirds (ptilonorhynchidae) The second avian family that has received great attention from ornithologists (e.g., G. Borgia, J. Diamond, C. Frith, D. Frith, T. Gilliard, M. Pruett-Jones, and S. Pruett-Jones) is the bowerbird family (Ptilonorhynchidae), much of which has been incorporated in an excellent review of the family (Frith and Frith 2004). The bowerbirds are exceptional because in many species the males construct structures that act as attractions, display arenas, and courting grounds. These ‘‘bowers’’ can be anything from a cleared area with a few leaves placed for decoration, up to massive structures of twigs, decorated with hundreds of colorful items like fruit or flowers, that take months to construct. One of the notable things about the bowerbirds is how they use ornaments around their bower as attractants to females. Often these ornaments are hard-tofind, colorful items that are replaced often as they fade or spoil (e.g., flower petals and colored fruits). Males spend considerable effort collecting these articles and defending their decorations from marauding male conspecifics, so the degree of decoration can act as a measure of male quality to a potential female mate (Diamond 1982a; Diamond 1986; Hunter and Dwyer 1997).

nidification Most tropical passerine birds lay only two eggs at a time, but might have several nesting attempts in a year if the first is unsuccessful. Rates of nest predation are typically high in rainforests and rainforest birds may re-nest several times before successfully fledging. The nidification (i.e., nest building) of many Papuan bird species has not been described (Coates 1985, 1990), and there are very few detailed studies of nesting biology and nest success (e.g., Frith and Frith 1993, 1994). A little appreciated ecological role of some birds in Papua is that some, most notably Psittacids, excavate and enlarge cavities in trees. Because there are no woodpeckers in the region’s rainforests, the wide diversity of species that occupy nest holes (from rats to arboreal marsupials to hornbills) might rely on the relatively few avian species that actively excavate cavities. For many species, suitable cavity nest sites could be a major limiting factor to populations (Gibbs et al. 1993). One consequence of logging operations can be the removal of large trees that have cavities. The hunting practice of chopping down trees with cavities in them to

................. 16157$

CH42

02-08-07 10:43:59

PS

PAGE 671

672 / a n d rew m a c k & ja c k d um b ac her

extract prey also reduces availability of snags and nesting cavities. Reduction of snag and cavity availability through these activities could have serious deleterious effects on hole-nesting species of birds, especially parrots (Monterrubio et al. 2002; Seixas and Mourao 2002), and mammals (Laurance and Laurance 1996). These effects have not yet been assessed in New Guinea. Undoubtedly the New Guinea birds with the most fascinating nesting biology are the megapodes. This family of birds is unique in that all species use some form of environmental heat to incubate their eggs—neither the male or female broods the eggs, which makes them similar to the putative ancestor to modern birds. Much of what is known about megapodes has been summarized by Jones et al. (1995), although there has been some more recent work in the New Guinea area since the book was published (Sinclair 2000, 2001; Sinclair et al. 2002). Some species use their large, strong feet to scrape together large mounds of forest leaf litter, which generates heat in the center as it decomposes. The eggs are laid in the heart of these large mounds, covered, and allowed to incubate. In other species, the female lays her eggs in earth heated by geothermal sources or, in some instances, in solar-heated sand. In all cases, the young hatch, dig their way to the surface, and then are completely self sufficient. Some young can fly within 24 hours of emerging from their subterranean nest site, making them the most precocial birds in the world, with absolutely no post-hatching parental care. Because the chick develops completely without parental care, the female invests a great deal in egg yolk. Megapode eggs are the most nutritious eggs laid by birds. This, combined with the conspicuous and specialized places where they lay their eggs, means that human hunters highly prize megapode eggs. They are a nutritious delicacy for people across New Guinea, so megapodes are under heavy pressure from people wherever they live near humans. Indeed the Maleo, a megapode species from Sulawesi, is highly endangered from over-exploitation (Baker and Butchart 2000; Butchart and Baker 2000). Megapodes were once much more numerous and diverse across the Pacific region, but human hunters and possibly rats helped wipe them out in many places, and many species have already gone extinct (Steadman 1991; Steadman et al. 1999, 2002). Without proper management, many other megapode species could follow the Maleo to extinction. The Waigeo Brush Turkey (Aepypodius bruijnii), a megapode endemic to upland Waigeo Island, Papua, is one of the most highly endangered species in the region.

mixed-species flocks Around the world many birds form mixed species flocks (Ficken 2000; RagusaNetto 2002; Styring and Ickes 2001; Thiollay 1999, 2002; Vuilleumier 1967) and the birds of New Guinea are no different. Flocks can be aggregations at locally rich resources, as is common with nectar- and fruit-eating species. At the dense fruit or nectar resource many species can compete through exploitative and interference competition (Terborgh and Diamond 1970). Where resources become sufficiently abundant, it is possible that normally nonterritorial species temporarily defend

................. 16157$

CH42

02-08-07 10:44:00

PS

PAGE 672

Birds of Papua / 673

resources sufficiently rich to justify defense (Pratt 1984). How long such nectarivores and frugivores remain at the resource helps determine rates of pollen flow and seed dispersal (Pratt and Stiles 1983). But of perhaps greater interest are the groups of primarily insectivorous species that seem to move in loose parties through the forest but that employ different strategies to find insects. Some search foliage, some examine limbs and trunks, while others sally and grab flying insects (Bell 1983b). Whether joining flocks improves survivorship is difficult to tell, because many variables must be controlled, but at least in some cases survivorship for neotropical flocking species is higher than for non-flocking species (Jullien and Clobert 2000). Higher survivorship could be due to greater vigilance and predator avoidance (Thiollay 1999), or that foraging birds act as ‘‘beaters,’’ flushing insects that other individuals capture, thereby conferring a foraging advantage to insectivores that travel in groups rather than solo.

Systematics Avian systematics is experiencing a revolution as a result of recent advances in molecular biology. Although there are still relatively few studies that focus exclusively on Papuan birds, many studied avian groups have important centers of diversity in Indonesia and Australia. Study of these groups should help clarify our understanding of the evolution of avian diversity across the region. Among the most recent and important are two studies that examine the systematics of the Passeriform radiation (songbirds, or perching birds; Barker et al. 2002, 2004). Their work shows that the basal Oscine lineages appear to have evolved in Australasia, and that many of the most basal groups (Menuridae, Ptilonorhynchidae, Climacteridae, Meliphagoideae) have centers of diversity in Australasia, often centering on or near New Guinea. Twelve species of berrypecker, placed in two families, Melanocharitidae and Paramythiidae (Sibley and Monroe 1990), are the only widely recognized bird families endemic to New Guinea, although some other Papuan radiations may soon be recognized as families (e.g., Cnemophilinae, sensu Barker et al. 2004). The number of species recognized in Papua is liable to change substantially when modern phylogenetic analyses are undertaken. This is only in part the result of the much narrower species definition that is replacing the broader polytypical concept that dominated in the 1960s and 1970s. For example, within two New Guinean genera (Aegotheles and Pitohui) that have been studied using modern phylogenetic techniques, three to four new species have been revealed (Dumbacher and Fleischer 2001; Dumbacher et al. 2003; Pratt 2000). Many more groups currently viewed as species will likely be found to be composed of genetically welldifferentiated taxa that have been unrecognized because they are morphologically similar or poorly represented in systematic collections. Additional species, particularly pelagic species, might be expected to occur in Papua, but have not yet been recorded.

................. 16157$

CH42

02-08-07 10:44:01

PS

PAGE 673

674 / a n d rew m a c k & ja c k d um b ac her

There are probably relatively few new species remaining to be discovered in Papua, but scientists are likely to name new species of birds based on examination of specimens. Collection of additional specimens or closer examination of existing specimens is likely to reveal differences that have not been appreciated. A recent example is the Starry Owlet-nightjar (Aegotheles tatei), that has been elevated to species, based on a study of several definitive morphological characters that revealed significant differences. Several other groups probably need revision (e.g., Chaetorhynchus, Petroicidae, Melampitta, Pachycephalidae, and Ptilorrhoa).

Research Needs Field research has declined in Papua since the 1960s. Its avifauna is one of the least-studied in the world partly because of the inaccessibility of much of the region, but also because of the shortage of experienced ornithologists adequately familiar with the avifauna (Beehler et al. 1995). What recent knowledge we have of the Papuan birds comes mostly from research in the PNG side of the island. Some survey work is still being completed in PNG, but surveys in PNG are poor substitutes for elucidating the distribution of birds in Papua. Very few formal surveys have been published from Papua in the past twenty years (Diamond 1982b; Mack and Alonso 2000; Richards and Suryadi 2002) and only a handful of informal accounts from short visits by ornithologists and birdwatchers have appeared (e.g., Eastwood 1996; Gibbs 1994; Melville 1979, 1980). A large amount of information was collected when planning protected areas in Papua (Petocz et al. 1983; J. Diamond, unpub. data), but these were mostly short surveys in areas proposed as protected areas. Thus, despite some recent fieldwork, there is still considerable room for improvement in the state of the overall knowledge of avian distributions in Papua. Better knowledge of distributions of birds (as well as other taxa) is needed to guide land use planning and conservation actions (e.g., Benayas and de la Montana 2003). Without a fundamental knowledge of the distribution of biodiversity in Papua, site-based conservation interventions can be little more than best guesses (Supriatna 1999). The second major deficiency in knowledge that will hamper conservation and management needs comes from ignorance of basic natural history and ecology. For management of hunting offtake (e.g., megapode egg harvesting), it is necessary to know basic natural history—population density, home range size, longevity, age of first reproduction, and annual reproductive output (Alvard et al. 1997; Novaro et al. 2000; Robinson et al. 1999; Sanderson et al. 2002). Traditional use of cassowaries is probably not sustainable in many areas, but we lack the basic natural history data to conduct population viability analyses and to make informed management decisions (Johnson et al. 2004). Basic ecological knowledge is required in order to manage ecosystems, both in protected areas and in other land use zones, in order to minimize impact (Brussard 1991). The third area in which research would be beneficial is the realm of pure science. Many important scientific innovations are derived from studies of birds in

................. 16157$

CH42

02-08-07 10:44:02

PS

PAGE 674

Birds of Papua / 675

New Guinea (Diamond 1973; Diamond et al. 1989; Mayr 1963; Mayr and Diamond 1976; Wallace 1869). Because New Guinea has a unique biota with an evolutionary history independent of most of the rest of the world, it offers a special opportunity both to test ideas that are derived from other areas without merely being pseudoreplication (Hurlbert 1984) and to generate novel ideas that can be applied to other parts of the world (Westoby 1988). Better knowledge of avian distributions can help elucidate the complex evolutionary history and biogeography of the region (Heads 2001a,b, 2002). We do not have good basic distributional data for any widespread group of organisms in New Guinea except the birds, so improved knowledge of avian distributions provides the best hope for generating novel findings in the fields of biogeography and evolutionary biology.

Conservation Papua is home to a substantial number of endemic species that are primarily confined to small islands (e.g., Aepypodius bruijnii, Megapodius wallacei, Eos cyanogenia, Centropus chalybeus, Myiagra atra, Monarcha julianae) or, in some cases, to small areas on the mainland, particularly certain montane species (e.g., Anurophasis monorthonyx, Sericornis rufescens, Petroica archboldi, Lonchura teerinki). Many of these endemics are included among the 129 species from the New Guinea region listed by the IUCN and BirdLife International at some level of conservation concern, and 74.4% of the listed species are endemic to the New Guinea region (Appendix 8.2). Papua still has a relatively small human population and large areas of intact forest, but conservation of this province is threatened by Indonesia’s rapid human population growth and increasing exploitation of natural resources. Conservation threats to birds in Papua can be classified in five general categories, each of which we discuss below.

traditional and subsistence use of birds The people of Papua have developed hundreds of distinct cultures, each with its own traditions, myths, and uses of wildlife, in which birds play a central role. Traditional adornment for ceremonies, weddings, and rituals often includes bird plumage, sometimes in large quantities (Healey 1986; Heaney 1982). The group most used for traditional adornment, the birds of paradise, was also at one time very heavily exploited for the millinery trade overseas. At the peak of the trade, export of bird of paradise plumes was measured in tons of feathers per year, yet no species went extinct (Swadling 1995). This is because only fully-plumaged males were taken and subadult males could reproduce in their absence, and because the most desired species (Paradisaea spp.) are typically widespread from the lowlands to middle elevations. Traditional plume usage is liable to have an impact on species that are strongly and widely desired (e.g., Harpyopsis novaeguineae, Psittrichas fulgidus, Epimachus fastuosus; Mack and Wright 1998; Watson and Asoyama 2001) or which might be less desired, but have restricted ranges or populations (e.g., Cicinnurus respublica).

................. 16157$

CH42

02-08-07 10:44:02

PS

PAGE 675

676 / a n d rew m a c k & ja c k d um b ac her

Although larger mammals are particularly targeted by subsistence hunting, many bird species also face serious pressure from hunting (Mack and West 2005). Because the human population is growing rapidly in Papua and there is little domestic livestock or access to commercially produced meat, wildlife faces inordinate pressure from subsistence hunting. In most of rural and montane New Guinea the majority of people obtain a significant portion of their dietary protein from wild game (see Dwyer 1974, 1982, 1983; Morren 1986). Much of this comes from birds. There are very few data on consumption of small birds, which are often consumed before researchers can register the event. This kind of hunting seems to have a significant effect on small bird populations around many human population centers (Mack, pers. obs.) but is unlikely to threaten small bird species that are fairly widespread. The birds most threatened directly by subsistence hunting and egg-taking are cassowaries (3 spp.), Goura pigeons (3 spp.) and other large terrestrial pigeons, hornbills (1 sp.), megapodes (9 spp.), and waterfowl (at least 1 sp.). Larger ground-nesting birds such as terns, waterfowl, and shorebirds probably suffer high nest predation wherever they nest near human population centers. Many of the larger birds are reasonably secure from traditional hunting with bow and arrow but they are vulnerable to shotguns and, in the case of cassowaries, snares. Thus technological innovations like shotguns and wire can have serious impacts when introduced. Megapodes are particularly vulnerable to depletion. Megapodes produce large and highly nutritious eggs and lay them in conspicuous mounds of rotting vegetation or in geothermally heated rookeries well known to local hunters. Such highly concentrated, easily located, and nutritious eggs are heavily exploited across the region. However, megapode eggs can be sustainably harvested with proper scientifically based management. Not coincidentally, one of the rarest birds in Papua is a megapode, Aepypodius bruijnii. This species is close to extinction, almost certainly due to past harvest practices that were unsustainable. It is reasonable to expect that other species and populations of megapodes will face the same fate if management practices do not begin soon.

agricultural conversion Agricultural conversion includes both large-scale industrial production and smallscale subsistence. Large areas of Papua have already been cleared for industrial agriculture or are planned to be cleared for plantations (Supriatna 1999). In the lowlands, conversion is primarily for oil palm plantations. Oil palm plantations are essentially sterile in terms of biodiversity, and they also pollute freshwater systems. Because they are monocultures, they have little biological or structural variability and thus make a poor substrate for bird populations. Sometimes some predatory birds, especially owls, do well in oil palm plantations due to the large numbers of rats supported by the palm nuts, but overall, oil palm plantations do not support substantial diversity.

................. 16157$

CH42

02-08-07 10:44:03

PS

PAGE 676

Birds of Papua / 677

industrial logging Large-scale logging presents a major threat to many birds in Papua. Impacts derive from the direct effects of cutting and extracting timber, and indirect effects such as elevated hunting and introduction of pests. The magnitude of direct effects will vary according to the intensity of the extraction and the practices used. At the most damaging extreme is clear-felling, in which the majority of trees are removed (e.g., for pulp wood). This type of logging is presently rare in Papua, but projections of future demand from Asia, particularly China, suggest the demand for New Guinea pulp could increase. At the other end of the spectrum is small-scale, environmentally friendly selective logging. Many conservationists are developing practices for reduced impact forestry and at the same time developing markets for certified lower-impact timber. There is still a great deal to learn about how to minimize the impact of logging, and we need data on the effects of different forestry practices. Nonetheless, some information is available, particularly from tropical forests in other parts of the world. Many bird species depend on hollow trees and snags for roosting and nesting sites (Gibbs et al. 1993). Logging practices that reduce damage to roost trees can help maintain populations of parrots and other cavity-nesting organisms in Australia (Brigham et al. 1998; Gibbons and Lindenmayer 2002). The impacts of moderate levels of timber removal on Papuan birds are difficult to measure, partly due to the inherent problems associated with censusing birds in rainforests (Driscoll 1984). However evidence suggest that while some bird species increase in regrowth and others decrease after logging, large tracts of second growth have depauperate avifaunas (Driscoll 1984). Because many birds of the New Guinea rainforests depend on either nectar or fruit as dietary staples, logging practices that reduce nectar- and fruit-bearing trees can have substantial effects on a large number of bird species. This impact will be more dramatic if keystone fruit or nectar resources are removed (van Schaik et al. 1993). Keystone resources are loosely defined as resources upon which a disproportionate number of species are strongly dependent, and whose removal would have large, cascading effects on the ecosystem (Paine 1995; Simberloff 1998). For example, figs or other fruits that are available during times of low overall fruit availability act as keystone resources (Kinnaird et al. 1999; Patel 1997; Peres 2000; Shanahan et al. 2001). Low impact logging should be designed to protect keystone resources so that key fruit and nectar resources are sufficient to maintain bird populations. Although data are sparse from Papua, it appears that keystone fruit resources would at least include some species of Ficus, Elmerrillia, and Calophyllum, but it is extremely difficult to clearly identify keystone fruit resources in New Guinea as some are important in different years or different seasons (Wright 2005). Regeneration after logging can be enhanced if bird populations are robust and able to maintain natural patterns of seed dispersal. Birds and other frugivores move seeds into disturbed areas and thus enhance forest regeneration (Gorchov

................. 16157$

CH42

02-08-07 10:44:03

PS

PAGE 677

678 / a n d rew m a c k & ja c k d um b ac her

et al. 1993; Holbrook et al. 2002; Wunderle 1997). Thus birds and other seed dispersing animals can assist revegetation of logged areas through natural processes that would be costly for humans to undertake. In the mountainous terrain of Papua, seed dispersing frugivores like cassowaries are essential in order to move seeds uphill. In the absence of dispersers, many plant populations will shift downhill over time merely by the agency of gravity moving seeds of each successive generation downhill (Mack et al. 1999).

commercial bird trade Few data exist on the numbers of birds exported from Papua for the live bird trade, but what information there is suggests a considerable trade in live birds from Papua into the markets of Southeast Asia (Nash 1992, 1994; Rumbiak 1984). The international live animal trade is a contributing factor in the conservation status of hundreds of threatened and endangered species on the IUCN Red List (IUCN 2002). Large numbers of birds in Southeast Asia move through the major bird markets of Jakarta, Singapore, and Hong Kong. For some, particularly parrots and lories, the live pet trade might present a significant threat. Closer monitoring of the markets and exportations from Papua is needed.

introduced exotic species New Guinea is fortunately free of many of the catastrophic introductions that have devastated many bird populations around the world. Introduced birds brought avian malarias to Hawaii that helped bring about the extinction of many native bird species (Warner 1968). On Guam, the inadvertent introduction of the Brown Tree Snake (Boiga irregularis) from the New Guinea region has caused the extinction of several endemic species (Fritts and Rodda 1998; Rodda et al. 1997). Throughout the Pacific, the introduction of the Polynesian rat to remote islands has caused a massive pulse of avian extinctions (Kirch et al. 1992; Owens and Bennett 2000). On Vanuatu, the introduced fire ant Wasmannia is wiping out birds that have altricial nestlings. These are just a few examples; many conservationists see the transportation and introduction of invasive exotic species as one of the biggest global threats to biodiversity. Although it may be possible to correct many environmental wrongs through rehabilitation, reintroduction, or restoration, it is almost impossible to eradicate an invasive species once it becomes established. There are few data for New Guinea on introduced and invasive species. Invasive plants and fungi of significant economic impact have received some attention. But the roles of introduced Rattus, cats, and other species likely to impact birds have received little attention. The numbers of invasive species that directly impact birds are probably lower than some areas, such as New Zealand or Australia, where many birds are threatened by stoats, cats, rats, and other predators. But conservationists, policy makers, and enforcement agencies should not be complacent. The figures for New Guinea are low, not because it is harder for invasive species to establish in New Guinea, but because there has been less economic activity and

................. 16157$

CH42

02-08-07 10:44:04

PS

PAGE 678

Birds of Papua / 679

commerce of the sort that brings invasive species. As commerce increases in Papua and there is more exchange with the outside world, more exotic species will land in Papua. Strict customs and quarantine regulations coupled with effective enforcement are needed. In Papua there is currently deep concern over the introduction of macaques, in the Jayapura area (Chapter 7.7). This species has had devastating effects on the native fauna and flora of other islands to which it has been introduced (e.g., the Mascarene Islands). The monkeys compete with other animals by consuming fruits and other resources and directly prey upon some animals, particularly nesting birds. Because New Guinea does not have any native placental predatory mammals, release of species such as cats or monkeys would pose a significant threat to the birds of Papua.

Literature Cited Abbott, L.D., E.A. Silver, R.S. Anderson, R. Smith, J.C. Ingle, S.A. Kling, D. Haid, E. Small, J. Galewsky, and W. Sliter. 1997. Measurement of tectonic surface uplift rate in a young collisional mountain belt. Nature 385: 501–507. Alvard, M.S., J.G. Robinson, K.H. Redford, and H. Kaplan. 1997. The sustainability of subsistence hunting in the neotropics. Conserv Biol 11: 977–982. Ashton, P.S. 2003. Floristic zonation of tree communities on wet tropical mountains revisited. Perspect Plant Ecol Evol Syst 6: 87–104. Baker, G.C., and S.H.M. Butchart. 2000. Threats to the maleo Macrocephalon maleo and recommendations for its conservation. Oryx 34: 255–261. Barker, F.K., G.F. Barrowclough, and J.G. Groth. 2002. A phylogenetic hypothesis for passerine birds: taxonomic and biogeographic implications of an analysis of nuclear DNA sequence data. Proc R Soc Lond B Biol Sci 269: 295–308. Barker, K.F., A. Cibois, P. Schikler, J. Feinstein, and J. Cracraft. 2004. Phylogeny and diversification of the largest avian radiation. Proc Natl Acad Sci U.S. 101: 11040– 11045. Bawa, K.S., and S.L. Krugman. 1991. Reproductive biology and genetics of tropical trees in relation to conservation and management. Pp. 119–136 in Gomez-Pompa, A., T.C. Whitmore, and M. Hadley (eds.) Rain Forest Regeneration and Management. Parthenon Publishing Co., Park Ridge, New Jersey. Beehler, B.M. 1982. Ecological structuring of forest bird communities in New Guinea. Pp. 837–860 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Beehler, B.M. 1985. Adaptive significance of monogamy in the Trumpet Manucode Manucodia keraudrenii (Aves: Paradisaeidae). Pp. 83–99 in Gowaty, P.A., and D.W. Mock (eds.) Avian Monogamy. American Ornithologists’ Union, Washington, D.C. Beehler, B.M. 1987. Birds of paradise and mating system theory—predictions and observations. Emu 87: 78–89. Beehler, B.M. 1988. Lek behavior of the Raggiana Bird of Paradise. Nat Geog Soc Res Rep 4: 343–358. Beehler, B.M. 1989. Patterns of frugivory and the evolution of birds of paradise. Proceedings of the XIX International Ornithological Congress: 816–826.

................. 16157$

CH42

02-08-07 10:44:05

PS

PAGE 679

680 / a n d rew m a c k & ja c k d um b ac her Beehler, B.M. 1994. Canopy-dwelling honeyeater aggressively defends terrestrial nectar source. Biotropica 26: 459–461. Beehler, B.M., W. Crill, B. Jefferies, and M. Jefferies. 1992. New Guinea Harpy Eagle attempts to capture a monitor lizard. Emu 92: 246–247. Beehler, B.M., and J.P. Dumbacher. 1996. More examples of fruiting trees visited predominantly by birds of paradise. Emu 96: 81–88. Beehler, B.M., and B.W. Finch. 1985. Species Checklist of the Birds of New Guinea. Royal Australasian Ornithologists Union, Victoria. Beehler, B.M., T.K. Pratt, and D.A. Zimmerman. 1986. Birds of New Guinea. Princeton University Press, Princeton. Beehler, B.M., J.B. Sengo, C. Filardi, and K. Merg. 1995. Documenting the lowland rainforest avifauna in Papua New Guinea—effects of patchy distributions, survey effort and methodology. Emu 95: 149–161. Bell, H.L. 1982a. A bird community of lowland rain forest in New Guinea. 2. Seasonality. Emu 82: 65–74. Bell, H.L. 1982b. A bird community of lowland rainforest in New Guinea. 1. Composition and density of the avifauna. Emu 82: 24–41. Bell, H.L. 1982c. A bird community of New Guinean lowland rainforest. 3. Vertical distribution of the avifauna. Emu 82: 143–162. Bell, H.L. 1983a. A bird community of lowland rainforest in New Guinea. 6. Foraging ecology and community structure of the avifauna. Emu 84: 142–158. Bell, H.L. 1983b. A bird community of lowland rainforest in New Guinea. 5. Mixedspecies flocks. Emu 82: 256–275. Benayas, J.M.R., and E. de la Montana. 2003. Identifying areas of high-value vertebrate diversity for strengthening conservation. Biol Conserv 114: 357–370. Bond, W.J. 1994. Do mutualisms matter? assessing the impact of pollinator and disperser disruption on plant extinction. Philos Trans R Soc Lond B Biol 344: 83–90. Brandl, R., A. Kristin, and B. Leisler. 1994. Dietary niche breadth in a local community of passerine birds, an analysis using phylogenetic contrasts. Oecologia 98: 109–116. Brigham, R.M., S.J.S. Debus, and F. Geiser. 1998. Cavity selection for roosting, and roosting ecology of forest-dwelling Australian Owlet-nightjars (Aegotheles cristatus). Aust J Ecol 23: 424–429. Brown, E.D., and M.J.G. Hopkins. 1995. A test of pollinator specificity and morphological convergence between nectarivorous birds and rainforest tree flowers in New Guinea. Oecologia 103: 89–100. Brown, E.D., and M.J.G. Hopkins. 1996. How New Guinea rainforest flower resources vary in time and space: implications for nectarivorous birds. Aust J Ecol 21: 363–378. Brussard, P.F. 1991. The role of ecology in biological conservation. Ecol Appl 1: 6–12. Bulmer, S. 1982. Human ecology and cultural variation in prehistoric New Guinea. Pp. 169–206 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Butchart, S.H.M., and G.C. Baker. 2000. Priority sites for conservation of maleos (Macrocephalon maleo) in central Sulawesi. Biol Conserv 94: 79–91. Cadee, G.C. 1985. Some data on seabird abundance in Indonesian waters, July–August 1984. Ardea 73: 183–188. Christidis, L., and R. Schodde. 1993. Sexual selection for novel partners, a mechanism for accelerated morphological evolution in the Birds-of-Paradise Paradisaeidae. Bull Brit Orn Club 113: 169–172. Coates, B.J. 1985. Birds of Papua New Guinea: Non-passerines. Dove, Alderley.

................. 16157$

CH42

02-08-07 10:44:06

PS

PAGE 680

Birds of Papua / 681 Coates, B.J. 1990. Birds of Papua New Guinea: Passerines. Dove, Alderley. Collins, B.G., and D.C. Paton. 1989. Consequences of differences in body mass, wing length and leg morphology for nectar-feeding birds. Aust J Ecol 14: 269–289. Connell, D.W., C.N. Fung, T.B. Minh, S. Tanabe, P.K.S. Lam, B.S.F. Wong, M.H.W. Lam, L.C. Wong, R.S.S. Wu, and B. Richardson. 2003. Risk to breeding success of fish-eating ardeids due to persistent organic contaminants in Hong Kong: evidence from organochlorine compounds in eggs. Water Res 37: 459–467. Connell, D.W., B.S.F. Wong, P.K.S. Lam, K.F. Poon, M.H.W. Lam, R.S. Wu, B.J. Richardson, and Y.F. Yen. 2002. Risk to breeding success of ardeids by contaminants in Hong Kong: evidence from trace metals in feathers. Ecotoxicology 11: 49–59. Cracraft, J., and J. Feinstein. 2000. What is not a bird of paradise? molecular and morphological evidence places Macgregoria in the Meliphagidae and Cnemophilinae near the base of the corvoid tree. Proc R Soc Lond B 267: 233–241. Croxall, J.P. 1977. Feeding behaviour and ecology of New Guinea rainforest insectivorous passerines. Ibis 119: 113–146. Davies, H.L., R.D. Winn, and P. KenGemar. 1996. Evolution of the Papuan Basin—a view from the orogen. Pp. 53–62 in Third PNG Petroleum Convention, Port Moresby, Papua New Guinea. De Luca-Abbott, S.B., B.S.F. Wong, D.B. Peakall, P.K.S. Lam, L. Young, M.H.W. Lam, and B.J. Richardson. 2001. Review of effects of water pollution on the breeding success of waterbirds, with particular reference to ardeids in Hong Kong. Ecotoxicology 10: 327–349. Diamond, J.M. 1966. Zoological classification system of a primitive people. Science 151: 1102–1104. Diamond, J.M. 1972. Avifauna of the Eastern Highlands of New Guinea. Nuttall Ornithological Club, Cambridge. Diamond, J.M. 1973. Distributional ecology of New Guinea birds. Science 179: 759–769. Diamond, J.M. 1978. Niche shifts and the rediscovery of interspecific competition. Am Sci 66: 322–331. Diamond, J.M. 1981. Birds of paradise and the theory of sexual selection. Nature 293: 257–258. Diamond, J.M. 1982a. Evolution of the bowerbirds’ bowers: animal origins of the aesthetic sense. Nature 297: 99–102. Diamond, J.M. 1982b. Rediscovery of the Yellow-fronted Gardner Bowerbird. Science 216: 431–434. Diamond, J.M. 1983. Melampitta gigantea: possible relation between feather structure and underground roosting habits. Condor 85: 89–91. Diamond, J.M. 1985. New distributional records and taxa from the outlying mountain ranges of New Guinea. Emu 85: 65–91. Diamond, J.M. 1986. Animal art: variation in bower decorating style among male bowerbirds Amblyornis inornatus. Proc Natl Acad Sci U.S. 89: 3042–3046. Diamond, J.M. 1987. Bower building and decoration by the bowerbird Amblyornis inornatus. Ethnology 74: 177–204. Diamond, J., S.L. Pimm, M.E. Gilpin, and M. Lecroy. 1989. Rapid evolution of character displacement in Myzomelid honeyeaters. Am Nat 134: 675–708. Dingle, H. 2004. The Australo-Papuan bird migration system: another consequence of Wallace’s Line. Emu 104: 95–108. Dow, D.B. 1977. A geological synthesis of Papua New Guinea. Bull Aust Bur Min Res 201. Dow, D.B., and R. Sukamto. 1984. Late Tertiary to Quaternary tectonics of Irian Jaya. Episodes 7: 3–9.

................. 16157$

CH42

02-08-07 10:44:07

PS

PAGE 681

682 / a n d rew m a c k & ja c k d um b ac her Driscoll, P.V. 1984. The effects of logging on bird populations in lowland New Guinea rainforest. Ph.D. diss., University of Queensland, Brisbane. Dumbacher, J.P., and R.C. Fleischer. 2001. Phylogenetic evidence for colour-pattern convergence in toxic pitohuis: Mu¨llerian mimicry in birds? Proc R Soc Lond B 268: 1971–1976. Dumbacher, J.P., T.K. Pratt, and R.C. Fleischer. 2003. Phylogeny of the owlet-nightjars (Aves: Aegothelidae) based on mitochondrial DNA sequence. Mol Phyl Evol 29: 540–549. Dwyer, P.D. 1974. The price of protein: five hundred hours of hunting in the New Guinea highlands. Oceania 44: 278–293. Dwyer, P.D. 1982. Prey switching: a case study from New Guinea. J Anim Ecol 51: 529–542. Dwyer, P.D. 1983. Etolo hunting performance and energetics. Human Ecol 11: 145–174. Eastwood, C. 1996. A trip to Irian Jaya. Muruk 8: 12–23. Emlen, S.T., and L.W. Oring. 1977. Ecology, sexual selection, and evolution of mating systems. Science 197: 215–223. Feld, S. 1990. Sound and Sentiment: Birds, Weeping, Poetics, and Song in Kaluli Expression. University of Pennsylvania Press, Philadelphia. Ficken, M.S. 2000. Call similarities among mixed species flock associates. Southwestern Nat 45: 154–158. Frith, C.B. 1971. Some undescribed nests and eggs of New-Guinea birds. Bull Brit Orn Club 91: 46–49. Frith, C.B. 1981. Displays of Count Raggi’s Bird of Paradise Paradisaea raggiana and congeneric species. Emu 81: 193–201. Frith, C.B., and B.M. Beehler. 1998. The Birds of Paradise: Paradisaeidae. Oxford University Press, Oxford. Frith, C.B., and D.W. Frith. 1993. The nesting biology of the Ribbon-tailed Astrapia Astrapia mayeri (Paradisaeidae). Emu 93: 12–22. Frith, C.B., and D.W. Frith. 1994. The nesting biology of Archbold’s Bowerbird Archboldia papuesis and a review of that of other bowerbirds (Ptilonorhynchidae). Ibis 136: 153–160. Frith, C.B., and D.W. Frith. 1996. Description of the unique Parotia lawesii and Paradisaea rudolphi hybrid bird of paradise (Paradisaeidae). Rec Aust Mus 48: 111–116. Frith, C.B., and D.W. Frith. 2004. The Bowerbirds: Ptilonorhynchidae. Oxford University Press, Oxford. Frith, D.W., and C.B. Frith. 1988. Courtship display and mating of the Surperb Bird of Paradise Lophorhina surperba. Emu 88: 183–188. Frith, H.J., F.H.J. Crome, and T.O. Wolfe. 1976. Food of the fruit-pigeons in New Guinea. Emu 76: 49–58. Fritts, T.H., and G.H. Rodda. 1998. The role of introduced species in the degradation of island ecosystems: a case history of Guam. Ann Rev Ecol Syst 29: 113–140. Fuller, E. 1995. The Lost Birds of Paradise. Swan Hill Press, Shrewsbury, England. Gibbons, O., and D.B. Lindenmayer. 2002. Tree Hollows and Wildlife Conservation in Australia. CSIRO Publishing, Collingwood, Victoria. Gibbs, D. 1994. Undescribed taxa and new records from the Fakfak Mountains, Irian Jaya. Bull Brit Orn Club 114: 4–12. Gibbs, J.P., M.L. Hunter, and S.M. Melvin. 1993. Snag availability and communities of cavity-nesting birds in tropical versus temperate forests. Biotropica 25: 236–241. Gorchov, D.L., F. Cornejo, C. Ascorra, and M. Jaramillo. 1993. The role of seed dispersal

................. 16157$

CH42

02-08-07 10:44:08

PS

PAGE 682

Birds of Papua / 683 in the natural regeneration of rain forest after strip-cutting in the Peruvian Amazon. Vegetatio 107/108: 339–349. Hall, R. 2001. Cenozoic reconstructions of SE Asia and the SW Pacific: changing patterns of land and sea. Pp. 35–56 in Metcalfe, I., J.M.B. Smith, M. Morwood, and I.D. Davidson (eds.) Faunal and Floral Migrations and Evolution in SE Asia-Australasia. Swets & Zeitlinger Publishers, Lisse, Netherlands. Hall, R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. J Asian Earth Sci 20: 353–431. Hamann, A., and E. Curio. 1999. Interactions among frugivores and fleshy fruit trees in a Philippine submontane rainforest. Conserv Biol 13: 766–773. Hardin, G. 1960. The competitive exclusion principle. Science 131: 1292–1297. Hartert, E., K. Paludan, L. Rothschild, and E. Stresemann. 1936. Die Vogel des WeylandGebriges und seines Vorlandes. Mitteilungen aus dem Zoolischen Museum 21: 11–186. Heads, M. 2001a. Birds of paradise (Paradisaeidae) and bowerbirds (Ptilonorhynchidae): Regional levels of biodiversity and terrane tectonics in New Guinea. J Zool (London) 255: 221–339. Heads, M. 2001b. Birds of paradise, biography and ecology in New Guinea: a review. J Biogeogr 28: 893–925. Heads, M. 2001c. Regional patterns of biodiversity in New Guinea plants. Bot J Linn Soc 136: 67–73. Heads, M. 2002. Birds of paradise, vicariance biogeography and terrane tectonics in New Guinea. J Biogeogr 29: 1–23. Healey, C.J. 1986. Men and birds in the Jimi Valley. The impact of man on birds of paradise in the Papua New Guinea highlands. Muruk 1: 34–71. Healey, C. 1993. Folk taxonomy and mythology of birds of paradise in the New Guinea highlands. Ethnology 32: 19–34. Heaney, W. 1982. The changing roles of birds of paradise plumes in bridewealth in the Wahgi Valley. Pp. 227–231 in Morauta, L., J. Pernetta, and W. Heaney (eds.) Traditional Conservation in Papua New Guinea, Implications for Today. Institute of Applied Social and Economic Research, Boroko. Hickey, J.J., and D.W. Anderson. 1968. Chlorinated hydrocarbons and eggshell changes in raptorial and fish-eating birds. Science 162: 271–273. Holbrook, K.M., T.B. Smith, and B.D. Hardesty. 2002. Implications of long-distance movements of frugivorous rain forest hornbills. Ecography 25: 745–749. Hunter, C.P., and P.D. Dwyer. 1997. The value of objects to Satin Bowerbirds Ptilonorhynchus violaceus. Emu 97: 200–206. Hurlbert, S.H. 1984. Pseudoreplication and the design of ecological field experiments. Ecol Monogr 54: 187–211. Hyndman, D.C., and J.I. Menzies. 1990. Rain forests of the Ok Tedi headwaters, New Guinea: an ecological analysis. J Biogeogr 17: 241–273. IUCN. 2002. IUCN Red List of Threatened Species. IUCN, Gland, Switzerland. Johns, R.J. 1976. A classification of the montane forests of Papua New Guinea. Science in New Guinea 4: 105–128. Johnson, A., R. Bino, and P. Igag. 2004. A preliminary evaluation of the sustainability of cassowary (Aves: Casuariidae) capture and trade in Papua New Guinea. Anim Conserv 7: 129–137. Jones, D.N., R.J. Dekker, and C.S. Roselaar. 1995. The Megapodes. Oxford University Press, Oxford.

................. 16157$

CH42

02-08-07 10:44:08

PS

PAGE 683

684 / a n d rew m a c k & ja c k d um b ac her Jullien, M., and J. Clobert. 2000. The survival value of flocking in neotropical birds: reality or fiction? Ecology 81: 3416–3430. Karasov, W.H., and D.J. Levey. 1990. Digestive trade-offs and adaptations of frugivorous passerine birds. Physiol Zool 63: 1248–1270. Kearns, C.A., D.W. Inouye, and N.M. Waser. 1998. Endangered mutualisms: the conservation of plant-pollinator interactions. Ann Rev Ecol Syst 29: 83–112. Kinnaird, M.F., T.G. O’Brien, and S. Suryadi. 1996. Population fluctuation in Sulawesi red-knobbed hornbills: tracking figs in space and time. Auk 113: 431–440. Kinnaird, M.F., T.G. O’Brien, and S. Suryadi. 1999. Importance of figs to Sulawesi’s imperiled wildlife. Trop Biodiv 6: 5–18. Kirch, P.V., J.R. Flenley, D.W. Steadman, F. Lamont, and S. Dawson. 1992. Ancient environmental degradation. Res Explor 8: 166–179. Kitayama, K. 1992a. An altitudinal transect study of the vegetation on Mount Kinabalu, Borneo. Vegetatio 102: 149–171. Kitayama, K. 1992b. Comparative vegetation analysis on the wet slopes of two tropical mountains: Mt Haleakala, Hawaii and Mt. Kinabalu, Borneo. Ph.D. diss., University of Hawaii. Kocher-Schmid, C. 1991. Of People and Plants: A Botanical Ethnography of Nokopo Village, Madang and Morobe Provinces, Papua New Guinea. Ethnologisches Seminar der Universita¨t und Museum fu¨r Vo¨lkerkunde, in Kommission bei Wepf und Co., Basel. Kocher-Schmid, C. 1993. Birds of Nokopo. Muruk 6: 1–15. Laurance, W.F., and S.G.W. Laurance. 1996. Responses of five arboreal marsupials to recent selective logging in tropical Australia. Biotropica 28: 310–322. Legge, S., S. Murphy, P. Igag, and A.L. Mack. 2004. Territoriality and density of an Australian migrant, the Buff-breasted Paradise Kingfisher, in the New Guinean nonbreeding grounds. Emu 104: 15–20. Lomolino, M.V. 2001. Elevation gradients of species-density: historical and prospective views. Global Ecol Biogeogr Lett 10: 3–13. Mack, A.L. 1993. The sizes of vertebrate-dispersed fruits: a neotropical-paleotropical comparison. Am Nat 142: 840–856. Mack, A.L., and L.E. Alonso (eds.). 2000. A biological assessment of the Wapoga River area of northwestern Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14. Conservation International, Washington, D.C. Mack, A.L., K. Ickes, J.H. Jessen, B. Kennedy, and J.R. Sinclair. 1999. Ecology of Aglaia mackiana (Meliaceae) seedlings in a New Guinea rain forest. Biotropica 31: 111–120. Mack, A.L., and P. West. 2005. Ten thousand tonnes of small animals: wildlife consumption in Papua New Guinea, a vital resource in need of management. Resource Management in Asia-Pacific Working Paper 61: 1–23. Mack, A.L., and D.D. Wright. 1996. Notes on the occurrence and feeding of birds at Crater Mountain Biological Research Station, Papua New Guinea. Emu 96: 89–101. Mack, A.L., and D.D. Wright. 1998. The Vulturine Parrot, Psittrichas fulgidus, a threatened New Guinea endemic: notes on its biology and conservation. Bird Conserv Intl 8: 185–194. Mack, A.L., and D.D. Wright. 2005. The frugivore community and the fruiting plant flora in a New Guinea rainforest: identifying keystone frugivores. Pp. 184–203 in Dew, L., and J.P. Boubli (eds.) Tropical Fruits and Frugivores: The Search for Strong Interactors. Springer, The Netherlands. Majnep, I.S., and R. Bulmer. 1977. Birds of My Kalam Country. Auckland University Press, Auckland.

................. 16157$

CH42

02-08-07 10:44:09

PS

PAGE 684

Birds of Papua / 685 Mayr, E. 1941. List of New Guinea Birds: A Systematic and Faunal List of the Birds of New Guinea and Adjacent Islands. American Museum of Natural History, New York. Mayr, E. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Massachusetts. Mayr, E., and J.M. Diamond. 1976. Birds on islands in the sky: origin of the montane avifauna of northern Melanesia. Proc Natl Acad Sci U.S. 73: 1765–1769. McGoldrick, J.M., and R. MacNally. 1998. Impact of flowering on bird community dynamics in some central Victorian eucalypt forests. Ecol Res 13: 125–139. McNab, B.K. 1988. Food habits and the basal rate of metabolism in birds. Oecologia 77: 343–349. McNab, B.K. 2005. Food habits and the evolution of energetics in birds of paradise (Paradisaeidae). J Comp Physiol B Biochem Syst Environ Physiol 175: 117–132. Mees, G.F. 1980. Supplementary notes on the avifauna of Misool. Zool Meded 55: 1–10. Mees, G.F. 1982. Birds from the lowlands of southern New Guinea (Merauke and Koembe). Zoologische Verhandelingen 191: 1–188. Melville, D. 1979. Ornithological notes on a visit to Irian Jaya. PNG Bird Society Newsletter 161: 3–22. Melville, D. 1980. Some observations on birds in Irian Jaya, New Guinea. Emu 80: 89–91. Mols, C.M.M., and M.E. Visser. 2002. Great tits can reduce caterpillar damage in apple orchards. J Appl Ecol 39: 888–899. Monterrubio, T., E. Enkerlin-Hoeflich, and R.B. Hamilton. 2002. Productivity and nesting success of Thick-billed Parrots. Condor 104: 788–794. Morren, G.E.B. 1986. The Miyanmin: Human Ecology of a Papua New Guinea Society. UMI Research Press, Ann Arbor. Nash, S.V. 1992. Parrot trade records for Irian Jaya, Indonesia, 1985–1990. TRAFFIC Bulletin 13: 42–45. Nash, S.V. 1994. Further parrot trade records for Irian Jaya, Indonesia. TRAFFIC Bulletin 14: 121–124. Novaro, A.J., K.H. Redford, and R.E. Bodmer. 2000. Effect of hunting in source-sink systems in the neotropics. Conserv Biol 14: 713–721. Nunn, G.B., and J. Cracraft. 1996. Phylogenetic relationships among the major lineages of the birds-of-paradise (Paradisaeidae) using mitochondrial DNA gene sequences. Mol Phyl Evol 5: 445–459. O’Brien, T.G., M.F. Kinnaird, E.S. Dierenfeld, N.L. Conklin-Brittain, R.W. Wrangham, and S.C. Silver. 1998. What’s so special about figs? Nature 392: 668–668. Ogilvie-Grant, W.R. 1915. Report on the birds collected by the British Ornithologists’ Union expedition and Wollaston expedition in Dutch New Guinea. Ibis Tenth Series, Jubilee Supplement no. 2: 1–336. Owens, I.P.F., and P.M. Bennett. 2000. Ecological basis of extinction risk in birds: habitat loss versus human persecution and introduced predators. Proc Natl Acad Sci U.S. 97: 12,144–12,148. Paijmans, K. 1970. An analysis of four tropical rain forest sites in New Guinea. J Ecol 58: 77–101. Paijmans, K. 1976. New Guinea Vegetation. National University Press, Canberra. Paine, R.T. 1995. A conversation on refining the concept of keystone species. Conserv Biol 9: 962–964. Patel, A. 1997. Phenological patterns of Ficus in relation to other forest trees in southern India. J Trop Ecol 13: 681–695. Paton, D.C., and B.G. Collins. 1989. Bills and tongues of nectar-feeding birds—a review

................. 16157$

CH42

02-08-07 10:44:09

PS

PAGE 685

686 / a n d rew m a c k & ja c k d um b ac her of morphology, function and performance, with intercontinental comparisons. Aust J Ecol 14: 473–506. Patterson, B.D., D.F. Stotz, S. Solari, J.W. Fitzpatrick, and V. Pacheco. 1998. Contrasting patterns of elevational zonation for birds and mammals in the Andes of southeastern Peru. J Biogeogr 25: 593–607. Pearson, D.L. 1977. A pantropical comparison of bird community structure on six lowland rain forest sites. Condor 79: 232–244. Peres, C.A. 2000. Identifying keystone plant resources in tropical forests: the case of gums from Parkia pods. J Trop Ecol 16: 287–317. Perfecto, I., J.H. Vandermeer, G.L. Bautista, G.I. Nunez, R. Greenberg, P. Bichier, and S. Langridge. 2004. Greater predation in shaded coffee farms: the role of resident neotropical birds. Ecology 85: 2677–2681. Petocz, R., M. Kirenius, and Y. de Fretes. 1983. Avifauna of the reserves in Irian Jaya. World Wildlife Fund Report. World Wildlife Fund, Bogor, Indonesia. Philpott, S.M., R. Greenberg, P. Bichier, and I. Perfecto. 2004. Impacts of major predators on tropical agroforest arthropods: comparisons within and across taxa. Oecologia 140: 140–149. Pigram, C.J., and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. BMR J Aust Geol Geophys 10: 193–211. Pigram, C.J., and P.A. Symonds. 1991. A review of the timing of the major tectonic events in the New Guinea orogen. J So East Asian Earth Sci 6: 307–318. Polhemus, D.A. 1996. Island arcs and their influence on Indo-Pacific biogeography. Pp. 51–56 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. Academic Publishing, Amsterdam. Pratt, T.K. 1982. Biogeography of birds in New Guinea. Pp. 815–836 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. W. Junk, The Hague. Pratt, T.K. 1984. Examples of tropical frugivores defending fruit-bearing plants. Condor 86: 123–129. Pratt, T.K. 2000. Evidence for a previously unrecognized species of owlet-nightjar. Auk 117: 1–11. Pratt, T.K., and E.W. Stiles. 1983. How long fruit-eating birds stay in the plants where they feed: implications for seed dispersal. Am Nat 122: 797–805. Pruett-Jones, S.G., and M.A. Pruett-Jones. 1988. The use of court objects by Lawes’ Parotia. Condor 90: 538–545. Ragusa-Netto, J. 2002. Vigilance towards raptors by nuclear species in bird mixed flocks in a Brazilian savannah. Studies on Neotropical Fauna and Environment 37: 219–226. Rahbek, C. 1997. The relationship among area, elevation, and regional species richness in neotropical birds. Am Nat 149: 875–902. Ramsey, M.W. 1988. Differences in pollinator effectiveness of birds and insects visiting Banksia menziesii (Proteaceae). Oecologia 76: 119–124. Ramsey, M.W. 1989. The seasonal abundance and foraging behavior of honeyeaters and their potential role in the pollination of Banksia menziesii. Aust J Ecol 14: 33–40. Rand, A.L., and E.T. Gilliard. 1967. Handbook of New Guinea Birds. Weidenfeld and Nicolson, London. Richards, S.J., and S. Suryadi (eds.). 2002. A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. Conservation International, Washington, D.C. Richardson, A.J., and I.R. Taylor. 2003. Are rice fields in southeastern Australia an

................. 16157$

CH42

02-08-07 10:44:10

PS

PAGE 686

Birds of Papua / 687 adequate substitute for natural wetlands as foraging areas for egrets? Waterbirds 26: 353–363. Ridgely, R.S., D. Agro, and L. Joseph. 2005. Birds of Iwokrama forest. Proc Acad Nat Sci Phil 154: 109–121. Robinson, J.G., K.H. Redford, and E.L. Bennett. 1999. Wildlife harvest in logged tropical forests. Science 284: 595–596. Rodda, G.H., T.H. Fritts, and D. Chiszar. 1997. The disappearance of Guam’s wildlife. Bioscience 47: 565–574. Rumbiak, A.M. 1984. Observations on the trade in birds of paradise in Bomakia District of Kouh, Region of Merauke. Sanchez-Cordero, V. 2001. Elevation gradients of diversity for rodents and bats in Oaxaca, Mexico. Global Ecol Biogeogr Lett 10: 63–76. Sanderson, E.W., K.H. Redford, A. Vedder, P.B. Coppolillo, and S.E. Wardrecord. 2002. A conceptual model for conservation planning based on landscape species requirements. Lands Urban Plan 58: 41–56. Seixas, G.H.F., and G.D. Mourao. 2002. Nesting success and hatching survival of the Bluefronted Amazon (Amazona aestiva) in the Pantanal of Mato Grosso do Sul, Brazil. J Field Orn 73: 399–409. Shanahan, M., S. So, S.G. Compton, and R. Corlett. 2001. Fig-eating by vertebrate frugivores: a global review. Biol Rev 76: 529–572. Sibley, C.G., and B.L.J. Monroe. 1990. Phylogeny and Classification of Birds of the World. Yale University Press, New Haven. Simberloff, D. 1998. Flagships, umbrellas, and keystones: is single-species management passe in the landscape era? Biol Conserv 83: 247–257. Sinclair, J.R. 2000. The behaviour, ecology and conservation of three species of megapode in Papua New Guinea. M.Sc. thesis, University of Otago, Otago. Sinclair, J.R. 2001. Temperature regulation in mounds of three sympatric species of megapode (Aves: Megapodiidae) in Papua New Guinea: testing the ‘‘Seymour Model.’’ Aust J Zool 49: 675–694. Sinclair, J.R., T.G. O’Brien, and M.F. Kinnaird. 2002. The selection of incubation sites by the Philippine Megapode, Megapodius cumingii, in North Sulawesi, Indonesia. Emu 102: 151–158. Stattersfield, A.J., M.J. Crosby, A.J. Long, and D.C. Wege. 1998. Endemic Bird Areas of the World: Priorities for Biodiversity Conservation. BirdLife International, Cambridge. Steadman, D.W. 1991. The identity and taxonomic status of Megapodius stairi and M. burnabyi (Aves, Megapodiidae). Proc Biol Soc Wash 104: 870–877. Steadman, D.W., G.K. Pregill, and D.V. Burley. 2002. Rapid prehistoric extinction of iguanas and birds in Polynesia. Proc Natl Acad Sci U.S. 99: 3673–3677. Steadman, D.W., J.P. White, and J. Allen. 1999. Prehistoric birds from New Ireland, Papua New Guinea: extinctions on a large Melanesian island. Proc Natl Acad Sci U.S. 96: 2563–2568. Styring, A.R., and K. Ickes. 2001. Woodpecker participation in mixed species flocks in peninsular Malaysia. Wilson Bull 113: 342–345. Supriatna, J. 1999. The Irian Jaya Biodiversity Conservation Priority-setting Workshop: Final Report. Conservation International, Washington, D.C. Swadling, P. 1995. Plumes from Paradise: Trade Cycles in Outer Southeast Asia and Their Impact on New Guinea and Nearby Islands until 1920. Robert Brown and Assoc., Coorparoo. Terborgh, J.W., and J.M. Diamond. 1970. Niche overlap in feeding assemblages of New Guinea birds. Wilson Bull 89: 29–52.

................. 16157$

CH42

02-08-07 10:44:10

PS

PAGE 687

688 / a n d rew m a c k & ja c k d um b ac her Thiollay, J.M. 1999. Frequency of mixed species flocking in tropical forest birds and correlates of predation risk: an intertropical comparison. J Avian Biol 30: 282–294. Tremblay, A., P. Mineau, and R.K. Stewart. 2001. Effects of bird predation on some pest insect populations in corn. Agric Ecosyst Environ 83: 143–152. van Bael, S.A., and J.D. Brawn. 2005. The direct and indirect effects of insectivory by birds in two contrasting neotropical forests. Oecologia 143: 106–116. van Bael, S.A., J.D. Brawn, and S.K. Robinson. 2003. Birds defend trees from herbivores in a neotropical forest canopy. Proc Natl Acad Sci U.S. 100: 8304–8307. van Balen, S. 1998. Tropical forest raptors in Indonesia: recent information on distribution, status, and conservation. J Raptor Res 32: 56–63. van Schaik, C.P., J.W. Terborgh, and S.J. Wright. 1993. The phenology of tropical forests—adaptive significance and consequences for primary consumers. Ann Rev Ecol Syst 24: 353–377. Vuilleumier, F. 1967. Mixed species flocks in Patagonian forests with remarks on interspecies flock formation. Condor 69: 400–404. Wallace, A.R. 1869. The Malay Archipelago and the Land of the Orang-utan and the Bird of Paradise. Richard Clay and Sons, Suffolk. Wallace, A.R. 1876. The Geographical Distribution of Animals: With a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth’s Surface. Macmillan, London. Warner, R.E. 1968. The role of introduced diseases in the extinction of the endemic Hawaiian avifauna. Condor 70: 101–120. Waser, N.M., L. Chittka, M.V. Price, N.M. Williams, and J. Ollerton. 1996. Generalization in pollination systems, and why it matters. Ecology 77: 1043–1060. Watson, M., and S. Asoyama. 2001. Dispersion, habitat use, hunting behavior, vocalizations, and conservation status of the New Guinea Harpy Eagle (Harpyopsis novaeguineae). J Raptor Res 35: 235–239. Westoby, M. 1988. Comparing Australian ecosystems to those elsewhere. Bioscience 38: 549–556. Wollaston, A.F.R. 1912. Pygmies & Papuans: The Stone Age To-day in Dutch New Guinea. Smith, Elder & Co., London. Wooller, F.L.S., and K.C. Richardson. 1988. Morphological relationships of passerine birds from Australia and New Guinea in relation to their diets. Zool J Linn Soc 94: 193–201. Wooller, R.D., K.C. Richardson, and D.R. Wells. 1990. Allometric relationships of the gastrointestinal tracts of insectivorous passerine birds from Malaysia, New-Guinea and Australia. Aust J Zool 38: 665–671. Wright, D.D. 2005. Diet, keystone resources, and altitudinal movement of dwarf cassowaries in relation to fruiting phenology in a Papua New Guinean rainforest. Pp. 204–235 in Dew, J.L., and J. P. Boubli (eds.) Tropical Fruits and Frugivores: The Search for Strong Interactors. Springer, New York. Wunderle, J.M. 1997. The role of animal seed dispersal in accelerating native forest regeneration on degraded tropical lands. Forest Ecol Mangt 99: 223–235.

................. 16157$

CH42

02-08-07 10:44:11

PS

PAGE 688

4.10. A Taxonomic and Geographic Overview of the Mammals of Papua kristofer m. helgen h e co m p o si t i o n of the biota of the Australo-Papuan region is strikingly unique compared to other continental areas worldwide, a point well illustrated by the region’s native mammals. Nowhere else do all three fundamental clades of living mammals—monotremes, marsupials, and placentals—co-exist. Nowhere else are the marsupials so remarkably diverse in both morphology and ecology, and in no other continental fauna are the placental orders so poorly represented. As the explorer, collector, and biogeographer Alfred Russel Wallace famously observed, a striking faunal divide lies in the center of the Malay Archipelago, separating two highly distinctive biotic worlds: the disparate faunas of the Laurasian continent of Asia and the Gondwanan Australia (e.g., Wallace 1869). The western half of the archipelago, the Sunda Shelf—an extension of continental Asia—hosts a rich higher-taxonomic complement of placental mammals (12 eutherian orders are represented in the modern faunas of Sumatra, Java, and Borneo). In the east, the Moluccas, New Guinea, and Australia instead support marsupials, monotremes, and a much smaller complement of placental mammals—only bats and murine rodents. In between, on the island of Sulawesi, these great regional faunas meld together, with small frugivorous and large folivorous possums sharing the forests with endemic primates, squirrels, shrews, and civets. In terms of biodiversity, the jewel of the Australo-Papuan region is New Guinea, the world’s largest tropical island. The modern mammal fauna of New Guinea includes, for example, the world’s largest living monotreme, more arboreal marsupials than anywhere else, one of the richest regional complements of Old World fruit bats globally, and a spectacular ecomorphological radiation of murine rodents, ranging in size and habits from tiny terrestrial shrew-like ‘‘moss mice’’ and amphibious rats to the world’s largest-bodied extant murine (the montane genus Mallomys). Despite its global uniqueness, the mammal fauna of New Guinea remains very incompletely cataloged today, and the mammals of western New Guinea (Papua) are on the whole less well known than mammals in adjacent Papua New Guinea (PNG; Flannery 1995a). Heroic efforts to understand the regional mammal fauna of Papua were made by important collectors and taxonomists (e.g., Wollaston 1914; Ru¨mmler 1938; Brass 1941; Tate 1947, 1948a,b, 1951; van Steenis-Kruseman 1950; Laurie and Hill 1954; Ziegler 1977, 1982; Taylor et al. 1982; Hill 1990; Menzies 1991, 1993; Flannery 1990, 1995a,b; Figures 4.10.1, 2), particularly from the 1910s to 1950s, a time of seminal biological exploration. Nevertheless, Papua has probably received less intensive study to date than the mammal fauna of almost any other tropical area of comparative size worldwide.

T

Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

689

................. 16157$

CH43

02-08-07 10:44:21

PS

PAGE 689

690 / kristofer m . helgen

Figure 4.10.1. Cumulative taxonomic history of the mammal fauna of Papua. This cumulative graph marks the original scientific description of each currently recognized mammal species that occurs in Papua. Note that many species that occur in Papua were originally discovered in Papua New Guinea, Australia, or elsewhere. The earliest described species that occurs in Papua was named in 1766 (Phalanger orientalis, discovered in the Moluccas); the first species actually discovered in present-day Papua (Spilocuscus papuensis, in 1822) was not named until the early 19th century (Figure 4.10.2). The half-way point to the current total was reached in about 1905. Efforts to understand mammalian biodiversity in Papua are once again accelerating. In the last decade, many unknown or overlooked mammal species from western New Guinea have been described as new to science or elevated from taxonomic synonymy based on studies of both recently collected and historical museum specimens (Figures 4.10.1, 2; Groves and Flannery 1994; Flannery 1995a, b, 1999; Flannery et al., 1995, 1996; Menzies 1996a; Flannery and Groves 1998; Aplin et al., 1999; Bergmans 2001; Helgen and Flannery 2004a,b; Musser and Carleton 2005; Woolley 2005). Yet mammal assemblages from relatively large portions of Papua, from many discrete montane areas (e.g., the Foja, Van Rees, Tamrau, Fakfak, Kumawa, and Wandammen [or Wondiwoi] ranges), many lowland regions (e.g., the western Trans-Fly, southern Vogelkop, Bird’s Neck lowlands), and on large islands (e.g., Waigeo and Misool), remain largely or entirely

................. 16157$

CH43

02-08-07 10:45:13

PS

PAGE 690

Taxonomic and Geographic Overview of the Mammals / 691

Figure 4.10.2. Cumulative history of systematic mammalogy in Papua. Lines indicate cumulative numbers of currently recognized mammal species that were first discovered in Papua (solid line  marsupials and monotremes; dotted line  rodents; dashed line  bats). The curves are reasonable proxies for historical attention paid to taxa of the mammal fauna of Papua over the past two centuries; curves end when the most recent species was described. The shape of the curve for rodent descriptions is strikingly similar to that for marsupials and monotremes, though delayed by about 85 years, suggesting accelerating attention over time, especially from 1930 to 1950 (when several important regional rodent systematists were active) and again in recent decades. Insectivorous bats of Papua have received comparatively little attention and are overdue for serious study. unstudied. Many mammal species probably await discovery and classification in these and other areas of western New Guinea. Comparison of the mammal fauna of western New Guinea to that of other areas of the world is not the intent of this review. However, a few comments will serve to illuminate the notable diversity and endemism that characterize the native mammal fauna of Papua in global context (cf. Chapter 4.1). Biological inventory and taxonomic efforts to date have recorded a total of 191 native mammal species in Papua, including 40 endemic species (Tables 4.10.1–4). Though undoubtedly far from complete (e.g., Table 4.10.5), this total is equal to or greater than the number of native mammals recorded from other tropical and subtropical insular areas worldwide, including Madagascar (Goodman and Benstead 2005), the Phil-

................. 16157$

CH43

02-08-07 10:46:01

PS

PAGE 691

................. 16157$

CH43

02-08-07 10:46:01

PS

Zaglossus attenboroughi Flannery and Groves 1998

Zaglossus bruijnii (Peters and Doria 1876)

Zaglossus bartoni (Thomas 1907)

3

4

Dasyurus albopunctatus Schlegel 1880

Dasyurus spartacus van Dyck 1988

Murexia habbema (Tate and Archbold 1941)

5

6

7

Family Dasyuridae (14 species)

Tachyglossus aculeatus (Shaw 1792)

2

Number (see Chapter Appendix)

1

Family Tachglossidae (4 species)

Taxa 0–1,700

Endemic?1 Elevation2 (m) 2,700–2,800

⬍100

0–4,000

0–4,150

E 0–2,500

E 1,600

x

Northern flat lowlands3 Central Cordillera, northern4 x

North coast ranges5 ?b

x

Yapen Island* Numfoor Island• Biak, Supiori, Padaido islands• x

Vogelkop Peninsula lowlands x

x

Salawati Island* ?

x

Batanta Island• ?

?

Waigeo Island• Misool Island* ?

Vogelkop Peninsula mountains6 x

x

?

x

?

x

(x)

?

x

Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8

Table 4.10.1. Elevational and distributional ranges of the monotremes and marsupials of Papua (64 recorded species)

Central Cordillera, southern9 ?

x

? x

x

?d

x

x

Weyland Range10 Snow Mountains region11 Star Mountains10 ?

692 / kristofer m . helgen

PAGE 692

................. 16157$

CH43

Phascolosorex dorsalis (Peters and Doria 1876)

Phascolosorex brevicaudata (Rothschild and Dollman 1932)

Sminthopsis virginiae (Tarragon 1847)

16

17

18

Echymipera clara (Stein 1932)

Echymipera kalubu (Lesson 1828)

Echymipera rufescens (Peters and Doria 1875)

Isoodon macrourus (Gould 1842)

Microperoryctes aplini Helgen and Flannery 2004

19

20

21

22

23

Family Peramelidae (8 species, 9 counting unidentified)

Phascolosorex doriae (Thomas 1886)

Myoictis wallacei Gray 1858

13

15

Myoictis melas (Mu¨ller 1840)

12

Neophascogale lorentzii (Jentink 1911)

Murexia wilhelmina (Tate 1947)

14

Murexia naso (Jentink 1911)

11

Murexia melanurus (Thomas 1899)

9

10

Murexia longicaudata (Schlegel 1866)

8

02-08-07 10:46:02

PS

E 1,900–2,200

0–1,200

0–2,100

0–1,200

0–1,700

0–500

1,500–3,600

E 1,600–2,500

E 900–2,000

1,200–3,900

0–900

0–1,800

1,000–2,900

1,400–2,800

0–2,000

0–1,800

x

x

x

x

x

x

x

x

x

x

x

x

?

x

x

x

?b

?b

x

x ? x

x

x

?

x

x

x

x

x

?

?

x

x

?

?

? x

(x) x x

(x) x ?

x

x

x

x

x

x

x

x

?

?

x

?

?

?

?

?

?

?

x

?

?

?

x

x

x

x

x

?a

?

x

x

x

?

x

?d

?d

?d

x

?

?d

?d

x

x

(continued)

? x

x x

x x

x x

x x

x x

? x

x x

x x

x ?

Taxonomic and Geographic Overview of the Mammals / 693

PAGE 693

................. 16157$

CH43

02-08-07 10:46:02

PS

Microperoryctes murina Stein 1932

Microperoryctes undescribed species

Peroryctes raffrayana (MilneEdwards 1878)

25

26

27

Dendrolagus inustus Mu¨ller 1840

Dendrolagus mayri Rothschild and Dollman 1933

Dendrolagus mbaiso Flannery, Boeadi, and Szalay 1995

28

29

30

Family Macropodidae (13 species)

Microperoryctes longicauda (Peters and Doria 1876)

Taxa

24

Number (see Chapter Appendix)

Table 4.10.1. (Continued)

Endemic?1 Elevation2 (m) E 3,200–4,200

E 1,600

0–1,400

0–4,000

E 2,200–4,000

E 2,500

1,000–3,600

x

x

Northern flat lowlands3 Central Cordillera, northern4 x

North coast ranges5 x

x

Yapen Island* Numfoor Island• Biak, Supiori, Padaido islands• x

x

Vogelkop Peninsula lowlands x

x

(Q)

Batanta Island•

Salawati Island* (x) ?

Vogelkop Peninsula mountains6 Waigeo Island• Misool Island* x (x) x

x

x

x

?

Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8 x

?

?

?

?

Central Cordillera, southern9 x

x

x

x x

x x

x

Weyland Range10 Snow Mountains region11 Star Mountains10 x x

694 / kristofer m . helgen

PAGE 694

................. 16157$

Dendrolagus stellarum Flannery and Seri 1990

Dendrolagus ursinus (Temminck 1836)

Dorcopsis hageni Heller 1897

Dorcopsis luctuosa (D’Albertis 1874)

Dorcopsis muelleri (Schlegel 1866)

Dorcopsulus vanheurni (Thomas 1922)

Macropus agilis (Gould 1842)

Thylogale browni (Ramsay 1877)

Thylogale brunii (Schreber 1778)

32

33

34

35

36

37

38

39

40

CH43

02-08-07 10:46:03

Phalanger vestitus (Milne-Edwards 1877)

Spilocuscus maculatus (E. Geoffroy 1803)

47

Phalanger orientalis (Pallas 1766)

44

Phalanger sericeus Thomas 1907

Phalanger mimicus Thomas 1922

43

46

Phalanger gymnotis (Peters and Doria 1875)

42

45

Phalanger carmelitae Thomas 1898

41

Family Phalangeridae (10 species)

Dendrolagus pulcherrimus Flannery 1993

31

PS

0–1,400

1,200–2,200

1,500–3,900

0–1,500

0–800

0–2,700

1,350–3,800

⬍200

0–2,300

⬍500

800–3,200

E ⬍600

0–500

0–800

E 0–2,500

2,600–4,000

700–1,100

x

x

x

x

x

x

x

x

x

x

?

?b

?c ?

x

?

?c

(x)

x x

x x x

x

x

x

x

x

(Q?)

x

x

x

x

x

x

x

?

x

?

x

x x

x

? x

x

x

x

x

(Q)

x

(Q)

x

x

?

?

x

?

?

?

?

?

x

?

x

x

?a

x

x

x

x

x

x

x

?

?d

?d

?d

?d

?d

?d

x

(continued)

x ?

x

? x

x

x

x

x

Taxonomic and Geographic Overview of the Mammals / 695

PAGE 695

................. 16157$

CH43

02-08-07 10:46:03

PS

PAGE 696

52 Cercartetus caudatus (MilneEdwards 1877)

Family Burramyidae (1 species)

Distoechurus pennatus (Peters 1874)

Spilocuscus wilsoni Helgen and Flannery 2004

50 Family Acrobatidae (1 species)

Spilocuscus rufoniger (Zimara 1937)

49

51

Spilocuscus papuensis (Desmarest 1822)

48

Taxa

Table 4.10.1. (Continued)

Number (see Chapter Appendix)

1,000–3,700

0–1,900

0–1,200

E ⬍1,000

E ⬍1,000

Endemic?1 Elevation2 (m)

x

x

Northern flat lowlands3 Central Cordillera, northern4

x

x

x

?b

North coast ranges5

?

?

Yapen Island* Numfoor Island• Biak, Supiori, Padaido islands•

x

x

x

Vogelkop Peninsula lowlands

?

Salawati Island* ?

Batanta Island•

x

Waigeo Island• Misool Island*

x

x

x

Vogelkop Peninsula mountains6

?

x

?

?

?

Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8

?a ? x

?e x ?

Central Cordillera, southern9

x

x

Weyland Range10 Snow Mountains region11 Star Mountains10 696 / kristofer m . helgen

Dactylopsila trivirgata Gray 1858

55

................. 16157$

CH43

Pseudochirulus caroli (Thomas 1921)

Pseudochirulus forbesi (Thomas 1887)

Pseudochirulus mayeri (Rothschild and Dollman 1932)

Pseudochirulus schlegeli (Jentink 1884)

63

64

65

Pseuochirops cupreus (Thomas 1897)

60

62

Pseudochirops coronatus (Thomas 1897)

59

Pseudochirulus canescens (Waterhouse 1846)

Pseudochirops corinnae (Thomas 1897)

58

61

Pseudochirops albertisii (Peters 1874)

57

Family Pseudocheiridae (9 species)

Petaurus breviceps Waterhouse 1838

Dactylopsila megalura Rothschild and Dollman 1932

54

56

Dactylonax palpator (MilneEdwards 1888)

53

Family Petauridae (4 species)

02-08-07 10:46:03

PS

E 750–1,900

1,200–4,200

450–3,800

E 0–ⱕ2,200

0–1,700

1,350–4,000

E 1,900–2,500

900–2,900

300–2,500

3,000

0–2,400

1,000–2,300

850–3,000

x

x

x

x

x

x

?b

x

x

x

x

x

x

x x x

x

x

x

x

x

x

? ?

?

?

? x

x ?

x

?

x

x

?

x

x

?

x

x

?

?

?

?

?

?

x

?

?

x

x

x

x

?e

x

x

x x

x x

x x

x

x x

?e x x

x x

x x

x

x

?d

x

?d

x

?d

?d

Taxonomic and Geographic Overview of the Mammals / 697

PAGE 697

................. 16157$

CH43

02-08-07 10:46:03

PS

d e

(Q) a b c

Legend blank x (x) ?

not recorded and probably absent from this area recorded in this area by a vouchered museum specimen recorded in this area by a vouchered photograph or accredited sighting not recorded but possibly present in this area (e.g., unconfirmed sight records, local information, presence inferred from wide distribution and appropriate habitat, recorded in Quaternary record and probably still extant, or present in better-surveyed adjacent parts of PNG) present in the late Quaternary fossil record but probably now extinct in this area present in the Fly River drainage of PNG and likely to extend to the TransFly area of Papua present above 500 m in the North Coast Ranges of PNG and likely to extend to the North Coast Ranges of Papua present below 1,000 m along the northern margin of the Central Dividing Ranges of PNG and likely to extend to the northern margin of the Central Dividing Ranges in Papua present above 1,000 m in the Star Mountains and/or Telefomin area of PNG and likely to extend to the Star Mountains of Papua present below 1,000 m along the southern margin of the Central Dividing Ranges of PNG and likely to extend to the southern margin of the Central Dividing Ranges in Papua

Note: Taxonomy follows Appendix 4.10.1, which should be consulted for numbered notes on taxonomic issues. Additional species can be expected in Papua (see Table 4.10.5). Unless indicated, lowlands areas are below 500 m. 1 Endemic?: E  species is endemic; blank  species is nonendemic. 2 Elevational ranges are drawn from vouchered specimens with explicit, associated altitudinal data in world museums (collected from all of mainland New Guinea and immediately adjacent islands, not just from Papua). ‘‘⬍ X’’ denotes that the upper altitudinal bound is not well established, but is likely to lie at or near X meters. 3 Northern flat lowlands of Papua, north of the Central Cordillera 4 Northern margin of the Central Cordillera, below 1,000 m 5 North coast ranges include the Cyclops and Foja Mts, above 500 m 6 Vogelkop Peninsula mountains include Arfak and Tamrau Mts, above 500 m 7 Bird’s Neck mountains include Fakfak and Wondiwoi Mts, above 500 m 8 Southern lowlands of eastern Papua include drainages of Digul and Fly rivers and the Trans-Fly ecoregion 9 Southern margin of the Central Cordillera, extending to the drainages of the Mimika, Setakwa, Utakwa, and Lorentz rivers, below 1,000 m 10 above 1,000 m 11 Greater Snow Mountains region, sensu Beehler et al. 1986 * indicates land-bridge islands or near-oceanic islands

Table 4.10.1. (Continued)

698 / kristofer m . helgen

PAGE 698

................. 16157$

Number (see Chapter Appendix)

Taxa

CH43

02-08-07 10:46:03

PS

ⱕ 2000 E 1750–1765

6 Hydromys chrysogaster E. Geoffroy, 1804

7 Hydromys hussoni Musser & Piik, 1982

10 Lorentzimys nouhuysii Jentink, 1911

9 Leptomys undescribed species ⱕ 2800

E 1000

1400–4000

1200–3500

5 Crossomys moncktoni Thomas, 1907

8 Hyomys dammermani Stein, 1933

1900–4100

E 2800–3200

E 2800–3600

1200–2900

Endemic?1 Elevation2 (m)

4 Coccymys ruemmleri (Tate & Archbold, 1941)

3 Coccymys albidens (Tate, 1951)

2 Baiyankamys habbema (Tate & Archbold, 1941)

1 Anisomys imitator Thomas, 1904

?

x

x

Northern flat lowlands3 Central Cordillera, northern4 x

North coast ranges5 ?b

?

Biak, Supiori, Padaido islands•

Yapen Island* Numfoor Island• x (x) x

Vogelkop Peninsula lowlands ?

Salawati Island*

x

x

(x) ? x ? ?

? x

?

?

?

?

x

x

Batanta Island• Waigeo Island• Misool Island* Vogelkop Peninsula mountains6 Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8

Table 4.10.2. Elevational and distributional ranges of the rodents of Papua, family Muridae (61 recorded species)

Central Cordillera, southern9 x

x

Weyland Range10 Snow Mountains region11 x

?d

?d

?d

?

?d

x

Star Mountains10 (continued)

x

x x

?

? x

? x

x

x

x x

Taxonomic and Geographic Overview of the Mammals / 699

PAGE 699

................. 16157$

CH43

1100–2900

13 Mallomys aroaensis (De Vis, 1907)

02-08-07 10:46:04

PS

x

ⱕ 1500

19 Mammelomys rattoides (Thomas, 1922)

x

x

1000–3200 x

Northern flat lowlands3 Central Cordillera, northern4 ?c

North coast ranges5

18 Mammelomys lanosus (Thomas, 1922)

E 1600–2500

1200–3700

16 Mallomys rothschildi Thomas, 1898

17 Mallomys new species

1500–3900

15 Mallomys istapantap Flannery, Aplin, & Groves, 1989

E 3500–4400

700–1900

14 Mallomys gunung Flannery, Aplin, & Groves, 1989

E 1400–1800

12 Macruromys major Ru¨mmler, 1935

Taxa

Endemic?1 Elevation2 (m)

11 Macruromys elegans Stein, 1933

Number (see Chapter Appendix)

Table 4.10.2. (Continued)

Yapen Island* Numfoor Island• x

x

?

?

?

Batanta Island• Waigeo Island• Misool Island* Vogelkop Peninsula mountains6 Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8 ?

Weyland Range10 Snow Mountains region11 x

x

x x

x

x

x

? x

x

Star Mountains10 ?d

?d

?d

?

?d

?d

700 / kristofer m . helgen

PAGE 700

Central Cordillera, southern9

Salawati Island*

Vogelkop Peninsula lowlands

Biak, Supiori, Padaido islands•

?

?

?

1200–1700 1600–2800

28 Paraleptomys rufilatus Osgood, 1945

29 Paraleptomys wilhelmina Tate & Archbold, 1941

................. 16157$

?

CH43

02-08-07 10:46:04

PS

ⱕ 700 E ⱕ 600

38 Pogonomelomys bruijnii (Peters & Doria, 1876)

36 Paramelomys steini (Ru¨mmler, 1935)

37 Pogonomelomys brassi Tate & Archbold, 1941

E 2000–2600

35 Paramelomys rubex (Thomas, 1922)

x x

x

?

x

x

x ? ? ?

?

?

? ?

x

?

?e

? x

ⱕ 1500 900–3000

34 Paramelomys platyops (Thomas, 1906)

x

x

x

x

ⱕ 1500

x

x

x x

x

? x

?d

x

?d

?

?d

?d

x

?d

(continued)

x

x x

x

x x

x

x x

?e ? ?

33 Paramelomys naso (Thomas, 1911)

?

?

?

ⱕ 1400 x

x ?

1200–2500

x

x

? x ? ?

32 Paramelomys moncktoni (Thomas, 1904) x ?

x ?

31 Paramelomys mollis (Thomas, 1913)

x

x ?

ⱕ 900 ?

x

x ?

30 Paramelomys lorentzii (Jentink, 1908)

x

?c ?b

x

ⱕ 2800

x ?b

27 Parahydromys asper (Thomas, 1906)

x x

26 Microhydromys richardsoni Tate & Archbold, 1941

x

ⱕ 2400 400–1500

25 Melomys rufescens (Alston, 1877)

?

?a x

x

?

? x

?

ⱕ 200

?

24 Melomys muscalis Thomas, 1913

? ?

x x

ⱕ 1400 ⱕ 1500

23 Melomys lutillus (Thomas, 1913)

E 1600–2800

1400–2800

22 Melomys leucogaster (Jentink, 1908)

21 Melomys frigicola Tate, 1951

20 Mayermys ellermani Laurie & Hill, 1954

Taxonomic and Geographic Overview of the Mammals / 701

PAGE 701

................. 16157$

CH43

02-08-07 10:46:04

x

ⱕ 2000

41 Pogonomys macrourus (Milne-Edwards, 1877)

PS

2200–4100

E 2000

47 Rattus arrogans (Thomas, 1922)

E 2800

46 Rattus arfakiensis (Ru¨mmler, 1935)

2300–3600

E 2200–3200

45 Pseudohydromys undescribed species

44 Pseudohydromys occidentalis Tate, 1951

43 Pogonomys undescribed species

1300–2800

x

ⱕ 3000

40 Pogonomys sp. cf. loriae Thomas, 1897

42 Pogonomys sp. cf. sylvestris Thomas, 1920

x

Endemic?1 Elevation2 (m) ⱕ 1800

Taxa

x

x

Northern flat lowlands3 Central Cordillera, northern4

39 Pogonomelomys mayeri (Rothschild & Dollman, 1932)

Number (see Chapter Appendix)

Table 4.10.2. (Continued)

North coast ranges5 ?

x

Yapen Island* Numfoor Island• x

?

Vogelkop Peninsula lowlands x

Salawati Island* ?

x

?

x

? ? ? x

x ?

?

?

?

?

?

? x

Batanta Island• Waigeo Island• Misool Island* Vogelkop Peninsula mountains6 Bird’s Neck lowlands Bird’s Neck mountains7 Eastern Papua, southern lowlands8 ?

Weyland Range10 Snow Mountains region11

Central Cordillera, southern9

? x

? x

x

? x

?

?e x ?

x

x x

Star Mountains10 ?d

?

?d

x

?d

702 / kristofer m . helgen

PAGE 702

Biak, Supiori, Padaido islands•

................. 16157$

ⱕ 700 E 850–2800 E 1000–2500

55 Rattus sordidus (Gould, 1858)

56 Rattus steini Ru¨mmler, 1935

57 Rattus unicolor (Ru¨mmler, 1935)

CH43

x

?

? ? ? ?

?

?

x

?

x

02-08-07 10:46:04

PS

x

?

(x)

?

x

x x

x x

x x

x x

x x

x

x x

x x

?

?d

?d

?d

x

x

Notes: See Table 4.10.1 for notes and legend. Taxonomy follows Appendix 4.10.1, which should be consulted for numbered notes on taxonomic issues. Elevational ranges (in meters; symbols as for Table 4.10.1) are drawn from vouchered specimens with explicit, associated altitudinal data in world museums (collected from all of mainland New Guinea and immediately adjacent islands, not just from Papua). Geographic designations and symbols follow Table 4.10.1. Additional species can be expected in Papua (Table 4.10.5).

?

?e

?

x

x

x

ⱕ 1600

x

?

?

?

x

64 Xenuromys barbatus (Milne-Edwards, 1900)

?

x

x

x

?b ?

x

x

x x ? x

? ? ?

x

⬍ 200

x

x

(Q) ?

x

x

63 Uromys scaphax Thomas, 1913

x

x

x

ⱕ 1200

E ⬍ 1000

x ?

x

ⱕ 1900

?

?

x

x

E ⬍ 1000 x

x

x ?

62 Uromys nero Thomas, 1913

61 Uromys emmae Groves & Flannery, 1994

60 Uromys caudimaculatus (Gray, 1866)

59 Uromys boeadii Groves & Flannery, 1994

850–3000

E 3200–4500

54 Rattus richardsoni Tate, 1949

58 Uromys anak Thomas, 1907

ⱕ 1500

1500–2600

E 2900–4000

ⱕ 1200

E ⱕ 600

ⱕ 2800

53 Rattus praetor (Thomas, 1888)

52 Rattus pococki Ellerman, 1941

51 Rattus omichlodes Misonne, 1979

50 Rattus leucopus (Gray, 1867)

49 Rattus jobiensis Ru¨mmler, 1935

48 Rattus foersteri (Ru¨mmler, 1935)

Taxonomic and Geographic Overview of the Mammals / 703

PAGE 703

................. 16157$

CH43

02-08-07 10:46:05

Nyctimene albiventer (Gray 1863)

Nyctimene cyclotis Andersen 1910)

Nyctimene draconilla Thomas 1922

7

8

9

Paranyctimene tenax Bergmans 2001

Nyctimene aello Thomas 1900

6

Paranyctimene raptor Tate 1942

Macroglossus minimus (E. Geoffroy 1810)

5

12

Dobsonia moluccensis (Quoy and Gaimard 1830)

4

11

Dobsonia minor (Dobson 1879)

3

Nyctimene species

Dobsonia emersa Bergmans & Sarbini 1985

2

10

Dobsonia beauforti Bergmans 1975

Species

1

Number

PS

probably restricted to north and west New Guinea

probably restricted to south New Guinea

insular (Numfoor, Biak-Supiori)

distribution poorly understood

widespread in all montane areas

widespread

widespread

widespread

widespread

widespread

insular (Numfoor, Biak-Supiori)

insular (West Papuan Islands)

Distribution in Papua1

0–1,600

0–1,200

⬍ 200

0–100

800–2,800

0–1,900

0–1,000

0–1,200

0–2,700

0–700

⬍ 200

⬍ 200

Elevation2 (m)

Table 4.10.3. Distributional and elevational ranges of the fruit bats of Papua, family Pteropodidae (23 recorded species)

704 / kristofer m . helgen

PAGE 704

................. 16157$

Pteropus pohlei Stein 1933

19

CH43

Syconycteris hobbit Ziegler 1982

23

montane; restricted to Central Dividing Ranges

widespread

widespread

insular (Biak, Waigeo)

insular (Yapen, Numfoor)

insular (recorded in Papua from Gag)

widespread

insular (recorded in Papua from Misool)

widespread

widespread (coastal and insular)

insular (recorded in Papua from Kofiau)

⬍ 200

1,800–2,800

0–3,000

0–2,200

⬍ 200

⬍ 300

⬍ 200

0–1,400

⬍ 200

0–500

0–1,000

Notes: Taxonomy follows Appendix 4.10.1, which should be consulted for numbered notes on taxonomic issues. Additional species can be expected in Papua (see Table 4.10.5). 1 Because most species are relatively widespread, only broad distributional statements are provided; some judgments about species’ distributions (e.g. ‘‘widespread’’) are based on analyses of conspecific museum material from Papua New Guinea. 2 Elevational ranges are drawn from vouchered specimens with associated altitudinal data in world museums (collected from all of mainland New Guinea and immediately adjacent islands, not just from Papua). ‘‘⬍ X’’ denotes that the upper altitudinal bound is not well established, but is likely to lie at or near X meters.

Syconycteris australis (Peters 1867)

22

Rousettus amplexicaudatus (E. Geoffroy 1810)

Pteropus personatus Temmink 1835

18

21

Pteropus neohibernicus Peters 1876

17

Pteropus new species

Pteropus melanopogon Peters 1867

16

20

Pteropus macrotis Peters 1867

Pteropus conspicillatus Gould 1850

14

15

Pteropus chrysoproctus Temminck 1837

13

Taxonomic and Geographic Overview of the Mammals / 705

02-08-07 10:46:05

PS

PAGE 705

Distribution in Papua1 widespread widespread widespread widespread montane; restricted to Central Cordillera widespread widespread probably restricted to south New Guinea restricted to Vogelkop, West Papuan Islands probably restricted to south New Guinea widespread widespread but rare in New Guinea insular (recorded in Papua from Gag) widespread restricted to south New Guinea widespread insular (recorded in Papua from Yapen) widespread widespread but rarely collected

Species

Family Hipposideridae (10 species) Aselliscus tricuspidatus (Temminck, 1835) Hipposideros ater Templeton, 1848 Hipposideros calcaratus (Dobson, 1877) Hipposideros cervinus (Gould, 1854) Hipposideros corynophyllus Hill, 1985 Hipposideros diadema (E. Geoffroy, 1813) Hipposideros maggietaylorae Smith and Hill, 1981 Hipposideros muscinus (Thomas and Doria, 1886) Hipposideros papua (Thomas and Doria, 1886) Hipposideros wollastoni Thomas, 1913

Family Rhinolophidae (2 species) Rhinolophus euryotis Temminck, 1835 Rhinolophus philippinensis Waterhouse, 1843

................. 16157$

CH43

Family Emballonuridae (7 species) Emballonura cf. alecto Eydoux and Gervais, 1836 Emballonura beccarii Peters and Doria, 1880 Emballonura furax Thomas, 1911 Emballonura raffrayana Dobson, 1879 Emballonura serii Flannery, 1994 Mosia nigrescens Gray, 1843 Saccolaimus saccolaimus (Temminck, 1838)

02-08-07 10:46:05

⬍ 100 0–1,500 0–1,200 0–1,600 0–300 0–1,600 0–200

0–1,800 0–1,300

0–600 0–1,700 0–600 0–1,400 1,400–2,700 0–1,300 0–300 0–600 0–300 0–800

Elevation2

Table 4.10.4. Elevational and distributional ranges of the insectivorous bat families of Papua (38 recorded species).

1 2 1 5 1 7 1

6 1

4 5 6 7 1 10 3 1 3 1

Number of records3

706 / kristofer m . helgen

PS

PAGE 706

................. 16157$

CH43

02-08-07 10:46:05

widespread but rarely collected widespread but rarely collected montane; restricted to Central Cordillera

Family Molossidae (3 species) Chaerephon jobensis (Miller, 1902) Mormopterus beccarii (Peters, 1881) Tadarida kuboriensis McKean and Calaby, 1968

0–1,400 0–300 1,900–3,000

0–1,500 0–3,200 0–2,000 0–1,600 0–2,900 0–2,800 0–1,200 ⬍ 100 0–1,500 0–2,600 0–2,100 0–1,300 0–2,400 700–3,000 0–1,300 ⬍ 200 1 1 1

6 2 1 4 4 1 4 2 2 2 2 1 1 2 6 1

Notes: Taxonomy follows Appendix 4.10.1, which should be consulted for notes on taxonomic issues. Additional species can be expected in Papua (see Table 4.10.5). 1 Because most species are relatively widespread, only broad distributional statements are provided; some judgments about species’ distributions (e.g. ‘‘widespread’’) are based on analyses of conspecific museum material from Papua New Guinea. 2 Elevational ranges are drawn from vouchered specimens with associated altitudinal data in world museums (collected from all of mainland New Guinea and immediately adjacent islands, not just from Papua). ‘‘⬍ X’’ denotes that the upper altitudinal bound is not well established, but is likely to lie at or near X meters. 3 Number of locality records in Papua.

widespread widespread widespread widespread widespread widespread but rarely collected widespread insular (recorded in Papua from Raja Ampats) seemingly patchy widespread widespread but rarely collected seemingly patchy widespread montane; restricted to Central Cordillera widespread restricted in Papua to Trans-Fly

Family Vespertilionidae (16 species) Miniopterus australis Tomes, 1858 Miniopterus macrocneme Revilliod, 1914 Miniopterus magnater Sanborn, 1931 Miniopterus propitristis Peterson, 1981 Miniopterus schreibersii (Kuhl, 1819) Murina florium Thomas, 1908 Myotis moluccarum (Thomas, 1915) Myotis cf. stalkeri Thomas, 1910 Nyctophilus bifax Thomas, 1915 Nyctophilus microtis Thomas, 1888 Philetor brachypterus (Temminck, 1840) Phoniscus papuensis (Dobson, 1878) Pipistrellus angulatus (Peters, 1880) Pipistrellus collinus Thomas, 1920 Pipistrellus papuanus (Peters and Doria, 1881) Scotorepens sanborni (Troughton, 1937)

Taxonomic and Geographic Overview of the Mammals / 707

PS

PAGE 707

................. 16157$

CH43

Occurs in PNG Trans-Fly Occurs in PNG Trans-Fly

Family Macropodidae Lagorchestes conspicillatus Gould, 1842 Thylogale stigmatica (Gould, 1860) Widespread in PNG Central Cordillera Occurs in southwest lowlands PNG Occurs in PNG Trans-Fly Occurs in Prince Alexander Mts, PNG Occurs in southwest lowlands PNG Occurs in Torricelli Mts, PNG Occurs in Star Mts region, PNG Occurs in PNG Trans-Fly Widespread in PNG Central Cordillera Occurs in PNG Trans-Fly

Occurs in southwest lowlands PNG

Family Peramelidae Echymipera echinista Menzies, 1990

Family Muridae Abeomelomys sevia (Tate & Archbold, 1935) Chiruromys vates (Thomas, 1908) Conilurus penicillatus (Gould, 1842) Hydromys ziegleri Helgen, 2005 Leptomys signatus Tate and Archbold, 1938 ‘‘Microhydromys musseri’’ Flannery, 1989 Pogonomys championi Flannery, 1988 Pseudomys delicatulus (Gould, 1842) Rattus verecundus (Thomas, 1904) Xeromys myoides Thomas, 1889

Occurs in PNG Trans-Fly Occurs in PNG Trans-Fly

Reason to expect occurrence

Family Dasyuridae Planigale novaeguineae Tate and Archbold, 1941 Sminthopsis archeri van Dyck, 1986

Species

Table 4.10.5. Species unrecorded but expected to occur in Papua (32 selected species)

Star Mts Southeast lowlands Trans-Fly North coast ranges Southeast lowlands North coast ranges Star Mts Trans-Fly Star Mts Trans-Fly

Trans-Fly Trans-Fly

Southeast lowlands

Trans-Fly Trans-Fly

Where in Papua to expect

708 / kristofer m . helgen

02-08-07 10:46:05

PS

PAGE 708

................. 16157$

CH43

Widespread in lowlands of PNG Widespread in lowlands of PNG Occurs in south lowlands of PNG Occurs in PNG Trans-Fly Occurs in PNG Trans-Fly

Family Rhinolophidae Rhinolophus arcuatus Peters, 1871 Rhinolophus megaphyllus Gray, 1834

Family Emballonuridae Emballonura dianae Hill, 1956 Saccolaimus mixtus Troughton, 1925 Taphozous australis (Gould, 1854)

02-08-07 10:46:06

PS

Scattered occurrence in PNG Scattered occurrence in PNG

Lowlands Lowlands

Trans-Fly Lowlands Lowlands Lower montane

South lowlands Trans-Fly Trans-Fly

Lowlands Lowlands

North coast ranges North coast ranges

Trans-Fly Trans-Fly Trans-Fly

All of these species are known from parts of Papua New Guinea adjacent to Papua and are judged to be extremely likely to extend into appropriate (but generally less wellsurveyed) parts of Papua, as indicated.

Family Molossidae Otomops papuensis Lawrence, 1948 Otomops secundus Hayman, 1952

Occurs in PNG Trans-Fly Widespread in lowlands of PNG Widespread in lowlands of PNG Scattered occurrence in PNG

Occurs in Bewani Mts, PNG Occurs in Bewani Mts, PNG

Family Hipposideridae Hipposideros edwardshilli Flannery and Colgan, 1993 Hipposideros fasensis Flannery and Colgan, 1993

Family Vespertilionidae Chalinolobus nigrogriseus (Gould, 1856) Kerivoula muscina Tate, 1941 Miniopterus medius Thomas and Wroughton, 1909 Nyctophilus timoriensis (Geoffroy, 1806)

Occurs in PNG Trans-Fly Occurs in PNG Trans-Fly Occurs in PNG Trans-Fly (coastal)

Family Pteropodidae Nyctimene sp. cf. cephalotes (Pallas, 1767) Pteropus alecto Temminck, 1837 Pteropus scapulatus Peters, 1862

Taxonomic and Geographic Overview of the Mammals / 709

PAGE 709

710 / kristofer m . helgen

ippines (Heaney et al. 1998), and the Greater and Lesser Antilles (Da´valos 2004). Though more species-rich, Papua’s mammalian endemism is less impressive than in these other faunas, but the present boundaries of Papua reflect political rather than geological history, so that the overwhelming majority of the mammals of Papua also occur in Papua New Guinea (Flannery 1995a). When the island of New Guinea is considered as a whole, especially in tandem with adjacent Melanesian archipelagos like the Bismarcks and Solomons, levels of mammalian endemism in this overall region are remarkably high in global context (Flannery 1995a,b, 1996). The native mammal fauna of New Guinea comprises 16 taxonomic families classified in six orders (Tables 4.10.1–4). These include monotremes (order Monotremata, represented by the Tachyglossidae); marsupials (including the orders Dasyuromorphia, represented by the Dasyuridae), Peramelemorphia (represented by the Peramelidae), and Diprotodontia (represented by the Phalangeridae, Burramyidae, Acrobatidae, Petauridae, Pseudocheiridae, and Macropodidae); rodents (order Rodentia, represented by the Muridae); and bats (order Chiroptera, represented by the Pteropodidae, Hipposideridae, Rhinolophidae, Emballonuridae, Vespertilionidae, and Molossidae). The members and basic ecological attributes of each of these mammalian families (as represented in Papua) are briefly discussed below. Wild living populations of non-native mammals are also discussed in a final section on introduced species.

Mammalian Families of Papua

echidnas (family tachyglossidae) There are two extant genera of tachyglossids: the short-beaked echidnas (Tachyglossus), widespread in Australia and New Guinea, and the long-beaked echidnas (Zaglossus), endemic to New Guinea. Both genera are represented in Papua. Echidnas are adorned with black and white dorsal spines, have a long, narrow rostrum (a ‘‘beak’’), and lack teeth. Both genera are animalivores, procuring terrestrial invertebrates with elongate tongues. Tachyglossus primarily consumes ants and termites, while Zaglossus specializes on earthworms. There is only one species of Tachyglossus (T. aculeatus), which is smaller than the species of Zaglossus, and has shorter, more brownish fur and a straighter and markedly shorter beak. Ecologically versatile, this species thrives in many different habitats, from deserts to montane forests, throughout Australia and Papua New Guinea; in Papua it is recorded only from littoral and monsoonal woodlands, grasslands, and swamp forests in the Trans-Fly region. Compared to Tachyglossus, species of Zaglossus are larger-bodied, possess a relatively much more elongate, down-curved beak, and generally have thicker, more blackish fur. Flannery and Groves (1998) demonstrated that at least three species of Zaglossus occur in New Guinea, each of which occurs in Papua (two species are endemic to the western part of the island). The earliest of these to be discovered was Zaglossus bruijnii, a species endemic to the greater Vogelkop region. This species’ historic distribution apparently included the lowlands of the Vogelkop

................. 16157$

CH43

02-08-07 10:46:07

PS

PAGE 710

Taxonomic and Geographic Overview of the Mammals / 711

Peninsula and the adjacent land-bridge island of Salawati, the Arfak, Tamrau, and Fakfak Mountains, and the Bird’s Neck region, probably as far east as the Charles Louis Range. Its current distribution is poorly documented, but it has probably contracted markedly due to hunting and disturbance in recent decades (Pasveer and Aplin 1998). For example, despite various historic and Quaternary records from lowland areas in the Vogelkop, most or all recent records of Z. bruijnii originate from upper montane forests, which are generally less accessible from human population centers. It probably remains common only in the more remote and thinly settled portions of its former range. The echidna of the New Guinean Central Cordillera is Zaglossus bartoni, distributed from the Wissel Lakes in the west to the mountains of Milne Bay Province (PNG) in the east. Though there are a number of historical and recent records of Z. bartoni from lowland forest (as low as sea level) in far eastern New Guinea, the subspecies in Papua (Z. b. diamondi) is not known to occur at elevations lower than 1,300 m. Like other Zaglossus, this species is generally rare near settlements and disturbed areas, but may remain more common in sparsely inhabited areas across its expansive montane range. Flannery and Groves (1998) also described a new species of small-bodied Zaglossus, Z. attenboroughi, based on a museum specimen collected in 1961 from the highest reaches of the Cyclops Mountains, where it is likely highly endangered or even extinct today (Figure 4.10.3). This species is also known from subfossil remains from Late Quaternary deposits in north coastal Papua New Guinea, where it is definitely extinct today (K. Aplin in litt.). This species may be found in the future in the isolated northern mountain ranges of Papua, such as the Foja and Van Rees ranges, which remain poorly explored (Helgen 2005).

dunnarts, dasyures, and quolls (family dasyuridae) The dasyurids are a family of small- to medium-sized insectivorous and carnivorous marsupials distributed throughout Australia and New Guinea (Krajewski et al. 2000). New Guinean dasyurid genera include the small-bodied dunnarts (Sminthopsis) and planigales (Planigale), generally weighing less than 40 g; the small- to medium-sized dasyures (Murexia, Phascolosorex, Neophascogale, and Myoictis) which weigh 30–300 g; and the quolls (Dasyurus), which can attain a weight of about one kilogram. The smaller dasyurids are primarily insectivorous, while the larger species feed especially on small vertebrates. Four dasyurid genera (Murexia, Phascolosorex, Neophascogale, and Myoictis), all of which occur in Papua, are endemic to New Guinea and adjacent land-bridge islands. Sminthopsis and Planigale are mainly Australian genera, and are represented in New Guinea only in the monsoonal woodlands and grasslands of the Trans-Fly, in the far south of the island. Only one species (Sminthopsis virginiae) is recorded from Papua (in the Trans-Fly region), but two others (Sminthopsis archeri and Planigale novaeguineae) occur in the eastern Trans-Fly (PNG), and with little doubt will be shown to extend to the Papua side as well. Murexia is a moderately diverse genus, distributed throughout New Guinea. Of

................. 16157$

CH43

02-08-07 10:46:07

PS

PAGE 711

712 / kristofer m . helgen

Figure 4.10.3. Three recently discovered species, all endemic to Papua. Left: Dendrolagus mbaiso, a secondarily terrestrial tree kangaroo from subalpine habitats in the Snow Mountains, described by Flannery et al. (1995). Top right: Zaglossus attenboroughi, a dwarf long-beaked echidna endemic to the Cyclops Mountains, described by Flannery and Groves (1998). Bottom right: Spilocuscus wilsoni, a dwarf spotted cuscus from the oceanic islands of Biak-Supiori, described by Helgen and Flannery (2004b). the five species in Papua, two are predominantly lowland species (M. longicaudata and M. melanura), while the remainder typically inhabit montane forests above 1,000 m (Table 4.10.1). Many of these species were formerly classified in Antechinus, an Australian genus to which they are not immediately related (e.g., Armstrong et al. 1998). Van Dyck (2002) recently erected multiple monotypic genera for these small dasyures, but I regard all of them as closely related species that comprise a monophyletic assemblage, and thus classify them in a single genus, Murexia (Table 4.10.1, Appendix 4.10.1). The medium-sized and rather colorful dasyures of the genera Phascolosorex and Neophascogale are scansorial and diurnal inhabitants of New Guinea’s montane forests (reviewed in phylogenetic context by Krajewski et al. 2004). The species of Phascolosorex are dark brown dasyures with silver tipping to the fur, a black dorsal stripe, and orange bellies. Three species occur in Papua, two of which are endemic to the western half of New Guinea (Table 4.10.1): P. doriae of the Arfak, Weyland, and Snow mountains, and P. dorsalis of the Arfaks. The single species of the mono-

................. 16157$

CH43

02-08-07 10:46:15

PS

PAGE 712

Taxonomic and Geographic Overview of the Mammals / 713

typic genus Neophascogale (N. lorentzii) is distributed along the Central Cordillera from the Weyland Range in the west to the Hagen Range of PNG. The three-striped dasyures (genus Myoictis) are widespread inhabitants of lowland and lower montane forests in New Guinea (Woolley 2005). Two species occur in Papua. Myoictis melas is distributed throughout almost all forested habitats of Papua below 1,800 m (including the Yapen, Salawati, Batanta, and Waigeo islands), with the exception of the Trans-Fly region, where M. wallacei occurs. New Guinea is home to two species of quolls (genus Dasyurus), both of which are found in Papua. Dasyurus albopunctatus occurs throughout New Guinea (apart from the Trans-Fly region) in forested areas from sea level to the tree line (Table 4.10.1), while Dasyurus spartacus is known only from the Trans-Fly region (van Dyck 1988), including Wasur National Park in Papua. Genetic studies have shown that the latter species is probably better recognized as a subspecies of the Australian species Dasyurus geoffroyi (Krajewski et al. 2000; K. Firestone, pers. comm.), with which it was regarded as conspecific in the past (Waithman 1979). Typically no more than three or four dasyurid species (in Papua, usually Dasyurus albopunctatus, Myoictis melas, Murexia longicaudata, and M. melanurus) cooccur locally in New Guinea’s lowland forests (⬍600–1,000 m). However, as many as six or seven dasyurids exist syntopically at higher altitudes. For example, six dasyurids have been collected together in forest situated at 2,800 m in the vicinity of Mt Trikora in the Snow Mountains (D. albopunctatus, Neophascogale lorentzii, Phascolosorex brevicaudata, M. habbema, M. naso, and M. wilhelmina). Dasyurid diversity decreases at yet higher elevations, such that only three species occur above 3,000 m (Table 4.10.1), and only one (N. lorentzii) typically extends above the tree line.

bandicoots (family peramelidae) Bandicoots are small- to medium-sized omnivorous marsupials, four genera of which are found in New Guinea (Echymipera, Peroryctes, Microperoryctes, and Isoodon). All species are terrestrial, generally living in burrows or nests on the ground by day and foraging for food along the ground at night. The larger forest-living bandicoots (Echymipera, Peroryctes, and the larger species of Microperoryctes) may feed especially on fallen fruits in addition to terrestrial invertebrates, while the smaller species (such as Microperoryctes aplini and M. murina) are probably more exclusively insectivorous. The bandicoots have usually been divided into two families (Peramelidae and Peroryctidae) following Groves and Flannery (1990), but recent studies have shown this arrangement to be cladistically untenable (e.g., Westerman et al. 1999), and I follow Groves (2005a) in classifying all New Guinean species in the family Peramelidae. Though most of New Guinea’s bandicoots live in closed forests, one species (Isoodon macrourus) typically inhabits monsoon woodland and grassland areas, and is present in Papua only in the Trans-Fly region. The species of Microperoryctes are exclusively montane, occurring in both mountain forests and subalpine grasslands along the Central Cordillera and in the Arfak Mountains. Three species

................. 16157$

CH43

02-08-07 10:46:16

PS

PAGE 713

714 / kristofer m . helgen

of Microperoryctes are endemic to Papua: M. murina, recorded only from Mt Sumuri in the Weyland Range; M. aplini of the Arfak Mountains (Helgen and Flannery 2004a); and an unnamed, relatively large species that occurs in montane forest and subalpine grasslands throughout the Snow Mountains (Table 4.10.1, Appendix 4.10.1). The sole species of the genus Peroryctes in Papua is P. raffrayana, a widespread species distributed in the Central Cordillera, Vogelkop Mountains, and the north coast ranges. Though common only above about 500 m, it extends down as low as sea level in foothill forests. Species of Echymipera are predominantly inhabitants of lowland forest, most commonly encountered below 1,000 m, though occasionally extending much higher (Table 4.10.1). Two species (E. kalubu and E. rufescens) co-occur throughout essentially all of lowland New Guinea and on many adjacent islands, and E. clara co-occurs locally with both of these throughout the northern lowlands. The poorly known species E. echinista is known only from the Fly and Strickland River drainages of south-central PNG, but probably extends to the southern lowlands of Papua as well. Bandicoots reach their greatest local diversity in forests along the slopes of the Central Cordillera. For example, exhaustive trapping along an altitudinal transect from 500 to 1,200 m along the northern margin of the Cordillera would reveal five species living in this same general area (E. clara, E. kalubu, E. rufescens, M. longicauda, and P. raffrayana). However, each of these species has different habitat and elevational preferences, and it would be unusual (if not impossible) to encounter all five species at exactly the same altitude. Despite wide recorded altitudinal ranges (Table 4.10.1), E. clara and E. rufescens are most common below 500 m, E. kalubu below 1,000 m, P. raffrayana from 800 to 2,000 m, and M. longicauda above 1,200 m. Three or four species generally co-exist locally in most lowland areas of New Guinea, but as a general rule no more than three bandicoot species occur in direct syntopy in New Guinean montane forests above 1,500 m, and only one species usually occurs in any given subalpine area.

cuscuses (family phalangeridae) The cuscuses are medium-sized (1.5–7 kg), nocturnal frugivore/folivores, most of which are arboreal. New Guinean species are classified in two genera, Phalanger (the cuscuses) and Spilocuscus (the spotted cuscuses). Three or four phalangerid species co-exist in most forested areas of New Guinea. Phalanger is one of the most diverse marsupial genera in the broader Australasian region (Flannery 1994b). Eight species occur on mainland New Guinea, six of which are present in Papua. Most species are arboreal and sleep in trees by day, though one species (P. gymnotis) is largely terrestrial and rests in burrows in the ground. Of the five species in Papua, two (P. sericeus and P. carmelitae) are restricted to montane forests along the Central Cordillera; one (P. vestitus) is widely but patchily distributed in a number of montane areas, including the Arfak, Tamrau, and Weyland ranges in Papua; one (P. gymnotis) is widespread both in lowland and montane forests up to 2,700 m, extending also to the land-bridge islands of Yapen, Misool, and Salawati in Papua; and two are committed inhabitants of

................. 16157$

CH43

02-08-07 10:46:17

PS

PAGE 714

Taxonomic and Geographic Overview of the Mammals / 715

lowland forests (P. orientalis in the northern and western lowlands and on many land-bridge and oceanic islands; P. mimicus in the southern lowlands). Species of Spilocuscus are restricted to lowland forests in New Guinea and tropical northern Australia, and on a number of adjacent islands. Two species inhabit the mainland of western New Guinea. One, the often beautifully patterned Common Spotted Cuscus (S. maculatus), is widespread in lowlands throughout New Guinea. The somewhat larger-bodied Black-spotted Cuscus (Spilocuscus rufoniger) is highly susceptible to local human hunting pressure and is patchily distributed throughout lowland forests north of New Guinea’s Central Cordillera, on the Bird’s Neck, and in the Vogelkop (Flannery 1995a; Feiler 1978; Helgen, unpublished data). The largest of the New Guinean cuscuses, both S. maculatus and S. rufoniger can exceed six kilograms in weight. Smaller-bodied endemic species of Spilocuscus occur on certain oceanic islands in the vicinity of New Guinea, including on Waigeo and Batanta (S. papuensis) and Biak-Supiori (S. wilsoni) in Papua (Helgen and Flannery 2004b).

pygmy possums (families burramyidae and acrobatidae) Two families of small-bodied possums (Burramyidae and Acrobatidae) occur in New Guinea (including Papua), where each is represented by a single widespread species endemic to New Guinea. Cercartetus caudatus is a very small burramyid possum (ca 20 g) that occurs at elevations above 1,000 m along the length of the Central Cordillera and in the Arfak Mountains. It is a scansorial insectivore common both in montane forests and subalpine grasslands. This species was formerly considered to occur also in northern Queensland (i.e., as the subspecies C. caudatus macrurus), but molecular studies suggest that the Australian population is better classified as a separate species (Osborne and Christidis 2002a). Distoechurus pennatus is a somewhat larger acrobatid possum (40–60 g) with a terminally distichous (‘‘feathered’’) tail. It is an arboreal insectivore and frugivore and is widespread in closed lowland and lower montane forests throughout the island of New Guinea below 1,900 m. A second acrobatid species, Acrobates pennatus, is known only from eastern Australia, but may also occur in New Guinea, from where it is provisionally recorded by two poorly provenanced specimens (Helgen, 2003).

striped possums and gliders (family petauridae) The marsupial family Petauridae is represented in New Guinea by the striped possums or trioks (Dactylopsila and Dactylonax) and the gliders (Petaurus). Trioks are small- to medium-sized (⬍600 g) nocturnal insectivores specializing on extracting wood-boring insects from tree cavities and fallen logs, with elongate and narrowed fourth fingers, grossly enlarged first incisors, and a bold black-andwhite dorsal striping pattern. The species of Dactylopsila are arboreal and nest in tree holes, while Dactylonax (represented by a single species, D. palpator) is more scansorial, nesting both in tree hollows and in underground burrows. Three

................. 16157$

CH43

02-08-07 10:46:18

PS

PAGE 715

716 / kristofer m . helgen

striped possums occur in Papua: Dactylopsila trivirgata (widespread, generally in lowland and lower montane forests), Dactylopsila megalura (in montane forests in the Snow and Star Mountains and the Weyland Range), and Dactylonax palpator (in montane forests throughout the Central Cordillera and in the Arfak Mountains). Dactylonax palpator commonly co-occurs with either D. trivirgata or D. megalura in montane forests, but the latter two species only rarely co-occur locally, with D. megalura typically occurring at higher elevations in areas of geographic overlap. The genus Petaurus is represented in Papua by a single species, the Sugar Glider (Petaurus breviceps), a common, small-bodied (50–100 g) species distributed throughout Australia, New Guinea from sea level to at least 3,000 m, and on many adjacent islands, including Misool, Salawati, Yapen, Numfoor, and Biak-Supiori in Papua. The Sugar Glider feeds on fruits and leaves, insects, nectar, and sap. It is nocturnal, arboreal, and nests in tree hollows, and is a versatile species that can be found in almost any treed habitat, from regrowth and floodplain forests to undisturbed montane forests. The larger Northern Glider (Petaurus abidi) is known only from hill and montane forests in the Torricelli Range of northern PNG (where it co-occurs with P. breviceps), but might well be found in the future to occur in the poorly surveyed north coast ranges of Papua.

ringtail possums (family pseudocheiridae) Ringtail possums are relatively small arboreal leaf eaters restricted to forested areas in New Guinea and Australia. Two genera occur in Papua: the smaller, more gracile species of Pseudochirulus (⬍1 kg), probably the smallest obligately folivorous mammals, and the larger, chunkier species of Pseudochirops (750–2,500 g). Most ringtails occur primarily in undisturbed forest. One species, Pseudochirulus canescens, is restricted to lowland and lower montane forests and several others extend on occasion into lowland forest, but the majority of pseudocheirids are montane (Table 4.10.1). In general most species are common only at or above about 1,200 m. As many as five ringtails species can be encountered along an elevational transect from 1,000 to 2,000 m along both the northern or southern slopes of Papua’s Central Cordillera, and up to four species (each of slightly different size and habits) can be found in direct syntopy in midmontane forests along much of the Cordillera. For example, Pseudochirulus mayeri, Pseudochirulus caroli, Pseudochirops albertisii, and Pseudochirops cupreus co-occur directly in Papua’s Weyland Range in forests situated at about 2,000 m; a somewhat different taxonomic assemblage (P. mayeri, P. cupreus, Pseudochirulus forbesi, and Pseudochirops cupreus) co-exists in forests situated at about 2,200 m in the Star Mountains in far eastern Papua (and in much of Papua New Guinea). Only two ringtails (P. cupreus and P. mayeri) occur in subalpine grasslands above the tree line. Presumably because of their exacting dietary requirements and typically montane occurrence, ringtails are poor overwater dispersers and do not extend to any oceanic islands. The majority of Papua’s nine species of ringtails have relatively wide distributions in New Guinea. However, three species are endemic to Papua, two of which

................. 16157$

CH43

02-08-07 10:46:18

PS

PAGE 716

Taxonomic and Geographic Overview of the Mammals / 717

(Pseudochirops coronatus and Pseudochirulus schlegeli) are known only from the Arfak Mountains; the other (P. caroli) is known from the western Central Cordillera, from the Baliem Valley to the Weyland Range. Another species (P. albertisii) has a disjunct distribution throughout the mountains of northern New Guinea (records in Papua are from the Arfak, Weyland, and Cyclops ranges, and from mountains on the island of Yapen). The painted ringtail, Pseudochirulus forbesi, is widespread in PNG but extends into Papua only in the Star Mountains; however, it also occurs in the Bewani and Torricelli ranges of northern PNG and may yet be found to occur in Papua’s disjunct northern ranges as well. Ringtails are found on only two land-bridge islands, Yapen and Salawati, both in Papua (P. canescens on both islands, and P. albertisii in the mountains of Yapen).

wallabies and kangaroos (family macropodidae) The kangaroos and wallabies are bipedally hopping, herbivorous marsupials. Most are obligately terrestrial, but a single genus, well represented in Papua (Dendrolagus, comprising the tree kangaroos), is almost exclusively arboreal. Five genera of macropodids occur in New Guinea, including Papua. Macropus, primarily an Australian genus, is represented in New Guinea by a single species, the Agile Wallaby (M. agilis), the largest extant native mammal in Melanesia (up to 25 kg). An inhabitant of monsoonal woodlands, grasslands, and open forests, the Agile Wallaby occurs throughout both southern New Guinea and northern Australia; in Papua it is restricted to the Trans-Fly region. The genus Dorcopsis comprises a number of small wallabies (averaging 5–6 kg) restricted to closed lowland forest habitats in New Guinea below 800 m elevation (Table 4.10.1; a record of D. hageni from 1,546 m discussed by Hedemark et al. 1997 cannot be substantiated). Three species with parapatric distributions occur in Papua, essentially with northern (D. hageni), western (D. muelleri), and southern (D. luctuosa) distributions. One of these, Dorcopsis muelleri, is endemic to Papua, and extends to the land-bridge islands of Yapen, Misool, and Salawati; it may also occur on Batanta and Waigeo but there are no confirmed records to date. The genus Dorcopsulus, endemic to New Guinea, consists of small-bodied wallabies (2–3 kg) restricted to closed montane forests above 800 m, where they replace the species of Dorcopsis found at lower elevations. One species, Dorcopsulus vanheurni, is found in Papua, where it is widely distributed throughout the mountain ranges of the Central Cordillera. Though extinct in outlying montane areas today, subfossil remains demonstrate that this species occurred in the Vogelkop (Aplin et al. 1999) and North Coast Ranges of PNG (K. Aplin in litt) until relatively recently. Pademelons (genus Thylogale) occur in Australia, New Guinea, and on a number of adjacent land-bridge and oceanic islands. Two living species are known from Papua: T. browni from the northern lowlands, and T. brunii from the southern lowlands. One additional species (T. stigmatica) occurs in the Trans-Fly of PNG and undoubtedly will be shown to extend to similar habitats in Papua. Two additional species (T. christenseni and T. sp. cf. calabyi) are known from late Qua-

................. 16157$

CH43

02-08-07 10:46:19

PS

PAGE 717

718 / kristofer m . helgen

ternary remains from the Snow Mountains (Flannery 1999). These montane species are thought to be extinct today (Flannery 1995a) but may yet be found to survive in some areas and should be sought in future surveys in the region. The most famous macropodids of New Guinea are the tree kangaroos (Dendrolagus), six species of which occur in Papua (Table 4.10.1). These include one Vogelkop area endemic (D. ursinus), one species widespread in the northern lowlands and the Vogelkop area (D. inustus), one species known from Papua only by a sight record from the isolated northern Foja Mountains (D. pulcherrimus), one known only from the north coastal Wandammen Mountains (D. mayri), and two species from the broad Snow Mountains region of the western Central Cordillera (D. stellarum and D. mbaiso). Most tree kangaroo species are fully arboreal forest dwellers, but the most recently discovered species, D. mbaiso (Figure 4.10.3), is a tree kangaroo that has secondarily evolved adaptations for terrestrial life and occurs in both upper montane forest and subalpine grassland (Flannery et al. 1995, 1996).

rats (family muridae) A single family of rodents (the Muridae) occurs in the greater Australo-Papuan region. Murids are by far the most diverse mammalian family in New Guinea, where they have undergone a series of impressive ecomorphological radiations. They are today represented by a large number of endemic genera of varying size and habits, as well as by genera that are geographically more widespread (such as Rattus and Melomys). Resolution of the systematic relationships of AustraloPapuan murids and the geographic and temporal patterns of their diversification is a longstanding but as yet unrealized goal in Australasian mammalogy, though relevant methods, samples, and molecular markers are now available for such a comprehensive study. A total of 23 genera (65 species) of murid rodents have been collected in Papua to date. No currently described genus is restricted to Papua, but many species are found nowhere else (Table 4.10.2). A detailed review of each genus is well beyond the scope of this chapter, and recent reviews of the systematics (Musser and Carleton 2005) and basic ecology (Flannery 1995a) of each species are available elsewhere. In broad summary, the murid fauna of Papua includes amphibious animalivores (Baiyankamys, Crossomys, Hydromys, Parahydromys), small terrestrial insectivores or animalivores (Leptomys, Mayermys, Pseudohydromys, Microhydromys), small scansorial omnivores (Lorentzimys), small scansorial and arboreal herbivores (Coccymys, Melomys, Pogonomys, Pogonomelomys), small- and medium-sized terrestrial herbivores and omnivores (Mammelomys, Paraleptomys, Paramelomys, Rattus), medium-sized scansorial herbivores (Anisomys), large scansorial herbivores (Mallomys, Uromys), and large terrestrial herbivores (Hyomys, Xenuromys). The general habits of at least one genus (Macruromys) remain almost completely obscure. Size variation in the family is enormous. The smallest New Guinean rodents (Microhydromys and Lorentzimys) weigh less than 15 grams, while species

................. 16157$

CH43

02-08-07 10:46:20

PS

PAGE 718

Taxonomic and Geographic Overview of the Mammals / 719

from a number of genera (Uromys, Xenuromys, Mallomys, and Hyomys) can exceed one kilogram in weight, with the larger species of Mallomys (probably the world’s largest living murines) weighing up to two kilograms or more. More than 30 murid rodent species have been collected in lower montane forest at a single site in PNG, Mt Sisa in Southern Highlands Province (Leary and Seri 1997; Aplin in litt.; Helgen, in prep.). Detailed, well-designed studies of murid diversity across local elevational gradients in New Guinea are as yet unavailable, but across the island, generic (and thus trophic) diversity appears to be greatest in lower and midmontane forests. Murid communities of most areas of Papua (essentially everywhere except the vicinity of Lake Habbema and Mt Trikora in the Snow Mountains) remain relatively poorly inventoried and remain an important priority for biological exploration in Melanesia. Many species new to science probably remain to be discovered or described, and many species currently recorded only from PNG will undoubtedly be shown to extend to Papua, especially in the Trans-Fly region, the Star Mountains, and the North Coast Ranges (Table 4.10.5).

fruit bats (family pteropodidae) Old World fruit bats, classified in the family Pteropodidae, are particularly diverse in Papua. The family is represented by large-bodied frugivorous flying foxes (Pteropus), small- to medium-sized frugivorous rousette bats (Rousettus) and barebacked fruit bats (Dobsonia), small frugivorous and insectivorous tube-nosed bats (Nyctimene and Paranyctimene), and the nectarivorous blossom bats (Syconycteris and Macroglossus). The upper elevational limit for the great majority of fruit bat species in Papua lies below 1,400 m, and pteropodid bats are thus most diverse and abundant in coastal and flat lowland forest. Indeed, below 1,000 m at least 13 species can occur in the same immediate area. Nevertheless, four species (Dobsonia moluccensis, Syconycteris australis, Rousettus amplexicaudatus, and Nyctimene albiventer) have wide altitudinal ranges and extend into midmontane forest or higher (Table 4.10.3) and two species (Nyctimene cyclotis and Syconycteris hobbit) are montane forest specialists that do not occur in the lowlands. The largest fruit bats of New Guinea—flying foxes of the genus Pteropus—are committed inhabitants of lowland habitats, especially coastal forests and forests on offshore islands. Only three species of Pteropus commonly occur on mainland New Guinea. Of these, one species (Pteropus conspicillatus) is generally restricted to coastal areas and nearby islands, including the Raja Ampats and the islands of Cenderawasih Bay. The remaining two species (P. neohibernicus and P. macrotis) inhabit the coast but also extend extensively inland, especially along river systems. Pteropus macrotis also occurs on a number of land-bridge islands surrounding New Guinea, including Salawati and Ara Island off the western Vogelkop, but does not extend to any oceanic islands. P. neohibernicus occurs patchily on islands both to the east and west of New Guinea (e.g., Gebe and Gag in Papua), but does not occur on islands in Cenderawasih Bay and is also unrecorded from the larger Raja Ampat Islands (e.g., Salawati and Waigeo). Five additional flying fox species are

................. 16157$

CH43

02-08-07 10:46:21

PS

PAGE 719

720 / kristofer m . helgen

known in Papua only from islands fringing New Guinea (Table 4.10.3); two of these (Pteropus pohlei and a related, undescribed species) are endemic to Papua. Across the geographic range of the bare-backed fruit bats (genus Dobsonia), no more than two species occur in syntopy. On the mainland of Papua and on Yapen, the two species present are D. moluccensis and D. minor, but different combinations prevail on other offshore islands, such as Biak-Supiori (D. emersa and D. beauforti) or the Raja Ampats (D. beauforti and D. moluccensis). In each case, these sympatric species generally segregate both by size and by roosting habits (one primarily in caves, another primarily in foliage). Dobsonia emersa (known only from Numfoor, Biak-Supiori, and the Padaido Islands) is endemic to Papua. The tube-nosed bats (Nyctimene and Paranyctimene) comprise one of the most distinctive lineages of Old World fruit bats. Though largely frugivorous, tubenosed bats are thought to rely on insects as well as fruit, at least to a greater extent than most other pteropodids. Though species of Nyctimene are distributed from the Philippines, Sulawesi, and the Lesser Sundas in the west to the Solomon Islands in the east, the genus is geographically centered on, and most diverse in, New Guinea, where three to four species commonly occur syntopically. The related genus Paranyctimene is restricted to mainland New Guinea and certain associated islands (e.g., Waigeo, Salawati, and Yapen in Papua), but taxonomic boundaries within the genus remain very poorly delineated (Bergmans 2001). The small blossom bat genus Syconycteris is another fruit bat lineage centered on New Guinea, extending elsewhere only to the Moluccas, Bismarcks, and East Papuan Islands. Two species occur in New Guinea, including Papua: the widespread and versatile S. australis, common in most habitats below 3,000 m, and S. hobbit, patchily distributed in moss forests above 1,800 m along much of the Central Cordillera. Many of Papua’s smaller species of fruit bats (such as Syconycteris australis, Macroglossus minimus, Rousettus amplexicaudatus, and Nyctimene albiventer) are common, highly adaptable species that occur in a wide variety of habitats, including both old growth and secondary forest and more degraded habitats such as native gardens and village environs. In fact, almost all lowland fruit bats thrive in gardens, and a number of species (especially M. minimus and R. amplexicaudatus) seem to occur at highest density in such anthropogenically disturbed habitats.

insectivorous bats (families hipposideridae, rhinolophidae, emballonuridae, vespertilionidae, and molossidae) The order Chiroptera (which includes all bats) has traditionally been classified into two suborders: the Megachiroptera (or ‘‘megabats’’), incorporating the single family Pteropodidae (the Old World fruit bats); and the Microchiroptera (or ‘‘microbats’’), comprising, at least in the Melanesian region, a number of families of generally smaller-bodied, insectivorous species. However, a considerable body of recent research, drawing largely from molecular comparisons, has now shown that

................. 16157$

CH43

02-08-07 10:46:22

PS

PAGE 720

Taxonomic and Geographic Overview of the Mammals / 721

microbats are not monophyletic (they represent a grade rather than a clade), and that some families are thus more closely related to megabats than to other lineages of microbats (e.g., Teeling et al. 2005). Although the distinction between megabats and microbats is not a natural one in a phylogenetic sense, it remains useful in terms of very broadly distinguishing basic ecological guilds (at least among Melanesian bats) into species specializing on fruit and nectar (Pteropodidae) and insectivorous species (other families). Mainland New Guinea’s microbats include species from five families: the leafnosed bats (Hipposideridae genera Aselliscus and Hipposideros), the horseshoe bats (Rhinolophidae: genus Rhinolophus), the sheath-tailed (or sac-winged) bats (Emballonuridae: genera Emballonura, Mosia, Saccolaimus, and Taphozous), the evening bats (Vespertilionidae: genera Chalinolobus, Myotis, Philetor, Pipistrellus, Scotorepens, Murina, Nyctophilus, Pharotis, Kerivoula, Phoniscus, and Miniopterus), and the free-tailed bats (Molossidae: genera Chaerephon, Mormopterus, Otomops, Tadarida). In addition, the family Megadermatidae is almost certainly represented in southeastern New Guinea by the genus Macroderma (Filewood 1983), but I know of no vouchered records to date. Each of these families and genera has distinctive ecomorphological features, but a detailed systematic and ecological review is beyond the scope of this chapter (see Bonaccorso 1998 and references therein). Though all of the genera listed above are recorded from PNG, five (Taphozous, Chalinolobus, Pharotis, Kerivoula, and Otomops) are not yet known from Papua. Though one (Pharotis) may well be legitimately absent from Papua, the lack of records for these other genera illuminates the paucity of scientific attention that the insectivorous bat fauna of western New Guinea has received to date. So, too, does the fact that that many particular species that are widespread in PNG or broader Melanesia (e.g., Rhinolophus arcuatus, Miniopterus medius, Emballonura dianae) have yet to be definitively recorded from western New Guinea, although they must occur there. Fresh efforts, including collection of specimens from many localities with concerted harp trapping, remote recording of echolocatory calls, and renewed taxonomic and molecular comparisons, will greatly clarify the systematic status and geographic distribution of Papua’s insectivorous bats, and will undoubtedly reveal them to be much more diverse than currently realized. Like fruit bats, New Guinean microbats reach their greatest diversity and abundance in coastal and flat lowland forests, and the great majority of New Guinean species (especially hipposiderids and emballonurids) do not extend above about 1,600 m (Table 4.10.4). Nevertheless, three species, including one molossid (Tadarida kuboriensis), one hipposiderid (Hipposideros corynophyllus), and one vespertilionid (Pipistrellus collinus) occur exclusively in montane forests along New Guinea’s Central Cordillera (in both Papua and PNG), and many vespertilionid species also extend into forests at higher elevations, with an upper altitudinal limit usually between 2,000 and 3,000 m. Currently no insectivorous bat species are known to be endemic to Papua.

................. 16157$

CH43

02-08-07 10:46:23

PS

PAGE 721

722 / kristofer m . helgen

non-native mammal species (dogs, pigs, rats and mice, shrews, deer, macaques, and cats) In addition to nearly 200 native species, a number of non-native mammals are represented in Papua today by wild-living populations. Some of these, such as dogs (Canis lupus familiaris), pigs (Sus scrofa), and Pacific rats (Rattus exulans) were apparently introduced in prehistoric times, probably 3,500 to 3,000 years ago, in tandem with Austronesian (Lapita) cultural expansion throughout the Pacific region (Kirch 1997). Others, including various commensal rodents, deer, and macaques result from introductions in the last few centuries or decades. Real and potential ecological impacts, negative or positive, of introduced mammal species on New Guinea’s native biota have received very little study to date and are worthy of further research. Dogs are represented in New Guinea both by domestic dogs, commonly used for hunting throughout the island, and by distinctive highland populations of feral dogs, often referred to as ‘‘New Guinea singing dogs’’ (Brisbin et al. 1994). Wildliving dogs have probably been present in New Guinea for only about 3,000 years (Hope 1981). Though highland ‘‘singing dog’’ populations are biologically interesting and deserve continued ecological study, neither molecular (e.g., Leonard et al. 2002) nor morphological evidence supports the taxonomic claim that these dogs represent a distinct canid species (‘‘Canis hallstromi’’), as some advocates have argued (see Koler-Matznick et al. 2003). Pigs (Sus scrofa) are widespread in New Guinea, both as wild-living and domesticated animals, and are the largest wild mammals found in Melanesia today. The fossil record demonstrates that pigs have been present in New Guinea for at least several thousand years and perhaps for much longer (6,000 to 10,000 years), though the accuracy of these earlier dates is uncertain (see Flannery 1995a; Larson et al. 2005). Pigs play a fundamental role in the economy and cultural heritage of Melanesian society (e.g., Hide 2004) and can be found today in essentially all geographic areas and at almost any elevation in Papua. The most common and widespread commensal murid in New Guinea is the Pacific Rat, Rattus exulans. This species is widespread throughout Southeast Asia, Melanesia, and on most island groups in the southwest Pacific (Flannery 1995b). In New Guinea it occurs from sea level to nearly 3,000 m, and is ecologically versatile, occurring in a wide array of habitats but preferring anthropogenically modified landscapes, especially village environs, gardens, and kunai grassland (Taylor et al. 1982). Rattus exulans is now relatively widespread in New Guinea and may have arrived several millennia ago in tandem with the Lapita expansion. Other non-native murids in New Guinea are more narrowly associated with human settlements and have probably arrived only in the last several hundred years. At least two members of the Rattus rattus species complex (identified as R. rattus and R. tanezumi by Musser and Carleton 1993, 2005) can be found in villages and towns in Papua, primarily in the lowlands (Taylor et al. 1982), as can the House Mouse, Mus musculus. The Brown Rat, Rattus norvegicus, is known from a few major port cities in New Guinea, including Jayapura in Papua, and

................. 16157$

CH43

02-08-07 10:46:24

PS

PAGE 722

Taxonomic and Geographic Overview of the Mammals / 723

two other Southeast Asian commensal rats, Rattus nitidus and R. argentiventer, are also established in Papua (in the Vogelkop Peninsula and Jayapura area, respectively). The House Shrew (Suncus murinus) is a lipotyphlan insectivore that lives in commensal contexts throughout south and Southeast Asia, including the Malay Archipelago as far east as the Moluccas and the Aru Islands (Corbet and Hill 1992). There is a single published record of S. murinus in New Guinea (Heim de Balsac and Heim de Balsac 1956), based on a specimen supposedly collected from the north coast of the Vogelkop Peninsula in 1877. No other New Guinea specimens have come to light, and Menzies (1996b) considers the record to be suspect. If S. murinus is present in Papua, it is probably restricted to major coastal towns and villages. Several other introduced mammal species are now established in Papua. Wildliving populations of the Rusa Deer (Cervus timorensis), native to Southeast Asia, occur across much of lowland Papua, including the Trans-Fly and entire southern lowlands, the Vogelkop Peninsula, and the vicinity of Jayapura (Flannery 1995a). A wild-living population of Long-tailed Macaques (Macaca fascicularis), a monkey species native to Southeast Asia, has become established in the immediate vicinity of Jayapura in recent decades (Kemp and Burnett 2003; Chapter 7.7). Continuing efforts are needed to eradicate, or at least prevent the spread of, macaques in Papua, as these versatile primates (prehistorically and historically established on many islands where they are not native, including Mauritius, Palau, and the Lesser Sundas) have the potential for considerable negative impact on native wildlife. Feral cats (Felis [sylvestris] catus) probably occur in many areas of Papua today, where they may negatively impact native small mammals and native predators, especially quolls (Flannery 1995a).

Distribution of the Mammals of Papua: Major Zoogeographic Regions, Regional Diversity, and Endemism Because the island of New Guinea is topographically and geologically so highly complex (cf. Chapter 2.1), it is not surprising that mammalian species richness and endemism are unevenly distributed across the island (Flannery 1995a; Heads 2002). Each of New Guinea’s mammal species has different ecological requirements and tolerances, such that some taxa are widely distributed both geographically and altitudinally, while many others have relatively restricted ranges, often limited to one or several discrete montane areas or particular offshore islands (Tables 4.10.1–2, Figures 4.10.4–8). The accounts below review the differential assemblages of mammals in various areas of western New Guinea, emphasizing zones of biogeographic importance, especially those notable for their rangerestricted mammalian endemism and those which remain important priorities for future study. While this review focuses largely on the modern fauna of western New Guinea, additional perspective from the Quaternary mammalian fossil record of Papua (especially for the Snow Mountains and the Vogelkop Peninsula) can be

................. 16157$

CH43

02-08-07 10:46:26

PS

PAGE 723

724 / kristofer m . helgen

Figure 4.10.4. Altitudinal distributions of Papuan mammal fauna taxa. Lowland: most common below 1,200 m, with upper altitudinal limit below 2,000 m. Montane: most common above 1,200 m, with lower altitudinal limit above 700 m on mainland New Guinea and above 300 m on adjacent islands. Versatile: spans a great range of elevations. derived from the work of Hope (1981), Aplin (1998), Aplin et al. (1999), Pasveer and Aplin (1998), and Flannery (1992, 1994a, 1995a, 1999). Because the geographic distributions of most insectivorous bat species in Papua remain especially poorly understood, the following discussion of geographic patterning in the New Guinea mammal fauna is drawn only from monotremes, marsupials, rodents, and pteropodid bats.

northern papua: the northern lowlands north coast ranges, and cenderawasih bay islands The lowland forests distributed throughout the central body of New Guinea north of the Central Cordillera are better known mammalogically than other lowland areas of the island. Historically, mammal assemblages from these northern lowland forests have been studied in detail in a few areas of Papua (Humboldt Bay and the foothills of the Cyclops, the island of Yapen, the Mamberamo Basin, and the foothills of the Snow Mountains along the Idenburg River) but also from a larger number of relevant sites in the northern lowlands of PNG. Drawing from inventory data generated at these sites, a number of observations and generalizations can be made about this northern fauna. For example, a core assemblage

................. 16157$

CH43

02-08-07 10:46:51

PS

PAGE 724

Taxonomic and Geographic Overview of the Mammals / 725

Figure 4.10.5. Altitudinal distributions of all Papuan mammal fauna (top) and endemic mammals of Papua (bottom). (For definitions, see Figure 4.10.4.) Half of all the mammals of Papua (top) are lowland species (split evenly between bats and nonvolant mammals), while a third (mostly nonvolant taxa) are restricted to montane areas. Papuan endemic mammals split almost evenly into montane and lowland species, and relatively few endemic mammals are versatile, occupying habitats at a range of altitudes. Papua’s endemic species are often geographically restricted taxa that have evolved in isolation on offshore islands (lowland habitats) or in discrete montane areas.

................. 16157$

CH43

02-08-07 10:47:27

PS

PAGE 725

726 / kristofer m . helgen

Figure 4.10.6. Biogeographic distributions of Papuan mammal fauna taxa. Mainland: occurs only on the New Guinea mainland. Continental: occurs on the New Guinea mainland and/or other continental landmasses (i.e., adjacent land-bridge islands and/or Australia). Oceanic: occurs only on oceanic (non-land-bridge) islands (e.g., Biak-Supiori). Widespread: occurs on both continental and oceanic landmasses. Papua’s monotremes and marsupials and its rodents have broadly similar distributions. However, more marsupials and monotremes extend to other land-bridge areas (mainly adjacent continental islands), which suggests that the extinction rate for rodents on islands is higher than for marsupials and monotremes. The greater percentage of rodents shared between oceanic and continental areas probably reflects a faster overwater dispersal rate, especially if rodent turnover rates are high. Bats have a very different distribution: only about one-quarter are restricted to the mainland or to adjacent continental areas; the rest extend to oceanic islands. This demonstrates the superior dispersal ability of bats relative to non-flying mammals, but may also reflect a taxonomic ‘‘lumping’’ in Papua’s bats, which have received less systematic attention than Papua’s nonvolant mammals (Figure 4.10.2). of 16 marsupial species (Dasyurus albopunctatus, Murexia longicaudata, Murexia melanura, Myoictis melas, Echymipera clara, Echymipera kalubu, Echymipera rufescens, Dendrolagus inustus, Dorcopsis sp. [D. hageni east of the Mamberamo River, D. muelleri to the west], Phalanger orientalis, Phalanger gymnotis, Spilocuscus maculatus, Spilocuscus rufoniger, Dactylopsila trivirgata, Petaurus breviceps, and Pseudochirulus canescens), at least seven rodent species (Hydromys chrysogaster, Mammelomys rattoides, Melomys rufescens, Paramelomys platyops, Pogonomys macrourus, Rattus praetor, and Uromys caudimaculatus), and ten fruit bat species (Dobsonia minor, Dobsonia moluccensis, Macroglossus minimus, Nyctimene aello,

................. 16157$

CH43

02-08-07 10:47:47

PS

PAGE 726

Taxonomic and Geographic Overview of the Mammals / 727

Figure 4.10.7. Distribution of all native mammals (top) and endemic mammals (bottom) of Papua. (For definitions, see Figure 4.10.6.) Compared with the mammal fauna of Papua as a whole, Papua’s endemic mammal species tend to be disproportionately restricted to oceanic islands or discrete areas of the mainland (many to particular mountain ranges; see Figures 4.10.5, 4.10.8). Nyctimene albiventer, Paranyctimene sp., Pteropus macrotis, Pteropus neohibernicus, Rousettus amplexicaudatus, and Syconycteris australis) are present almost everywhere in the lowland forests of this region, and most of these species (at least 82%) extend also to the land-bridge island of Yapen in Cenderawasih Bay. The majority of these taxa are restricted to (or overwhelmingly most common in) forests below 1,000 m (Tables 4.10.1–3), though a handful (e.g., Dasyrus albopunctatus, Petaurus breviceps, Syconycteris australis) thrive across a great elevational

................. 16157$

CH43

02-08-07 10:48:18

PS

PAGE 727

728 / kristofer m . helgen

Figure 4.10.8. Geographic distribution of Papua’s montane endemic mammals. ‘‘Broad’’ includes species distributed across the Vogelkop and Weyland ranges and the Snow Mountains. The expansive Snow Mountains are home to more than half the montane endemic mammals of Papua. The smaller Weyland and Arfak ranges support remarkably distinct mammal communities and exhibit levels of endemism disproportionate to their area. The unique Weyland mammal fauna, last studied in the 1920s, needs new mammal inventories. The distinctive Arfak mammal fauna signals a need to survey the mammal faunas of other Vogelkop-area ranges, such as Tamrau, Fakfak, and the Waigeo mountains. Mammal faunas from Papua’s North Coast Ranges (including the Cyclops and Wondiwois, each of which supports a single endemic mammal species, and the Fojas and Van Rees, which remain essentially unknown mammalogically) are particularly poorly known, and future surveys will likely reveal additional endemics. span. With sufficient survey efforts, this entire group of mammals can probably be expected at essentially any forested mainland site in northern Papua below about 500 m, though some of the larger-bodied, commonly-hunted species (in particular Dendrolagus inustus and Spilocuscus rufoniger) are often locally absent today in areas of higher human population density. Several additional mammals are widely distributed in northern New Guinea, but are excluded from this list for various reasons. Pteropus conspicillatus, though widespread in the northern lowlands, is generally restricted to near-coastal regions and adjacent islands, and rarely extends to forests far inland. Other species may well be widespread in the northern lowlands, but are probably uncommon, as relatively few vouchered records are available. Examples include Melomys leucogaster, previously thought to be restricted to southern New Guinea, but now recorded

................. 16157$

CH43

02-08-07 10:48:34

PS

PAGE 728

Taxonomic and Geographic Overview of the Mammals / 729

both from Jayapura and the Mamberamo Basin (Singadan and Patisellano 2002; Musser and Carleton 2005); Paramelomys moncktoni, the northern range of which is poorly defined at present (Flannery 1995a; Menzies 1996a; Musser and Carleton 2005); and probably Thylogale browni, recorded in Papua to date only from the Cyclops Range and from hill and mountain forests on Yapen (Flannery 1995a), though historically widespread in the northern lowlands of PNG. The apparent patchiness of this last species’ distribution again probably reflects human hunting pressure, with remaining populations being centered on remote upland forest areas as a result. An additional assemblage of species that is widespread (if uncommon) in northern New Guinea below 1,000 m includes the marsupials Peroryctes raffrayana and Distoechurus pennatus and the rodents Pogonomelomys mayeri, Uromys nero, Lorentzimys nouhuysi, and Xenuromys barbatus. Though most of these species have been collected on at least one occasion below 100 m, in the northern lowlands they usually occur in inland foothill forests, and are infrequently (if ever) encountered in coastal or expansive flat lowland forests below about 200–400 m. The tiny murine Microhydromys richardsoni is another very rarely collected inhabitant of hill and lower montane forests in New Guinea (between 500 and 1,500 m). Though recorded only once in Papua (at the Idenburg River along the northern margin of the Central Cordillera), it is likely more widespread in the northern lowlands than currently realized. Overall, no mammal species is endemic to the northern lowlands of Papua, but four mammals (Echymipera clara, Dorcopsis hageni, Mammelomys rattoides, and Pogonomelomys mayeri) occur only in lowland and lower montane forests north of the Central Cordillera in both Papua and PNG (i.e., from Yapen and the eastern shore of Cenderawasih Bay in the west to PNG’s Huon Peninsula in the east). Rising out of the expansive lowlands of northern New Guinea are a number of isolated mountain ranges, including the Van Rees, Foja, and Cyclops ranges in northern Papua and the Torricelli, Bewani, Prince Alexander, and Adelbert ranges in northern PNG. During cooler episodes in the Late Pleistocene, when sea levels were depressed, New Guinea’s montane forests extended to lower elevations than they do today, and may have been contiguous (or at least considerably more expansive) across some of these now isolated ranges. Mammal assemblages from two of Papua’s north coast ranges (the Van Rees and Foja ranges) remain almost entirely unknown at present, but the Cyclops Range has been the subject of moderate historical survey efforts. Several mammal species from the Cyclops’ montane forests (above about 800 m) are relatively widespread in montane habitats throughout New Guinea (e.g., Pseudochirops albertisii, Parahydromys asper, Mammelomys lanosus, and Paramelomys cf. rubex), extending elsewhere to the Central Dividing Ranges, the mountains of the Vogelkop Peninsula, or both. Though unrecorded as yet from the northern ranges of Papua, two other widespread montane mammals (Pseudochirulus forbesi and Nyctimene cyclotis) occur in the adjacent north coast ranges of PNG, and in the future may be found to occur in the Van Rees and Fojas if not in the Cyclops. Though most of these typically montane species are

................. 16157$

CH43

02-08-07 10:48:35

PS

PAGE 729

730 / kristofer m . helgen

occasionally encountered in foothill forests in the northern lowlands (down to about 300–500 m), all are ultimately reliant on montane forests. In addition to these more widespread montane species, the north coast ranges also support a considerable number of endemic mammal species, generally restricted to higher elevations (Helgen 2005). Three of these restricted-range species have been recorded from Papua to date: the dwarf echidna Zaglossus attenboroughi, known as a living animal only from the Cyclops (Flannery and Groves 1998); the tree kangaroo Dendrolagus pulcherrimus, recorded only from the Fojas and Torricellis (Flannery 1993; Helgen 2005); and the murine Paraleptomys rufilatus, endemic to the Cyclops, Torricelli, and Bewani ranges. Several additional north coast range endemic mammals (Dendrolagus scottae, ‘Microhydromys’ musseri, Hydromys ziegleri, and Hipposideros edwardshilli) are currently known only from northern PNG, but pending future survey efforts, these species may prove to extend to Papua’s littleknown northern mountain ranges as well (Helgen 2005). The oceanic islands of Biak, Supiori, and Numfoor, located in Cenderawasih (formerly Geelvink) Bay deserve particular mention in the context of northern New Guinean mammal biogeography. Like the avifauna of these islands, the entire native mammal fauna of Biak-Supiori and Numfoor shows signs of having arrived exclusively by overwater dispersal from northern New Guinea, and exhibits considerable endemism (Helgen and Flannery 2004b). Endemic mammal species include Uromys boeadii (an insular relative of U. nero), Uromys emmae (an insular relative of U. caudimaculatus), Spilocuscus wilsoni (an insular relative of S. maculatus), and Dobsonia emersa (an insular relative of D. moluccensis; Groves and Flannery 1994; Helgen and Flannery 2004b). Other distinctive near-endemics include the rat Rattus jobiensis (which extends elsewhere only to the adjacent land-bridge island of Yapen), as well as the flying fox Pteropus pohlei (which occurs only on Numfoor and Yapen) and its much smaller-bodied sister species (from Biak-Supiori and Waigeo), which remains undescribed. Finally, several other widespread mammal species (e.g., Echymipera kalubu, Petaurus breviceps) are represented on BiakSupiori by morphologically distinctive populations.

western papua: the western lowlands, vogelkop ranges, and raja ampat islands Mammal communities from lowland forests on the Vogelkop Peninsula and the adjacent Bird’s Neck region are less well inventoried than those of New Guinea’s north-central lowlands. Relevant data, drawn from scattered historical museum samples from a few lowland sites (primarily historical collections made at Sorong, Manokwari, Oransbari, and Triton Bay), and from an extensive subfossil record from two lowland sites near the Ayamaru Lakes (Aplin 1998; Aplin et al. 1999), suggest that the lowland Vogelkop fauna is similarly species-rich to that of the northern New Guinean lowlands but features a slightly different taxonomic assemblage. The original core mammal fauna of the coastal and flat western lowlands apparently comprised about 16 monotreme and marsupial species (Zaglossus

................. 16157$

CH43

02-08-07 10:48:36

PS

PAGE 730

Taxonomic and Geographic Overview of the Mammals / 731

bruijnii, Dasyurus albopunctatus, Murexia melanura, Myoictis melas, Echymipera kalubu, Echymipera rufescens, Dendrolagus inustus, Dendrolagus ursinus, Dorcopsis muelleri, Phalanger orientalis, Phalanger gymnotis, Spilocuscus maculatus, Spilocuscus rufoniger, Dactylopsila trivirgata, Petaurus breviceps, and Pseudochirulus canescens), at least six rodent species (Melomys rufescens, Paramelomys platyops, Pogonomelomys bruijnii, Pogonomys macrourus, Rattus praetor, and Uromys caudimaculatus), and 10 fruit bats (an assemblage identical to that of the northern lowlands), with Pteropus conspicillatus again an additional, common coastal inhabitant. (Only three of these core species—Z. bruijnii, D. ursinus, and P. bruijnii—are not found in the northern lowlands.) Given sufficient inventory efforts, most of the smaller-bodied species in this list could probably be expected at almost any forested site in the western lowlands (below about 500 m). However, certain larger-bodied species are locally extinct in many or most lowland areas today, with some (such as Zaglossus and Dendrolagus) now largely or entirely restricted to less accessible montane forests, and others (particularly Spilocuscus rufoniger) potentially extirpated entirely from the regional fauna (see Phalangeridae account, above). At least one species of Dorcopsulus (the montane species D. vanheurni) and a tree kangaroo related to Dendrolagus goodfellowi (probably D. cf. pulcherrimus) are additionally present in the Holocene subfossil fauna of the Vogelkop, but appear to be extinct in the modern Vogelkop fauna (Aplin et al. 1999). These perspectives drawn from the fossil record are especially important because they reveal that inferences drawn only from modern distributional data can lead to spurious ecological and biogeographic conclusions. This is especially true with regard to the larger mammals of New Guinea, whose distributions are presumably most highly impacted by past and current patterns of human hunting. Of the six additional species listed earlier as widespread inhabitants of foothill forest in the northern lowlands, two are also present in the modern fauna of the Vogelkop (Peroryctes raffrayana and Distoechurus pennatus), one is recorded in the subfossil but not yet from the modern fauna of the region (Xenuromys barbatus), and three (Lorentzimys nouhuysii, Pogonomelomys mayeri, and Uromys nero) are entirely unrecorded from the Vogelkop (see Aplin et al. 1999). Given that only a small amount of mammal survey work has commenced in the Vogelkop lowlands, it is too soon to say whether the members of the last group constitute legitimate absences, though I strongly suspect that at least Lorentzimys and U. nero will be shown to occur in the region. Additional species widespread in the northern lowlands that are unrecorded to date from the Vogelkop include Echymipera clara, Hydromys chrysogaster, and Mammelomys rattoides. Two of these species (E. clara and M. rattoides) are unrecorded in both the historical and subfossil faunas of the Vogelkop, and are probably truly absent from the western fauna. H. chrysogaster, however, is recorded both in the Vogelkop subfossil fauna and from islands to the west of the Vogelkop, such as Waigeo (Table 4.10.2), and there can be no doubt that it is present in the Vogelkop today. Another widespread and common species of the New Guinea lowlands, Uromys caudimaculatus, is known from the modern Vogelkop fauna only by trophy jaws collected from native hunters. The absence of

................. 16157$

CH43

02-08-07 10:48:36

PS

PAGE 731

732 / kristofer m . helgen

records for these species in the modern regional faunas reflects the paucity of effort invested in modern mammal collecting in the Vogelkop lowlands to date. In contrast to the still little-known Vogelkop lowlands, the mammals of the Vogelkop’s Arfak Mountains are relatively well known, thanks to extensive historical and recent inventory efforts (see Helgen and Flannery 2004a). The Arfaks support a diverse montane mammal fauna and are rich in montane endemics. Species globally restricted to forests above 700 m in the Arfaks include Phascolosorex dorsalis, Microperoryctes aplini, Pseudochirops coronatus, Pseudochirulus schlegeli, Rattus arfakiensis, and unnamed species of Leptomys and Mallomys (Tables 4.10.1–2; Appendix 4.10.1). All of these endemic species have either close phylogenetic relatives or close ecological vicars in montane forests along the Central Cordillera. In addition to these many endemics, the Arfaks also support a group of montaneadapted species that are more widespread throughout the mountains of New Guinea, particularly along the Central Cordillera (e.g., Phascolosorex doriae, Microperoryctes longicauda, Cercartetus caudatus, Dactylonax palpator, Hyomys dammermani, Parahydromys asper, Paramelomys mollis, Paramelomys rubex, Uromys cf. anak). Notably, this last species is recorded here from the Arfaks for the first time based on trophy jaws collected by T. Flannery, deposited at the Australian Museum in Sydney (cf. Tate 1951; Flannery 1995a). However, many other mammal species that are widespread throughout the length of the Central Cordillera are absent in the Arfaks, often without being replaced by a similar species (e.g., Anisomys imitator, Coccymys ruemmleri, Macruromys spp.). One species from the Arfaks and Tamraus (Pseudochirops albertisii) is absent from the Central Cordillera but shared with the Weyland Range and North Coast Ranges, a link echoed by the presence of Dendrolagus cf. pulcherrimus in the late Quaternary record of the Vogelkop (today known only from the Torricelli and Foja Ranges; see above). As noted above, several additional species (Zaglossus bruijnii and Dendrolagus ursinus) are also largely restricted to these montane forests today, although they probably had much wider elevational ranges prior to recent decades. Finally, two other marsupial species currently known only from Late Quaternary remains in the Vogelkop (the small possums Dactylopsila kambuayai and ‘‘Petauroides’’ ayamaruensis) are likely to survive undiscovered today in the Vogelkop’s mountain forests (see Aplin et al. 1999). Both of these intriguing animals should be especially sought in future biological surveys in the region. With renewed survey efforts, many of the Arfak Mountains’ seemingly unique mammal species (as well as additional, currently unknown mammals) may prove to occur in other mountain ranges in the greater Vogelkop region—especially in the expansive nearby Tamrau ranges, but also the Fakfak, Kumawa, and Wandammen Ranges in the Bird’s Neck region, all of which remain poorly known mammalogically. The Arfaks themselves, though better surveyed than other Vogelkop ranges, probably also support many other species that remain currently undetected. I suspect that these include small amphibious murines and tiny terrestrial moss mice, both of which are present throughout other montane areas of New Guinea but can be challenging to collect without pitfall techniques, which have

................. 16157$

CH43

02-08-07 10:48:36

PS

PAGE 732

Taxonomic and Geographic Overview of the Mammals / 733

only rarely been used to collect mammals in New Guinea (Flannery 1995a; Helgen 2005). A final important area to consider in western Papua is the Raja Ampat Islands, a group of western islands including the large islands of Misool, Salawati, Batanta, Waigeo, and the small islands of Kofiau and Gag, among others. With the arguable exception of Salawati, the mammal faunas of these islands are rather incompletely known (cf. Flannery 1995b; Meinig 2002), but some general comments can be made about their content. Misool and Salawati are land-bridge islands that support a depauperate subset of the same species found in the western lowlands of mainland New Guinea, and no endemics. Batanta and Waigeo are oceanic islands that in the late Pleistocene were united as a single, larger landmass. Like Salawati and Misool, these islands primarily support a reduced set of species that characterize the western lowlands, though at least one endemic species (the distinctive phalangerid Spilocuscus papuensis) is also present on these islands. An undescribed species of Pteropus (allied to P. pohlei of Numfoor and Yapen) is an additional Waigeo near-endemic, present elsewhere only on Biak-Supiori (Helgen, unpublished data). The fruit bat Dobsonia beauforti (a close ally of the Moluccan species D. crenulata and D. viridis) is another near-endemic species of the Raja Ampats (records are from Waigeo, Batanta, Salawati, Misool, and Gag), extending elsewhere only to Biak-Supiori and the North Moluccan island of Gebe. The oceanic islands of Kofiau and Gag, though included within the political boundaries of Papua, show a greater faunal resemblance to the Moluccas than to other parts of Papua. For example, the Moluccan flying foxes Pteropus chrysoproctus and P. personatus occur on Kofiau and Gag, respectively (Flannery 1995b; Maryanto and Kitchener 1999). All of the Raja Ampat islands, especially Waigeo, Batanta, and Kofiau (which have the highest potential among the Raja Ampats for supporting undiscovered mammal species) remain important priorities for biological inventory work.

southern papua: the southern lowlands and the trans-fly region The southern lowlands of Papua are bounded by the Mimika River to the west, the Central Cordillera to the north, and the Papua New Guinea border to the east. Faunistically, the southern lowlands can be divided into at least two distinctive subregions, divided more or less by the course of the Digul River. North of the Digul, lowland and swamp forests predominate; south of the Digul is the TransFly region, characterized by a highly seasonal complex mosaic landscape of extensive open habitats, such as savanna and woodland, interspersed with more closed habitats, especially monsoon and gallery forest. The mammal fauna of this region is recorded primarily by small collections. Specimens from the Mimika, Setakwa, and Utakwa river drainages are at the Natural History Museum (London); a larger collection from the vicinity of the Lorentz ( Noord) River is at the National Museum of Natural History (Leiden); and some scattered material from the Merauke area of the western Trans-Fly is housed in three collections (see below). This regional fauna is less well inventoried than

................. 16157$

CH43

02-08-07 10:48:37

PS

PAGE 733

734 / kristofer m . helgen

most areas of Papua, and as a result it is too early to fully identify a core complement of species present in most areas. Nevertheless, a number of marsupials (Myoictis melas, Echymipera kalubu, Echymipera rufescens, Dorcopsis muelleri, Phalanger mimicus, Phalanger gymnotis, Spilocuscus maculatus, Dactylopsila trivirgata, and Petaurus breviceps) and rodents (Hydromys chrysogaster, Melomys rufescens, Paramelomys lorentzii, and Rattus leucopus) are clearly widespread in western New Guinea both in hill forests along the southern margin of the Cordillera and in lowland forests north of the Trans-Fly. The same complement of lowland fruit bats found in the northern and western lowlands are also found in the southern lowlands (see above). A nearly identical complement of species also occurs in the Trans-Fly, except that Phalanger gymnotis is apparently absent, Dorcopsis muelleri is replaced by D. luctuosa, and Myoictis melas is replaced by M. wallacei. A nearendemic mammal species of Papua’s southern lowland rainforests is Uromys scaphax (see Appendix 4.10.1), which is recorded from the Mimika, Setakwa, and Lorentz river drainages, and is known otherwise only from the Strickland River drainage across the PNG border. Other species potentially unique to the southern lowlands of New Guinea (both Papua and PNG) are the bats Nyctimene draconilla and Hipposideros wollastoni (see Table 4.10.4, Appendix 4.10.1; but see also Bonaccorso 1998). Helgen and Oliver (2004) recently reviewed the mammal fauna of the TransFly region. Apart from the Bronze Quoll (Dasyrurus spartacus), which is probably conspecific with the Australian form D. geoffroyi (see Dasyuridae account above), the Trans-Fly has no endemic mammals. The Trans-Fly is faunistically notable mainly for supporting many species known otherwise only from Australia, such that the region is often thought of as a biotic extension of Australia (Schodde and Calaby 1977; Norris and Musser 2001; Chapter 5.12). Mammal species shared exclusively between the Trans-Fly and Australia include the dunnart Sminthopsis archeri, the macropodids Lagorchestes conspicillatus and Thylogale stigmatica; the rodents Conilurus penicillatus, Pseudomys delicatulus, and Xeromys myoides; and the fruit bats Nyctimene robinsoni and Pteropus scapulatus (Helgen and Oliver 2004). As noted above, Dasyurus [geoffroyi] spartacus probably falls into this category as well. With the exception of the last species, however, all of these mammals are so far recorded only from the eastern side of the Trans-Fly region (i.e., in PNG), but they all probably extend into contiguous habitats in Papua (see Table 4.10.5). Mammals recorded to date from the Papua side of the Trans-Fly comprise mostly widely distributed species (the monotreme Tachyglossus aculeatus, the marsupials Dactylopsila trivirgata, Dorcopsis luctuosa, Echymipera kalubu, E. rufescens, Isoodon macrourus, Macropus agilis, Myoictis wallacei, Petaurus breviceps, Sminthopsis virginiae, Spilocuscus maculatus, and Thylogale brunii; the rodents Melomys muscalis, Melomys rufescens, Paramelomys lorentzii, Paramelomys moncktoni, Pogonomelomys brassi, Pogonomys cf. macrourus [i.e. P. mollipilosus; see Appendix 4.10.1], Rattus leucopus, Rattus sordidus; and the fruit bats Dobsonia moluccensis, Pteropus macrotis, and Syconycteris australis). Essentially all vouchered records of mammals in the western Trans-Fly derive from the Merauke area, and are depos-

................. 16157$

CH43

02-08-07 10:48:37

PS

PAGE 734

Taxonomic and Geographic Overview of the Mammals / 735

ited today at the Naturalis Museum (Leiden), the Museum Zoologense Bogoriense (Cibinong, Indonesia), and the American Museum of Natural History (New York).

central papua: the central cordillera (star, snow, and weyland mts) The Central Cordillera is the montane backbone of the island of New Guinea, made up of many high, contiguous mountain ranges that span much of New Guinea’s east-west axis and split the island into isolated southern and northern lowland regions. The Cordillera extends from the Maneau Range of Milne Bay Province (PNG) in the far southeast to Papua’s Weyland Range in the west. For comparisons of mammalian zoogeography, the western body of the Cordillera (Papua’s portion) can be usefully divided three main zoogeographic divisions: the Star Mountains region, the Snow Mountains region, and the Weyland Range. The Star Mountains region straddles the border of Papua and PNG, and includes Papua’s Star Mountains as well as the Victor Emanuel and Hindenburg ranges of far western PNG (cf. Flannery and Seri 1990; Chapter 1.1). This region supports an extremely diverse montane mammal community. Based on extensive field survey efforts and studies in museums worldwide, Morren (1989) and Flannery and Seri (1990) were able to record 62 mammal species from forests and grasslands above 1,500 m on the PNG side of the Snow Mountains. (The identity of a few of these species has since been reassessed; e.g., records of Neohydromys fuscus and Hyomys goliath from the area actually represent Pseudohydromys occidentalis and Hyomys dammermani, respectively.) My studies of specimens deposited at the Bishop Museum (Honolulu), the National Museum of Natural History (Leiden), and the Papua New Guinea National Museum (Port Moresby) reveal that a number of additional species unrecorded by Morren (1989) or Flannery and Seri (1990) also inhabit high altitude forests in this region. These include Murexia habbema, Murexia wilhelmina, Neophascogale lorentzii, Phascolosorex brevicaudata, Mallomys istapantap, Parahydromys asper, an undescribed species of Microperoryctes, and a new genus and species of moss mouse (misidentified as Pseudohydromys occidentalis by Flannery and Seri 1990). The great majority of mammal species present in montane forests in the Star Mountains are widespread along the Central Cordillera. However, a handful seem to be shared exclusively with the expansive Snow Mountains region to the west (e.g., Murexia habbema [excluding hageni; see Appendix 4.10.1], Dendrolagus stellarum, Paraleptomys wilhelmina, and Pseudohydromys occidentalis) or with the Snow and Weyland mountains (Dactylopsila megalura). Several mammal species are known only from the Star Mountains region (e.g., Phalanger matanim, Pogonomys championi, and the unnamed moss mouse), but none of these is yet recorded from the Papua side of the border. Most mammal specimens collected to date from the Star Mountains of Papua (generally originating from the Sibil Valley; see Tables 4.10.1, 2) are held in the Naturalis Museum, Leiden. The heavily populated Baliem Valley forms the boundary between the Star and

................. 16157$

CH43

02-08-07 10:48:37

PS

PAGE 735

736 / kristofer m . helgen

Snow mountain regions. Though an important zoogeographic divide and a potentially important area of interchange, relatively little is known of the mammals of the Baliem area. Preliminary studies of mammal collections from the Baliem Valley and immediate vicinity (deposited at the Swedish Museum of Natural History, Stockholm, and the National Museum of Natural History, Leiden) have revealed unexpected discoveries. A vouchered skin from the Swart Valley (an extension of the Baliem) demonstrates that Crossomys moncktoni (previously unrecorded from Papua; see Appendix 4.10.1) extends much farther west than previously realized (cf. Musser and Carleton 2005). Further, several specimens of Pseudochirulus from Bokondini, previously identified as P. caroli (Flannery 1995a), may well represent an unnamed species (Hoogenboezem and Helgen, unpublished data). West of the Baliem Valley, the expansive Snow Mountains make up the western body of the Central Cordillera. The Weyland Range, forming the far western edge of the Cordillera, is often included in the definition of this larger Snow Mountains region (e.g., Beehler et al. 1986), but because the Weyland mammal fauna is highly distinctive, I treat it as a separate zoogeographic unit below. For the purposes of this review, the Snow Mountains are thus considered to extend from the Baliem Valley in the east to the Paniai Lakes in the west. The mammal fauna of the Snow Mountains region is more completely cataloged than that of any other area of Papua, and is extremely species rich. Especially important representative collections were made by Dutch, American, and British collectors during the first four decades of the twentieth century, particularly on the northern slopes of the Cordillera along the Idenburg River, in the neighborhood of Lake Habbema and Mt Trikora ( Mt Wilhelmina), on the southern slopes of the Cordillera along the Lorentz and Utakwa rivers, and around the Paniai ( Wissel) Lakes (e.g., see Wollaston 1914; Brass 1941; Tate 1951; Musser and Piik 1982; Flannery 1995a). More recent collections of both modern specimens and subfossil remains in the vicinities of Tembagapura, Kwiyawagi, and Mt Jaya have yielded many important newer discoveries to complement these earlier efforts (Hope 1981; Flannery 1995a, 1999; Flannery et al. 1995, 1996). Like the Star Mountains, the Snow Mountains are home to many species that are relatively widespread in montane forests along the Central Cordillera (Tables 4.10.1, 2). A few additional species extend elsewhere only to the mountains of the Vogelkop (e.g., Phascolosorex doriae, Rattus unicolor). The Snow Mountains also support many endemic species (Tables 4.10.1,2; Figure 4.10.8). Most of these are restricted to the region’s relatively extensive subalpine and alpine habitats (e.g., the rodents Mallomys gunung, Coccymys albidens, Rattus omichlodes, and Rattus richardsoni; and the wallaby Thylogale christenseni—a Snow Mountains endemic known as yet only from late Quaternary remains; Hope 1981) or to both upper montane forest and subalpine contexts (e.g., the tree kangaroo Dendrolagus mbaiso, the amphibious murine Baiyankamys habbema, undescribed species of Pogonomys and Microperoryctes; see Appendix 4.10.1). Of two remaining endemics, one is widespread in open areas at various altitudes above 1,500 m (the small rat Melomys frigicola) while another (the amphibious murine Hydromys hussoni)

................. 16157$

CH43

02-08-07 10:48:38

PS

PAGE 736

Taxonomic and Geographic Overview of the Mammals / 737

is known only from the Paniai Lakes (ca 1,800 m; Musser and Piik 1982; Helgen 2005). Despite considerable taxonomic attention paid to Snow Mountains mammals by early- and mid-twentieth-century taxonomists, many of these regional endemics were described only relatively recently (Tables 4.10.1, 2). Though a few of these endemic species appear to have immediately related sister taxa in similar montane areas elsewhere in New Guinea (e.g., Baiyankamys habbema, Microperoryctes sp., Pogonomys sp.), most of the Snow Mountains’ endemic mammals have no clearly identifiable vicars elsewhere. The Weyland Range lies at the far western margin of New Guinea’s expansive Central Dividing Ranges. This relatively small montane area is treated separately here because, based on current evidence, its mammal fauna appears highly distinctive in comparison with the rest of the Snow Mountains region. Like the Snows, the Weylands have been reasonably well studied. Extensive collections in the range were made in the 1930s in the Gebroeders area by F. Shaw Mayer (mainly deposited in the Natural History Museum, London); on Mount Kunupi, Mount Sumuri, and the Menoo Valley by Georg Stein (of the Zoology Museum in Berlin); and by A. E. Pratt (deposited in the Natural History Museum, London). Though most species recorded from the Weylands are relatively widespread along the Central Cordillera, the range supports at least three endemic mammals: the bandicoot Microperoryctes murina and the murine rodents Macruromys elegans and Paramelomys steini (Helgen and Flannery 2004a; Menzies 1996a; Musser and Carleton 2005). All three appear to be geographically relict forms, each of which is sister to much more widespread taxa. Microperoryctes murina is the sole species in the subgenus Microperoryctes, thought to be the sister taxon to the striped bandicoots (subgenus Ornoryctes), which occur throughout the entire length of the Central Cordillera and in the Arfak Mountains (Helgen and Flannery 2004a). Macruromys elegans is the sister species to M. major, which is widely distributed from the Snow Mountains to the Maneau Range in southeastern PNG, as well as in the Huon Peninsula. Finally, Paramelomys steini may be the sister species to P. rubex, a species widely distributed in montane areas throughout the island of New Guinea, including the Weyland Range, though the systematics of rubex (which currently incorporates a wide array of named forms) remain poorly understood despite a recent revision (Menzies 1996a). Interestingly, all three of these Weyland endemics are known only from the peaks of Mount Sumuri or Mount Kunupi; none having been collected in the nearby Gebroeders area. The Weyland Range is unusual in that it lacks certain species typical of most other montane areas in New Guinea, including echidnas. Such absences cannot be easily dismissed as an artifact of incomplete survey efforts, as the mammal fauna of the Weylands is one of the best inventoried in all of New Guinea. Absence of echidnas in the Weylands could result from an historical or prehistoric extirpation; alternately, it might be a natural absence that provides an insight into diversification of echidnas in western New Guinea. Zaglossus bartoni (a species of the Central Cordillera) occurs in the Paniai Lakes area to the immediate east of the Weylands, while Zaglossus bruijnii (typical of the Vogelkop and Bird’s Neck) is found to the

................. 16157$

CH43

02-08-07 10:48:38

PS

PAGE 737

738 / kristofer m . helgen

immediate southwest, in the Charles Louis Range (Flannery and Groves 1998). Is the Weyland Range a ‘‘drop-out’’ area that might have facilitated speciation of Zaglossus in allopatry? Or is it perhaps an area of former overlap or abutment between two closely related species? Similar questions apply to tree kangaroos, which are unknown from the Weylands but recorded in the historical fauna of the Paniai Lakes area (D. stellarum) and from the lowlands and mountains of the Bird’s Neck to the west (D. inustus, D. ursinus, D. mayri; Flannery et al. 1996). In summary, the Weyland Range poses several biogeographic riddles that encourage renewed research into the distributions and relationships of mammals at the western boundary of the Central Cordillera. The geographic coincidence of phylogenetically-important relict species, along with the Range’s seeming microgeographic faunal heterogeneity (i.e., potential differences in species composition between the Gebroeders and Kunupi/Sumuri), and conspicuous lack of certain widespread mammal lineages, definitively highlights the role that the Weyland Range will play in future efforts to unravel mammalian zoogeographic patterns in western New Guinea.

Acknowledgments I thank the editors of this volume for their friendship and assistance and Tim Flannery, Ken Aplin, and Guy Musser for their continuing guidance and collaboration. For access to specimens and other assistance, I am grateful to curators and staff at the American Museum of Natural History in New York (especially Darrin Lunde), the Naturalis Museum, Leiden (especially Chris Smeenk), the Australian Museum in Sydney (especially Sandy Ingleby and Tish Ennis), the Bishop Museum in Honolulu (especially Carla Kishinami and Allen Allison), the Natural History Museum, London (especially Paula Jenkins and Daphne Hills), the Museum fu¨r Naturkunde at Humboldt University in Berlin (especially Peter Giere and Rob Asher), the Museo Civico di Storia Naturale ‘‘Giacomo Doria’’ in Genoa (especially Guiliano Doria), the Swedish Museum of Natural History in Stockholm (especially Olavi Gro¨nwall), the Field Museum in Chicago (especially Bill Stanley and Larry Heaney), the Muse´um National d’Histoire Naturelle in Paris (especially Jacques Cuisin), and the Museum Zoologicum Bogoriense in Cibinong, Indonesia (especially Ibnu Maryanto). I also thank the U.S. National Science Foundation and the Smithsonian Institution for ongoing research support. Any errors are mine alone.

Literature Cited Aplin, K.P. 1998. Vertebrate zoogeography of the Bird’s Head of Irian Jaya, Indonesia. Pp. 803–890 in Miedema, J., C. Ode´, and R.A.C. Dam (eds.) Perspectives on the Bird’s Head of Irian Jaya, Indonesia. Rodopi, Amsterdam. Aplin, K.P., J.M. Pasveer, and W.E. Boles. 1999. Late Quaternary vertebrates from the Bird’s Head Peninsula, Irian Jaya, Indonesia, including descriptions of two previously

................. 16157$

CH43

02-08-07 10:48:38

PS

PAGE 738

Taxonomic and Geographic Overview of the Mammals / 739 unknown marsupial species. Records of the Western Australian Museum, Supplement 57: 351–387. Armstrong, L.A., C. Krajewski, and M. Westermann. 1998. Phylogeny of the dasyurid marsupial genus Antechinus based on cytochrome b, 12S rRNA, and protamine P1 genes. Journal of Mammalogy 81: 1008–1024. Beehler, B.M., T.K. Pratt, and D.A. Zimmermann. 1986. Birds of New Guinea. Princeton University Press, Princeton, New Jersey. Bergmans, W. 2001. Notes on distribution and taxonomy of Australasian bats. I. Pteropodinae and Nyctimeninae (Mammalia, Megachiroptera, Pteropodidae). Beaufortia 51: 119–152. Bonaccorso, F.J. 1998. Bats of Papua New Guinea. Conservation International Tropical Field Guide Series, Washington, D.C. Bowyer, J.C., G.R. Newell, C.J. Metcalfe, and M.B.D. Eldridge. 2003. Tree-kangaroos Dendrolagus in Australia: are D. lumholtzi and D. bennettianus sister taxa? Australian Zoologist 32: 207–213. Brass, L.J. 1941. The 1938–39 expedition to the Snow Mountains, Netherlands New Guinea. Journal of the Arnold Arboretum 22: 271–295, 297–342. Brisbin, I.L., Jr., R.P. Coppinger, M.H. Feinstein, and S.N. Austad. 1994. The New Guinea singing dog: taxonomy, captive studies, and conservation priorities. Science in New Guinea 20: 27–38. Colgan, D.J., and T.F. Flannery. 1992. Biochemical systematic studies in the genus Petaurus (Marsupialia: Petauridae). Australian Journal of Zoology 40: 245–256. Corbet, G.B., and J.E. Hill. 1992. The Mammals of the Indomalayan Region: A Systematic Review. Oxford University Press, Oxford. Da´valos, L.M. 2004. Phylogeny and biogeography of Caribbean mammals. Biological Journal of the Linnean Society 81: 373–394. Donnellan, S.C., T.B. Reardon, and T.F. Flannery. 1995. Electrophoretic resolution of species boundaries in tube-nosed bats (Chiroptera: Pteropodidae) in Australia and New Guinea. Australian Mammalogy 18: 61–70. ¨ ber artliche Abgrenzung und innerartliche Ausformung bei Phalanger Feiler, A. 1978. U maculatus. Zoologische Abhandlungen Staatliches Museum fu¨r Tierkunde in Dresden 35: 1–30. Filewood, L.W. 1983. The possible presence in New Guinea of the Ghost Bat (Macroderma gigas: Chiroptera, Megadermatidae). Australian Mammalogy 6: 35–36. Flannery, T.F. 1990. Mammals of New Guinea. Robert Brown and Associates, Carina, Queensland. Flannery, T.F. 1992. New Pleistocene marsupials (Macropodidae, Diprotodontidae) from subalpine habitats in Irian Jaya. Alcheringa 16: 321–331. Flannery, T.F. 1993. Taxonomy of Dendrolagus goodfellowi (Macropodidae: Marsupialia) with description of a new subspecies. Records of the Australian Museum 45: 33–42. Flannery, T.F. 1994a. The fossil land mammal record of New Guinea: a review. Science in New Guinea 20: 39–48. Flannery, T.F. 1994b. Possums of the World: A Monograph of the Phalangeroidea. GEO Productions, Sydney. Flannery, T.F. 1995a. Mammals of New Guinea, rev. ed. Cornell University Press, Ithaca, New York. Flannery, T.F. 1995b. Mammals of the South-West Pacific and Moluccan Islands. Cornell University Press, Ithaca, New York. Flannery, T.F. 1996. Mammalian zoogeography of New Guinea and the southwestern

................. 16157$

CH43

02-08-07 10:48:39

PS

PAGE 739

740 / kristofer m . helgen Pacific. Pp. 399–406 in Keast, A., and S.E. Miller (eds.) The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam. Flannery, T.F. 1999. The Pleistocene mammal fauna of Kelangurr Cave, central montane Irian Jaya, Indonesia. Records of the Western Australian Museum, Supplement 57: 341–350. Flannery, T.F., K.P. Aplin, C.P. Groves, and M. Adams. 1989. Revision of the New Guinean genus Mallomys (Muridae: Rodentia), with descriptions of two new species from subalpine habitats. Records of the Australian Museum 41: 83–105. Flannery, T.F., Boeadi, and A.L. Szalay. 1995. A new tree-kangaroo (Dendrolagus: Marsupialia) from Irian Jaya, Indonesia, with notes on ethnography and the evolution of tree-kangaroos. Mammalia 59: 65–84. Flannery, T.F., and D.J. Colgan. 1993. A new species and two new subspecies of Hipposideros (Chiroptera) from Western Papua New Guinea. Records of the Australian Museum 45: 43–57. Flannery, T.F., and C.P. Groves. 1998. A revision of the genus Zaglossus (Monotremata, Tachyglossidae), with description of new species and subspecies. Mammalia 62: 367–396. Flannery, T.F., R. Martin, and A. Szalay. 1996. Tree Kangaroos: A Curious Natural History. Reed Books, Melbourne. Flannery, T.F., and L. Seri. 1990. The mammals of southern West Sepik Province, Papua New Guinea: their distribution, abundance, human use, and zoogeography. Records of the Australian Museum 42: 173–208. Goodman, S.M., and J.P. Benstead. 2005. Updated estimates of biotic diversity and endemism for Madagascar. Oryx 39: 73–77. Groves, C.P. 2005a. Order Peramelemorphia. In Wilson, D.E., and D.R. Reeder (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed. Johns Hopkins University Press, Baltimore, Maryland. Groves, C.P. 2005b. Order Diprotodontia. In Wilson, D.E., and D.R. Reeder (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed. Johns Hopkins University Press, Baltimore, Maryland. Groves, C.P., and T.F. Flannery. 1990. Revision of the families and genera of bandicoots. Pp. 1–11 in Seebeck, J.H., P.R. Brown, R.L. Wallis, and C.M. Kemper (eds.) Bandicoots and Bilbies. Surrey Beatty & Sons, Norton, New South Wales. Groves, C.P., and T.F. Flannery. 1994. A revision of the genus Uromys Peters, 1867 (Muridae: Mammalia) with descriptions of two new species. Records of the Australian Museum 46: 145–169. Heads, M. 2002. Regional patterns of biodiversity in New Guinea animals. Journal of Biogeography 29: 285–294. Heaney, L.R., D.S. Balete, M.L. Dolar, A.C. Alcala, A.T.L. Dans, P.C. Gonzales, N.R. Ingle, M.V. Lepiten, W.L.R. Oliver, P.S. Ong, E.A. Rickart, B.R. Tabaranza Jr., and R.C.B. Utzurrum. 1998. A synopsis of the mammalian fauna of the Philippine Islands. Fieldiana Zoology n.s. 88: 1–61. Hedemark, M., S. Hamilton, and W. Takeuchi. 1997. Report on the First Bismarck-Ramu Biological Survey with Sociological and Logistical Comments. Department of Environment and Conservation, Papua New Guinea, Port Moresby. Heim de Balsac, H., and M.-H. Heim de Balsac. 1956. Suncus murinus (L.), a` la Re´union et en Nouvelle-Guine´e. Le Naturaliste Malgache 8: 143–147. Helgen, K.M. 2003. The feather-tailed glider (Acrobates pygmaeus) in New Guinea. Treubia 33: 107–111.

................. 16157$

CH43

02-08-07 10:48:39

PS

PAGE 740

Taxonomic and Geographic Overview of the Mammals / 741 Helgen, K.M. 2005. The amphibious murines of New Guinea (Rodentia, Muridae): the generic status of Baiyankamys and description of a new species of Hydromys. Zootaxa 913: 1–20. Helgen, K.M., and T.F. Flannery. 2004a. A new species of bandicoot, Microperoryctes aplini, from western New Guinea. Journal of Zoology (London) 264: 117–124. Helgen, K.M., and T.F. Flannery. 2004b. Notes on the phalangerid marsupial genus Spilocuscus, with description of a new species from Papua. Journal of Mammalogy 85: 825–833. Helgen, K.M., and P.M. Oliver. 2004. A review of the mammal fauna of the Trans-Fly ecoregion. Report to the WWF South Pacific Program, Papua New Guinea. Hide, R. 2004. Pig Husbandry in New Guinea: A Literature Review and Bibliography. Australian Centre for Agricultural Research, Canberra. Hill, J.E. 1990. A memoir and bibliography of Michael Rogers Oldfield Thomas, F.R.S. Bulletin of the British Museum of Natural History (Historical Series) 18: 25–113. Hope, J.H. 1981. A new species of Thylogale (Marsupialia: Macropodidae) from Mapala rock shelter, Jaya (Carstenz) Mountains, Irian Jaya (Western New Guinea), Indonesia. Records of the Australian Museum 33: 368–387. Kemp, N.J., and J.B. Burnett. 2003. Final report: a biodiversity risk assessment and recommendations for risk management of Long-tailed Macaques (Macaca fascicularis) in New Guinea. Indo-Pacific Conservation Alliance, Washington, D.C. Kirch, P.V. 1997. The Lapita Peoples. Blackwell, Malden, Massachusetts. Kitchener, D.J., N. Cooper, and I. Maryanto. 1995. The Myotis adversus (Chiroptera: Vespertilionidae) species complex in eastern Indonesia, Australia, Papua New Guinea and the Solomon Islands. Records of the Western Australian Museum 17: 191–212. Koler-Matznick, J., I.L. Brisbin Jr., M. Feinstein, and S. Bulmer. 2003. An updated description of the New Guinea singing dog (Canis hallstromi, Troughton 1957). Journal of Zoology (London) 261: 109–118. Krajewski, C., G.R. Moyer, J.T. Sipiorski, M.G. Fain, and M. Westerman. 2004. Molecular systematics of the enigmatic ‘‘phascolosoricine’’ marsupials of New Guinea. Australian Journal of Zoology 52: 389–415. Krajewski, C., S. Wroe, and M. Westerman. 2000. Molecular evidence for the timing of cladogenesis in dasyurid marsupials. Zoological Journal of the Linnean Society 130: 375–404. Larson, G., K. Dobney, U. Albarella, M. Fang, E. Matisoo-Smith, J. Robins, S. Lowden, H. Finlayson, T. Brand, E. Willerslev, P. Rowley-Conwy, L. Andersson, and A. Cooper. 2005. Worldwide phylogeography of wild boar reveals multiple centers of pig domestication. Science 307: 1618–1621. Laurie, E.M.O., and J.E. Hill. 1954. List of Land Mammals of New Guinea, Celebes, and Adjacent Islands, 1758–1952. British Museum (Natural History), London. Leary, T., and L. Seri. 1997. An annotated checklist of mammals recorded in the Kikori River Basin, Papua New Guinea. Science in New Guinea 23: 79–100. Leonard, J.A., R.K. Wayne, J. Wheeler, R. Valadez, S. Guillen, and C. Vila. 2002. Ancient DNA evidence for Old World origin of New World dogs. Science 298: 1613–1616. Maryanto, I., and D.J. Kitchener. 1999. Mammals of Gag Island. Treubia 31: 177–219. Meinig, H. 2002. New records of bats (Chiroptera) from Indonesian islands. Myotis 40: 59–79. Menzies, J.I. 1991. A Handbook of New Guinea Marsupials and Monotremes. Kristen Press, Madang, Papua New Guinea. Menzies, J.I. 1993. A history of mammalogy in New Guinea from 1726 to 1993. Science in New Guinea 19: 115–122.

................. 16157$

CH43

02-08-07 10:48:39

PS

PAGE 741

742 / kristofer m . helgen Menzies, J.I. 1996a. A systematic revision of Melomys (Rodentia: Muridae) of New Guinea. Australian Journal of Zoology 44: 367–426. Menzies, J.I. 1996b. Unnatural distribution of fauna in the East Malesian region. Pp. 31–38 in Kitchener, D.J., and A. Suyanto (eds.) Proceedings of the First International Conference on Eastern Indonesian-Australian Vertebrate Fauna. Western Australian Museum, Perth, Australia. Menzies, J.I., and J.C. Pernetta. 1986. A taxonomic revision of cuscuses allied to Phalanger orientalis (Marsupialia: Phalangeridae). Journal of Zoology (London) series B 1: 551–618. Morren, G.E.B., Jr. 1989. Mammals of the East Miyanmin area, Telefomin District, Papua New Guinea, with notes on folk knowledge and taxonomy. Science in New Guinea 15: 119–135. Musser, G.G., and M.D. Carleton. 1993. Family Muridae. Pp. 501–755 in Wilson, D.E., and D.R. Reeder (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 2nd ed. Smithsonian Institution Press, Washington, D.C. Musser, G.G., and M.D. Carleton. 2005. Family Muridae. In Wilson, D.E., and D.R. Reeder (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed. Johns Hopkins University Press, Baltimore, Maryland. Musser, G.G., and E. Piik. 1982. A new species of Hydromys (Muridae) from western New Guinea (Irian Jaya). Zoologische Mededelingen 56: 153–167. Norris, C.A., and G.G. Musser. 2001. Systematic revision within the Phalanger orientalis complex (Diprotodontia, Phalangeridae): a third species of lowland gray cuscus from New Guinea and Australia. American Museum Novitates 3356: 1–20. Osborne, M.J., and L. Christidis. 2002a. Systematics and biogeography of pygmy possums (Burramyidae: Cercartetus). Australian Journal of Zoology 50: 25–37. Osborne, M.J., and L. Christidis. 2002b. Molecular relationships of the cuscuses, brushtail, and scaly-tailed possums (Phalangerinae). Australian Journal of Zoology 50: 135–149. Pasveer, J.M., and K.P. Aplin. 1998. Late Pleistocene to modern vertebrate faunal succession and environmental change in lowland New Guinea: evidence from the Bird’s Head of Irian Jaya, New Guinea. Pp. 891–939 in Miedema, J., C. Ode´, and R.A.C. Dam (eds.) Perspectives on the Bird’s Head of Irian Jaya, Indonesia. Rodopi, Amsterdam. Petersen, R.L. 1991. Systematic variation in the megachiropteran tube-nosed bats Nyctimene cyclotis and N. certans. Bulletin of the American Museum of Natural History 206: 26–41. Ru¨mmler, H. 1938. Die Systematik und Verbreitung der Muriden Neuguineas. Mitteilungen Zoologischen Museum Berlin 23: 1–297. Schodde, R., and J.H. Calaby. 1972. The biogeography of the Australo-Papuan bird and mammal faunas in relation to Torres Strait. Pp. 257–300 in Walker, D. (ed.) Bridge and Barrier: The Natural and Cultural History of Torres Strait. Australian National University, Canberra. Singadan, R., and F. Patisellano. 2002. Small mammals of the Dabra area, Mamberamo River Basin, Papua, Indonesia. Pp. 89–91 in Richards, S.J., and S. Suryadi (eds.) A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. Conservation International (RAP Bulletin of Biological Assessment No. 25), Washington, D.C. Tate, G.H.H. 1947. Results of the Archbold Expeditions. No. 56. On the anatomy and classification of the Dasyuridae (Marsupialia). Bulletin of the American Museum of Natural History 88: 97–156.

................. 16157$

CH43

02-08-07 10:48:40

PS

PAGE 742

Taxonomic and Geographic Overview of the Mammals / 743 Tate, G.H.H. 1948a. Results of the Archbold Expeditions. No. 59. Studies on the anatomy and phylogeny of the Macropodidae (Marsupialia). Bulletin of the American Museum of Natural History 91: 233–352. Tate, G.H.H. 1948b. Results of the Archbold Expeditions. No. 60. Studies in the Peramelidae (Marsupialia). Bulletin of the American Museum of Natural History 92: 313–346. Tate, G.H.H. 1951. Results of the Archbold Expeditions. No. 65. The rodents of Australia and New Guinea. Bulletin of the American Museum of Natural History 97: 183–430. Taylor, J.M., J.H. Calaby, and H.M. van Deusen. 1982. A revision of the genus Rattus (Rodentia: Muridae) in the New Guinean region. Bulletin of the American Museum of Natural History 173: 177–336. Teeling, E.C., M.S. Springer, O. Madsen, P. Bates, S.J. O’Brien, and W.J. Murphy. 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307: 580–584. van Dyck, S. 1988. The Bronze quoll, Dasyurus spartacus (Marsupialia: Dasyuridae), a new species from the savannahs of Papua New Guinea. Australian Mammalogy 11: 145–156. van Dyck, S. 2002. Morphology-based revision of Murexia and Antechinus (Marsupialia: Dasyuridae). Memoirs of the Queensland Museum 48: 239–330. van Steenis-Kruseman, M.J. 1950. Malaysian plant collectors and collections. Flora Malesiana (1), Bogor and Leiden. Waithman, J. 1979. A report on a collection of mammals from southwest Papua, 1972– 1973. Australian Zoologist 20: 313–326. Wallace, A.R. 1869. The Malay Archipelago. Macmillan, London. Westerman, M., M.S. Springer, J. Dixon, and C. Krajewski. 1999. Molecular relationships of the extinct pig-footed bandicoot Chaeropus ecaudatus (Marsupialia: Perameloidea) using 12S rRNA sequences. Journal of Mammalian Evolution 6: 271–288. Wollaston, A.F.R. 1914. An expedition to Dutch New Guinea. Geographical Journal 43: 248–273. Woolley, P.A. 2005. Revision of the three-striped dasyures, genus Myoictis (Marsupialia: Dasyuridae), of New Guinea, with description of a new species. Records of the Australian Museum 57: 321–340. Ziegler, A.C. 1977. Evolution of New Guinea’s marsupial fauna in response to a forested environment. Pp. 117–138 in Stonehouse, B., and D. Gilmore (eds.) The Biology of Marsupials. Macmillan, London. Ziegler, A.C. 1982. An ecological check-list of New Guinea Recent mammals. Pp. 863–894 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea, vol. 2. Monographiae Biologicae, 42: 1–983 (2 vols).

Appendix 4.10.1: Taxonomic Notes Deviations from the taxonomy of Flannery (1995a, 1995b) or Musser and Carleton (2005) are discussed here. Marsupials and monotremes (numbering follows Table 4.10.1) 2–4. Taxonomy of Zaglossus follows Flannery and Groves (1998), a more realistic assessment of diversity than previous treatments. 4. Zaglossus bartoni (type locality Mt Victoria, PNG) as recognized by

................. 16157$

CH43

02-08-07 10:48:40

PS

PAGE 743

744 / kristofer m . helgen

7–11.

7.

9, 11.

12, 13.

16, 17.

19–27.

23. 24, 26.

29, 32.

Flannery and Groves (1998) may be a complex of related parapatric or allopatric species. The representative of this complex in Papua is Z. b. diamondi Flannery and Groves 1998. Extension of the genus Murexia to include the Antechinus-like New Guinea dasyures (sensu Flannery 1995a), as well as M. longicaudata and M. rothschildi (‘‘traditional’’ members of the genus), is supported by genetic studies (e.g., Armstrong, Krajewski, and Westerman 1998; Krajewski, Wroe, and Westerman 2000; contra van Dyck 2002), which highlight this grouping as a monophyletic assemblage distinct from other dasyurid clades. Murexia habbema (of the Snow and Star Mountains) is tentatively recognized as a species distinct from M. hageni (of east-central PNG) pending a study completed by D. Lunde (in litt.). These taxa were considered conspecific by van Dyck (2002). Based on examinations of relevant specimens, I provisionally recognize Murexia wilhelmina (widespread in the Central Dividing Ranges) as a species distinct from (and potentially syntopic with) M. melanura. These taxa were considered conspecific by van Dyck (2002). Populations traditionally identified as ‘‘Myoictis melas’’ (e.g., Flannery 1995a) represent a complex of at least four distinctive parapatric and allopatric species (P. Woolley 2005). Only M. melas (sensu stricto) and M. wallacei are recorded from Papua to date. Phascolosorex dorsalis (recorded only by the type and other specimens from the Arfak Mountains) differs in pelage coloration, body size (larger), and relative tail length (longer) compared to P. brevicaudata of the Central Cordillera (including whartoni). These allopatric taxa are ranked here as separate species pending more detailed study. Molecular data suggest that the bandicoot family ‘‘Peroryctidae,’’ as proposed by Groves and Flannery (1990), is paraphyletic; thus all New Guinean bandicoots are here placed in the family Peramelidae (Westerman et al. 1999). Helgen and Flannery (2004a) described Microperoryctes aplini as a distinctive new species endemic to the Arfak Mountains. Helgen and Flannery (2004a) argued for the specific-level separation of Microperoryctes longicauda and M. ornata, suggesting that these are morphologically distinct species that overlap geographically. My subsequent studies of museum holdings suggest that an additional unnamed species of Microperoryctes, the largest in the genus, occurs at high altitudes in the Snow Mountains. Drawing from comparisons of craniodental and external morphology as well as a relevant but limited molecular study (Bowyer et al. 2003), I provisionally recognize Dendrolagus mayri (known only

................. 16157$

CH43

02-08-07 10:48:40

PS

PAGE 744

Taxonomic and Geographic Overview of the Mammals / 745

30.

31.

42.

43. 47.

50. 53–55.

from the Wondiwoi Range on the Bird’s Neck) and D. stellarum (from the Snow and Star Mountains) as allopatric species distinct from D. dorianus of south-eastern PNG, with which they are traditionally considered conspecific (D. notatus of east-central PNG probably represents a fourth distinct species in this complex). Even with this splitting, relatively extensive DNA sequence divergence between Star and Snow Mts populations of stellarum (7.8% in cyt b haplotypes) potentially suggests that more than one species could be included within stellarum (Bowyer et al. 2003). This morphologically-isolated tree kangaroo was described as new to science by Flannery, Boeadi, and Szalay (1995); this is the ‘‘Dendrolagus sp.’’ of Flannery (1995a). Though described as a subspecies of Dendrolagus goodfellowi by Flannery (1993), D. pulcherrimus differs from goodfellowi in color pattern, in its slightly smaller size, and in its wide premolars, and has been recently ranked as a distinct allopatric species (Helgen 2005; Groves 2005). However, it is morphologically extremely close to goodfellowi and its appropriate taxonomic rank deserves further study. The record of pulcherrimus from Papua is based on a sighting by Jared Diamond in the Foja Range reported by Flannery, Martin, and Szalay (1997), which requires further confirmation but is provisionally accepted here. Morphological variation (individual, geographic, and elevational) in Phalanger gymnotis is poorly understood, and it is possible that more than one species may be involved. Various past authors have discussed how animals from higher elevations (⬎ ca 1,500 m) seem to differ from those from the lowlands (in their smaller body size, longer fur, relatively larger teeth, pale terminal tail-tip, etc.; Menzies and Pernetta 1986; Flannery and Seri 1990). Evaluating the systematic significance of these and other morphological traits requires a renewed investigation drawing from available museum samples of gymnotis (relatively common in world museums), ideally including molecular comparisons. Phalanger mimicus is recognized as distinct from P. intercastellanus following Norris and Musser (2001). ‘‘Spilocuscus maculatus,’’ as currently defined (e.g., Flannery 1994; Helgen and Flannery 2004b), probably represents a speciescomplex of parapatric and allopatric lowland species. Discerning systematic boundaries and distinctions between the various taxa included in maculatus is an important priority for future study. Spilocuscus wilsoni of Biak-Supiori was recently described by Helgen and Flannery (2004b). In my view, Dactylonax Thomas, 1910 (a monotypic genus comprising D. palpator), is morphologically far-removed from other striped

................. 16157$

CH43

02-08-07 10:48:41

PS

PAGE 745

746 / kristofer m . helgen

56.

possums (Dactylopsila spp.) and should be recognized as a separate genus. This arrangement is supported by limited molecular comparisons (Osborne and Christidis 2002b), but molecular studies with more complete sampling of petaurids are needed. More than one species may be included in Petaurus breviceps as currently recognized (see Colgan and Flannery 1992)—a complex problem that has received insufficient study. Species boundaries could almost certainly be resolved with morphometric and molecular comparisons drawing from currently available museum material and associated tissue samples. Further study of available series of biacensis from Biak-Supiori suggest that this population is not specifically distinct from breviceps (contra Flannery 1994) and differs mainly in its higher incidence of melanism.

Rodents (numbering follows Table 4.10.2) 2. Baiyankamys is recognized as a genus distinct from Hydromys following Helgen (2005). 3. Description of a new genus is being prepared for the distinctive murine Coccymys albidens (G. Musser and D. Lunde, in litt.). 5. Crossomys moncktoni is recorded here from Papua for the first time based on a specimen from the Swart Valley in the Swedish Museum of Natural History, Stockholm. 7. Hydromys hussoni of the Wissel Lakes is recognized as endemic to Papua following the description of H. ziegleri (based on a specimen from PNG formerly identified as hussoni) by Helgen (2005). 9. An undescribed species of Leptomys occurs in the Arfak Mountains (Helgen and Flannery 2004a; Musser, Helgen, and Lunde, unpublished). 10. Populations currently referred to Lorentzimys nouhuysii (Flannery 1995a; Musser and Carleton 2005) appear to represent a complex of species, the taxonomy of which requires detailed study. 13. Mallomys aroaensis is now known to occur throughout the Snow Mountains, not just in the mountains of Papua New Guinea (Helgen, unpublished; contra Flannery et al. 1989). 17. An undescribed species of Mallomys occurs in the Arfak Mountains (Helgen and Flannery 2004a). 21, 23, 24. Melomys frigicola is recognized as a species distinct from lutillus following Menzies (1996a) and Musser and Carleton (2005). Musser and Carleton (2005) provisionally applied the name Melomys burtoni to the Trans-Fly population usually referred to lutillus (implying its conspecificity with an Australian species), but the name M. muscalis (type locality ‘‘Lower Fly River’’) is conservatively used for this taxon here. Further study is needed (K. Aplin, in litt.). 29. Populations currently referred to Paraleptomys wilhelmina (Flannery

................. 16157$

CH43

02-08-07 10:48:41

PS

PAGE 746

Taxonomic and Geographic Overview of the Mammals / 747

1995a; Musser and Carleton 2005) appear to represent a complex of species, the taxonomy of which requires detailed study. 30–36. Use of Paramelomys Ru¨mmler, 1936 at generic level follows Menzies (1996a) and Musser and Carleton (2005). 34. The taxonomy of Paramelomys platyops is poorly understood. As currently recognized (Flannery 1995a, 1995b; Menzies 1996a; Musser and Carleton 2005), it probably represents a complex of related species widely distributed in lowland habitats throughout New Guinea and on adjacent islands. 35, 36. The taxonomy of Paramelomys rubex is poorly understood. As currently recognized (Menzies 1996a; Musser and Carleton 2005), it probably represents a complex of species widely distributed in montane habitats throughout New Guinea. Paramelomys steini is recognized as a species distinct from P. rubex following Menzies (1996a) and Musser and Carleton (2005). 37, 38. The allopatric (or potentially parapatric) taxa Pogonomelomys bruijnii and P. brassi differ in color and cranial size and I regard them as closely-related but distinct species pending further study. 39. Specimens currently allocated to Pogonomelomys mayeri may represent at least two distinct, geographically overlapping species. 40. Populations currently referred to Pogonomys loriae (e.g., by Flannery 1995a) represent a complex of species (see, e.g., Aplin, Pasveer, and Boles 1999), the taxonomy of which requires detailed study. 41, 42. Specimens of ‘‘Pogonomys sylvestris’’ from the Arfak Mountains reported by Rothschild and Dollman (1933) may represent macrourus, as discussed by Tate (1936), but a species similar to P. sylvetris does apparently occur in the Arfaks (T. Flannery, in litt.). Pogonomys mollipilosus of the Trans-Fly is provisionally included in the synonymy of P. macrourus here, as recently advocated by Musser and Carleton (2005), though my own recent examination of the type specimen of mollipilosus (which is a larger-toothed rat than P. macrourus) suggests that the two are not conspecific. 43. An undescribed Pogonomys species similar to P. sylvestris occurs in the Snow Mountains (G. Musser and D. Lunde, in litt.). 46, 47, 53. I agree with Musser and Carleton (2005) that Rattus arrogans and Rattus pococki should be recognized as valid species that co-occur along much of the western Central Cordillera, and that Rattus arfakiensis (represented both by the holotype and by additional specimens at the Australian Museum in Sydney) probably also represents a distinct species (all of these taxa were formerly lumped in the ‘‘Stenomys niobe’’ species-complex). However, given its collection altitude (ca 1,700 m) and published measurements (e.g., Tate, 1951), I am unconvinced that the holotype of klossi ( haymani Ellerman, 1941) is an example of arrogans; rather, it may represent

................. 16157$

CH43

02-08-07 10:48:41

PS

PAGE 747

748 / kristofer m . helgen

a third, lower-elevation species in this complex (cf. Flannery and Seri 1990: 191). Clearly a detailed taxonomic review is necessary. 48, 57. Rattus steini of west-central New Guinea (including baliemensis) is larger and differs in its mammae formula compared to geographically disjunct populations referred to as ‘‘R. steini’’ from the Star Mountains (including the Sibil Valley of Papua) and elsewhere in PNG (see Taylor et al. 1982). In my opinion these eastern populations must be regarded as representing a separate species, the name for which is R. foersteri (Ru¨mmler, 1935), with synonyms hageni and rosalinda (see Taylor, Calaby, and van Deusen 1982). 51. The name Rattus omichlodes was misspelled as ‘‘omlichodes’’ by Flannery (1995a). 57. Rattus unicolor (of montane western New Guinea) is generally regarded as a synonym of R. verecundus but differs from that geographically-disjunct species in its relatively shorter tail (subequal to head-body length), smaller auditory bullae, narrower incisive foramina, and uniformly dark tail (see Taylor et al., 1982). In my opinion it should be regarded as a species distinct from R. verecundus, which is thereby restricted to PNG and not reviewed here. Nevertheless, R. verecundus probably extends into Papua in the western Star Mountains region (Table 4.10.5), though there are no records to date. 59, 60, 61, Uromys scaphax and U. nero are species distinct from U. caudimacula63. tus, a point I will review in a forthcoming paper (Helgen, unpublished). The insular species U. boeadii (Biak-Supiori) is most closely allied to the mainland U. nero. Fruit-bats (numbering follows Table 4.10.3) 4. Though often regarded as distinct species (e.g., Flannery 1995a), Dobsonia magna (of New Guinea and Australia) and D. moluccensis (of the Moluccas) are extremely closely allied and better recognized as conspecific (Helgen, unpublished; D. Byrnes, in litt.). 7. Nyctimene albiventer as recognized here (i.e., incorporating papuanus Andersen, 1910) is likely to represent multiple distinct species (N. Irwin, pers. comm.). 8. Though Nyctimene cyclotis and Nyctimene certans are commonly recognized as distinct, sympatric montane species following Peterson (1991) and Flannery (1995a), more recent morphological (Helgen, unpublished) and electrophoretic (Donnellan et al., 1996) analyses offer no support for their separation. They should be regarded as a single species, N. cyclotis, widespread in montane contexts. 10. An unidentified species of Nyctimene occurs on the islands of BiakSupiori and Numfoor in Cenderawasih Bay. This is not N. cyclotis (contra Flannery 1995b) and may represent a new species.

................. 16157$

CH43

02-08-07 10:48:42

PS

PAGE 748

Taxonomic and Geographic Overview of the Mammals / 749

11, 12.

16.

19, 20.

Geographic distributions for Paranyctimene raptor and P. tenax were not delineated by Bergmans (2001) in his description of P. tenax and thus remain confused. Paranyctimene tenax may be widely distributed in the northern lowlands of New Guinea and P. raptor in the south (N. Irwin, pers. comm.). Pteropus melanopogon is newly recorded here from Papua based on a specimen at the Naturalis Museum (Leiden, Netherlands) from the Raja Ampat island of Misool (number 37766). An undescribed species of Pteropus (allied to P. pohlei) occurs on the oceanic islands of Biak-Supiori and (probably) Waigeo in Papua.

Insectivorous bats (see Table 4.10.2) Hipposideros wollastoni: taxonomy of this species is not well understood, and I provisionally separate the subspecies proposed by Flannery and Colgan (1993) as separate species, restricting H. wollastoni (s.s.) to the southern lowlands of New Guinea. Emballonura cf. alecto: recorded from Gag Island by Maryanto and Kitchener (1999). Emballonura serii: newly recorded from Papua based on a specimen from Yapen at the American Museum of Natural History, New York (AMNH 221958; D. Lunde, in litt.); this specimen was misidentified as Emballonura furax by Bonaccorso (1998). Myotis moluccarum: Recognized as distinct from Myotis adversus following Kitchener, Cooper, and Maryanto (1995). Myotis cf. stalkeri: M. stalkeri or a related large-bodied Myotis has been collected from Waigeo by Meinig (2002) and from Batanta by M. Farid (in litt., 2005).

................. 16157$

CH43

02-08-07 10:48:42

PS

PAGE 749

................. 16157$

CH43

02-08-07 10:48:42

PS

PAGE 750

Marshall, Andrew J., and Bruce M. Beehler. Ecology of Indonesian