The Ecology of Papua: Part Two [6, 1 ed.] 0794604838, 9780794604837

Table of contents :
Intro
Contents
Abbreviation
Section V: Natural Ecosystems
5.1. The Diversity and Conservation of Papua’s
Ecosystems
5.2. The Ecology of Papuan Coral Reef
5.3. Seagrass Ecosystems of Papua
5.4. Mangrove Forests of Papua
5.5. Inland Water Ecosystems in Papua:Classification, Biota, and Threats
5.6. Coastal Vegetation of Papua
5.7. Lowland Swamp and Peat Vegetation of Papua
5.8. Lowland Vegetation of Papua
5.9. Heath Vegetation of Papua
5.10. Montane Vegetation of Papua
Gallery
5.11. Subalpine and Alpine Vegetation of Papua
5.12. Grassland and Savanna Ecosystems of the Trans-Fly, Southern Papua
5.13. Caves of Papua
Section VI: Human-Ecosystem Interactions
6.1. The History of Human Impact on New Guinea
6.2. A Brief Social and Political History of Papua, 1962–2005
6.3. The Agricultural Systems of Papua
6.4. Patterns of Commercial and Industrial Resource Use in Papua
6.5. Natural Resource Economics of Papua
Section VII: Conservation of Papuan Natural Resources
7.1. Threats to Biodiversity
7.2. Setting Priorities and Planning Conservation in Papua
7.3. The Protected Area System in Papua
7.4. Conservation Laws, Regulations, and Legislation in Indonesia, with Special Reference to Papua
Gallery
7.5. Opportunities and Challenges for Doing
Conservation in Papua
7.6. Community-Based Conservation in the Trans-Fly Region
7.7. A Non-native Primate (Macaca fascicularis) in Papua: Implications for Biodiversity
7.8. Exotic Herpetofauna: A New Threat to New Guinea’s Biodiversity?
Section VIII: Appendices
8.1. Glossary of botanical terms
8.2. Provisional species list of birds occurring in Papua
8.3. The vertebrate fauna of the Fly and Purari Deltas and Bintuni Bay mangroves
8.4. Preliminary checklist of amphibians and reptiles reported from Papua and the Aru Islands
8.5. Fish records in the Trans-Fly
8.6. Mammal records in the Trans-Fly
8.7. IUCN Protected Area Management categories
8.8. Protected areas in Papua
8.9. Habitat coverage within protected areas in Papua
8.10. Nongovernmental institutions in Papua that are working on environmental and conservation issues
8.11. Government institutions that are working on environmental and conservation issues in Papua
Index

Citation preview

EcologyOf Papua Vol.2

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THE ECOLOGY OF INDONESIA SERIES Volume VI

The Ecology of Papua Part Two

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 Toni Whitten

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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

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EcologyOf Papua Vol.2

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THE ECOLOGY OF INDONESIA SERIES Volume VI

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

EcologyOf Papua Vol.2

5/30/07

10:06 AM

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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: 978-1-4629-0680-2 (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 This Volume ix SECTION FIVE. NATURAL ECOSYSTEMS 5.1. The Diversity and Conservation of Papua’s Ecosystems 753 andrew j. marshall 5.2. The Ecology of Papuan Coral Reefs 771 douglas fenner 5.3. Seagrass Ecosystems of Papua 800 len mckenzie, rob coles, and paul erftemeijer 5.4. Mangrove Forests of Papua 824 daniel m. alongi 5.5. Inland Water Ecosystems in Papua: Classification, Biota, and Threats 858 dan a. polhemus and gerald r. allen 5.6. Coastal Vegetation of Papua 901 robert j. johns, garry a. shea, and pratito puradyatmika 5.7. Lowland Swamp and Peat Vegetation of Papua 910 robert j. johns, garry a. shea, and pratito puradyatmika 5.8. Lowland Vegetation of Papua 945 robert j. johns, garry a. shea, and pratito puradyatmika 5.9. Heath Vegetation of Papua 962 garry a. shea, robert j. johns, willem vink, and pratito puradyatmika 5.10. Montane Vegetation of Papua 977 robert j. johns, garry a. shea, willem vink, and pratito puradyatmika 5.11. Subalpine and Alpine Vegetation of Papua 1025 robert j. johns, garry a. shea, and pratito puradyatmika 5.12. Grassland and Savanna Ecosystems of the Trans-Fly, Southern Papua 1054 michele bowe, neil stronach, and renee bartolo 5.13. Caves of Papua 1064 louis deharveng, tony whitten, and philippe leclerc SECTION SIX. HUMAN-ECOSYSTEM INTERACTIONS 6.1. The History of Human Impact on New Guinea 1087 geoffrey s. hope 6.2. A Brief Social and Political History of Papua, 1962–2005 1098 jaap timmer 6.3. The Agricultural Systems of Papua 1125 manuel boissie` re and yohanes purwanto vii

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viii / contents

6.4. Patterns of Commercial and Industrial Resource Use in Papua 1149 dessy anggraeni 6.5. Natural Resource Economics of Papua 1167 james b. cannon SECTION SEVEN. CONSERVATION OF PAPUAN NATURAL RESOURCES 7.1. Threats to Biodiversity 1199 scott frazier 7.2. Setting Priorities and Planning Conservation in Papua 1230 john burke burnett 7.3. The Protected Area System in Papua 1251 yance de fretes 7.4. Conservation Laws, Regulations, and Legislation in Indonesia, with Special Reference to Papua 1276 suer suryadi, agustinus wijayanto, and james b. cannon 7.5. Opportunities and Challenges for Doing Conservation in Papua 1311 yance de fretes 7.6. Community-Based Conservation in the Trans-Fly Region 1327 michele bowe 7.7. A Non-native Primate (Macaca fascicularis) in Papua: Implications for Biodiversity 1348 neville j. kemp and john burke burnett 7.8. Exotic Herpetofauna: A New Threat to New Guinea’s Biodiversity? 1365 burhan tjaturadi, stephen richards, and keliopas krey SECTION EIGHT. APPENDICES 8.1. Glossary of Botanical Terms 1371 8.2 Provisional Species List of Birds Occurring in Papua 1386 8.3 The Vertebrate Fauna of the Fly and Purari Deltas and Bintuni Bay Mangroves 1404 8.4 Preliminary Checklist of Amphibians and Reptiles Reported from Papua and the Aru Islands 1410 8.5 Fish Records in the Trans-Fly 1416 8.6 Mammal Records in the Trans-Fly 1422 8.7 IUCN Protected Area Management Categories 1425 8.8 Protected Areas in Papua 1426 8.9 Habitat Coverage within Protected Areas in Papua 1431 8.10 Nongovernmental Institutions in Papua That Are Working on Environmental and Conservation Issues 1434 8.11 Government Institutions That Are Working on Environmental and Conservation Issues in Papua 1437 Index

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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 ix

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x / ab b r ev ia ti ons used in th is v ol ume

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)

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ab b r evi a ti ons used in thi s v ol ume / xi 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)

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xii / abbreviations u sed i n this v olume

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)

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a b b rev i a tio ns used i n thi s v olu me / xiii 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)

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xiv / 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

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a b br ev ia ti ons used in this v ol um e / xv 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)

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section five 

Natural Ecosystems

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5.1. The Diversity and Conservation of Papua’s Ecosystems andrew j. marshall h e ter m ‘ ‘e c o s ys t e m’’ refers to a biological community and its physical environment. Sir Arthur Tansley, an English botanist who was a pioneer in the study of plant ecology, coined the term in 1935 in recognition of the fact that a true understanding of ecological processes requires consideration of organisms and their habitats as a single, integrated system (Tansley 1935). Some ecologists extended this view and argued that the ecosystem should be considered the basic unit of ecological investigation (e.g., Evans 1956; Rowe 1961). Although modern ecology incorporates research on a variety of scales, from populations of single species, through landscapes and ecoregions, to the entire biosphere, the consideration of ecosystems as functional units has produced important insights into a range of important ecological processes, such as primary production, energy flow, and nutrient cycling. In this section we take a broad, ecosystem-level view of the Papuan environment. This level of analysis allows us to consider issues of biodiversity, conservation, and human well-being from a broader perspective than is possible when these issues are examined at smaller spatial scales. In this introductory chapter I comment briefly on some general concepts related to ecosystem classification, diversity, services, and conservation, and consider how these concepts can be applied to the management and preservation of Papua’s ecosystems. In the following twelve chapters, experts provide overviews of the ecology, organization, and conservation of Papua’s most important ecosystem types. First, a comment on terminology. In ecology, as in many other scientific disciplines, terminology is both a blessing and a burden. When clearly defined and applied, specific terms unambiguously convey meaning and permit relevant debate. Unfortunately, ecological terms are frequently used in contexts other than those in which they were originally applied, without appropriate definition or clarification. Such misuses of terminology obscure meaning and can result in vigorous debates that create much heat while shedding little light on the issues under discussion. The term ‘‘ecosystem’’ is used frequently and in a wide variety of contexts without formal definition. In this volume we use the term to classify specifically delineated parts of the environment and all biological organisms that inhabit them. For example, lower montane forest is a particular ecosystem type that encompasses the physical structure of a mountain (e.g., bedrock, soil) found between roughly 650 and 1,500 meters elevation and all of the flora and fauna living within this structure (Chapter 5.10). It is distinct from the alpine ecosystem type typically found at higher elevations and the lowland forest ecosystem type found below. We do not use the term ‘‘ecosystem’’ to refer to the habitat occupied by a particular

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species, as this term is highly-species specific: the habitat of one species may include many ecosystem types (e.g., many bird species), while another species might only be found in one particular subset of an ecosystem type (e.g., a tree species limited to a particular soil type).

Ecosystem Classification One of the complications that has long plagued ecosystem ecologists is the difficulty in identifying the boundaries of an ecosystem. Sharp lines can rarely be drawn that delineate the extent of a particular ecosystem type or contain all relevant ecological processes and interactions (Whittaker 1970). Even boundaries that initially appear to be clearly delineated are revealed upon careful examination to be porous and dynamic. For example, at first glance few ecosystems would seem to be more clearly distinct than the abutting marine and terrestrial ecosystems found along coastlines around the world. However, closer investigation reveals that energy and nutrients flow between these ecosystems, organisms move back and forth between them, and that the health and stability of one can profoundly effect the other. These interactions make categorization of ecosystems as discrete entities somewhat artificial. For the purposes of description, we have classified Papua’s ecosystems into twelve broad categories. Nevertheless, it is important to remember that these classifications are simplifications made to facilitate discussion, and that in reality ecosystems are highly interconnected and interdependent. The principle division of aquatic ecosystem types is based on water salinity, and two major categories are typically considered: saltwater (or marine) and freshwater ecosystems. Various ecosystem types are defined in each of these broad categories based on physical features such as substrate, temperature, water depth, and dominant vegetation type (Smith and Smith 2003). In this chapter we consider four major categories of aquatic ecosystems in Papua: coral reefs, seagrass ecosystems, mangroves, and inland water ecosystems. The world’s major terrestrial ecosystem types (often referred to as biomes) are classified by vegetation type, which is largely dependent on rainfall and temperature (Whittaker 1970). Within these biomes, separate ecosystems can also be defined according to the composition and structure of the plant community. We follow this convention by considering six distinct vegetative formations (i.e., ecosystems) within the tropical forest biome, following a roughly altitudinal gradient from coastal ecosystems to alpine vegetation. We also consider the extensive monsoon grassland and savanna ecosystems found in plains and deltas of the great rivers in southern New Guinea. Finally, we discuss the unique and little-known cave ecosystems of Papua.

Ecosystem Diversity in Papua Many of the terrestrial ecosystem types discussed in this section are further subdivided based on dominant vegetation, altitude, soil type, and degree of human

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disturbance. The wide array of ecosystem types found in Papua helps to explain why this is an area of such high biodiversity and a major center of endemism in many distinct taxonomic groups. From the reefs that contain the most coral species in the world to the cryovegetation communities growing in the ice and snow atop its highest mountains, Papua’s ecosystem diversity creates a wide range of ecological conditions, each of which supports a highly specialized community of flora and fauna. Some ecosystems are fairly well characterized and understood (e.g., seagrass ecosystems, coastal vegetation), while others are scarcely known and the diversity and ecological interactions contained therein have yet to be discovered (e.g., cave ecosystems). Yet the uniqueness, complexity, and diversity of each of these ecosystem types is abundantly clear, and helps to make Papua one of the most biologically important regions on earth (Supriatna 1999). Papua’s high diversity of terrestrial ecosystems is largely due to its wide altitudinal range (Figure 5.1.1). Accurate measures of the extent of different ecosystem types in Papua are difficult to calculate, both because of difficulties in classifying ecosystems and complications in recognizing these ecosystem types on images obtained through remote sensing. However, based on general land cover classifications (Hansen et al. 1998) and recent Landsat 7 ETM imagery of Papua (1999–2000), the extent of broad land classes can be mapped (Figure 5.1.2). Anal-

Figure 5.1.1. Surface elevation and ocean depth in Papua. The wide range of altitudes leads to a diversity of ecosystem types.

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Figure 5.1.2. Forest cover in Papua. The picture is an interpretation of Landsat 7 ETM imagery of Papua, using a combination of images acquired in 1999 and 2000. Source: Forest Watch Indonesia–Conservation International–Ministry of Forestry.

ysis of the resulting map provides estimates of the extent of each major land class in Papua. When forests are broadly defined to include all land cover classes with greater than 10% tree or shrub canopy cover, roughly 85% of Papua was forested in 2000 (Table 5.1.1). Over 60% of these forests were lowland evergreen forests (51% of Papua’s total area), making Papua home to the largest remaining tracts of lowland tropical evergreen forest in Indonesia. Large areas of mangrove forest (15,124 km2, 4.3% of forested land), swamp ecosystems (68,312 km2, 19.5% of forested land), and montane forest (36,032 km2, 10.3% of forested land) are also found, in addition to several other ecosystem types, each of which comprise more than 1% of forested area in Papua (Table 5.1.1). The distribution and diversity of ecosystem types across the island of New Guinea are similar to those found in Papua (Figure 5.1.3). Due to differences in data quality and forest classification, figures for New Guinea are not directly comparable to those from Papua. However, analyses show that in 2000 the island of New Guinea was overwhelmingly forested, containing almost 657,000 km2 (82% of the land area) of broadleaf forest and woodland (Table 5.1.2). For this reason New Guinea is considered one of the world’s three great lowland tropical rainforest Wilderness Areas (Mittermeier et al. 2003).

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Table 5.1.1. Major land classes in Papua Land classes

Area (km2)

Mangrove forest Swamp

% of forested land

% of total land

15,124

4.3

3.6

7,465

2.1

1.8

Swamp brush

10,559

3.0

2.5

Swamp forest

50,288

14.3

12.1

Lowland evergreen rainforest

213,627

60.8

51.3

Lower montane rainforest

8,658

2.5

2.1

Upper montane rainforest

27,373

7.8

6.6

Subalpine forest

4,266

1.2

1.0

Brush

4,490

1.3

1.1

Savanna

9,298

2.6

2.2

Total forest cover Bare ground, rice paddies, transmigration settlements Total land area

351,147

84.4

64,982

15.6

416,129

100.0

Numbers may not accord perfectly with data listed in other chapters in this section due to differences in habitat classification and methods used to estimate the extent of each habitat type. Listed are estimates from Interpretation of Landsat 7 ETM imagery of Papua, using a combination of images acquired in 1999 and 2000. Source: Forest Watch Indonesia–Conservation International–Ministry of Forestry.

the importance of ecosystem diversity for papuan fauna Landscapes containing several ecosystem types have higher species richness than equivalent areas containing only a single ecosystem type (Figure 5.1.4). Thus Papua’s high ecosystem diversity helps to explain the high diversity found in a number of taxonomic groups of flora (Section 3) and fauna (Section 4). In addition, many vertebrate species rely on more than one ecosystem type, often utilizing different ecosystem types for breeding, nesting, and foraging. For example, several species of sea turtles feed in open oceans and seagrass ecosystems, but rely on coastal beaches to lay their eggs (Chapter 4.6). Similarly, many species of mammals, insects, and birds breed in mangrove forests but live and forage mainly in adjacent terrestrial or marine habitats (Chapter 5.4). Greater Melampittas nest in cave ecosystems but forage daily in nearby forest ecosystems (Chapter 5.13), and many other bird species utilize several forest types at a variety of altitudes during their normal life cycles (Chapter 4.9). Therefore, preservation of the full complement of ecosystem types in Papua is necessary both to preserve its high biodiversity and to provide the habitat requirements for a number of threatened vertebrate species.

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Figure 5.1.3. Forest cover in New Guinea. The picture is an interpretation of Landsat 7 ETM imagery of Papua, using a combination of images acquired in 1999 and 2000. Source: Forest Watch Indonesia–Conservation International–Ministry of Forestry.

Table 5.1.2. Major land classes in New Guinea Land classes

Area (km2)

Evergreen broadleaf forest

540,418

Deciduous broadleaf forest

% of land area 67.3

12,130

1.5

104,369

13.0

Wooded grassland

82,196

10.2

Closed shrubland

1,745

0.2

Open shrubland

16,254

2.0

Grassland

35,668

4.4

Cropland

6,359

0.8

Bare ground

4,358

Woodland

Urban and built-up area

0.5

75

Total land area

0.01

803,572

100.0

Land classes in this table are different from those presented in Table 5.1.1. Source: Hansen et al. (1998).

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Figure 5.1.4. Schematic diagram of species richness and ecosystem diversity. The graph shows species-area curves for two landscapes. Landscape 1 (solid line) is comprised of a single ecosystem type (A). As more area is sampled the total number of species recorded increases, but slope decreases as an increasing proportion of the total species richness of ecosystem A is recorded. Landscape 2 (dashed line) is comprised of three ecosystem types (A, B, C). The species-area curves for the two landscapes are equivalent as long as sampling is confined to ecosystem A. However, as sampling begins in ecosystem B the species-area curve in Landscape 2 increases sharply as many new species are recorded in this new ecosystem type. Sampling of ecosystem C results in another rapid increase in total species richness in Landscape 2. This schematic shows that for any sampling area (e.g., a⬘) species richness is higher in landscapes containing multiple ecosystem types than in landscapes comprised of a single ecosystem type (e.g., s2 ⬎ s1).

interactions among ecosystem types As is the case in many subjects within ecology and conservation biology, the more we learn about ecosystems the more we realize how connected and interdependent they are. As noted above, classification of ecosystems into discrete ‘‘types’’ masks the fact that there are many important interactions among them. For example, seagrass ecosystems provide an important functional link and buffer between reefs and mangrove ecosystems (Chapters 5.3 and 5.4) and forest ecosystems provide key nutrient inputs into aquatic and cave ecosystems (Chapters 5.5 and 5.13). This interdependence means that when one ecosystem is damaged it can have strong and often unforeseen effects on adjacent ecosystems. For example, uncontrolled clear-cutting of forest not only negatively effects forest ecosystems; the resultant erosion can also lead to detrimental siltation of downstream aquatic ecosystems (Chapter 5.5) and sediment deposition that can cause major harm to coral reefs (Chapter 5.2). Similarly, the smoke resulting from large-scale burning of lowland forests and peat swamps can have effects on other ecosystem types. For instance,

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the fires that occurred in Sumatra and Kalimantan in 1997 led to red tide phytoplankton blooms that caused large-scale death of coral reefs in the Mentawai Islands (Abram et al. 2003, 2004) and Bali (van Woesik 2004). We are only beginning to understand the complexity of interactions among ecosystem types, but these examples warn us that degradation of one ecosystem can have broad cascading effects on other ecosystems.

Papuan Conservation: An Ecosystem Perspective Conservationists address questions across a broad range of spatial scales. Each of these approaches can yield valuable insights and have important implications for the preservation of biodiversity. Here I briefly consider some of the issues relevant to conservation of entire ecosystems. I consider the services provided by Papua’s ecosystems, discuss research into the relationship between biodiversity and ecosystem function, assess the representation of different ecosystems in Papua’s protected areas network, and consider the implications of an ecosystem perspective on Papuan conservation issues.

ecosystem services The earth’s ecosystems provide a wealth of services necessary for human health and well-being, many of which are taken for granted or severely undervalued. Such services include purification of air and drinking water, reduction in the severity of droughts and floods, generation and preservation of soils and soil fertility, pollination of crops, nutrient cycling, climate stabilization, carbon sequestration, control of infectious disease, and erosion protection (Daily 1997; Krebs 2001). The relatively new field of natural resource economics has helped to raise awareness of the immense financial value of ecosystem services (Balmford et al. 2002, 2003; Balmford and Whitten 2003; Costanza 1991; Costanza et al. 1997; James, Gaston, and Balmford 1999; Peet 1992; Chapter 6.5), but the true benefits and costs of ecosystem services and their loss are rarely incorporated into decisions about natural resource management, particularly in developing countries. The financial costs associated with loss of ecosystem services resulting from degradation are rarely (or never) fully offset by those perpetrating the degradation, and the social and health costs are frequently disproportionately paid by people in lower economic groups. For example, the health costs alone associated with the Indonesian forest fires in 1997 have been estimated at 145 million U.S. dollars, with the majority of morbidity and mortality falling upon the poorest people in the region (Barber and Schweithelm 2000). Similar fires burn almost yearly and those who profit financially from these ecological disasters are not held accountable. Papua’s ecosystems provide environmental services of immense local, regional, and global importance. For example, Papua’s forests maintain water quality and prevent soil erosion for numerous local communities. Regionally, Papua’s mangroves serve as important breeding grounds for endangered vertebrates and commercially important marine invertebrates, sequester pollutants and environmental

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contaminants, protect against coastal erosion, and can even serve as physical barriers protecting humans from tsunami (Alongi 2002; Danielsen et al. 2005). More broadly, Papua’s extensive forests and seagrass ecosystems serve as globally important sites of carbon sequestration that help to ameliorate global climate change. Therefore sound management and conservation of Papua’s ecosystems will ensure that the valuable environmental services they provide will enhance human health and well-being for future generations.

diversity and ecosystem function Because ecosystems provide such a wide range of services crucial to human health, substantial theoretical, empirical, and experimental work has addressed the relationship between diversity (or, more specifically, species richness) and ecosystem function. Although the theoretical roots of this discussion go back decades (MacArthur 1955; May 1972; Statzner and Moss 2004), the unprecedented extinction rates resulting from human degradation of natural ecosystems have made this issue one of considerable practical relevance in recent years (Cameron 2002; Kinzig 2001; Loreau et al. 2001, 2002; Naeem et al. 1994; Schwartz et al. 2000). Examination of this topic is complicated by several issues. First, until recently, unusually contentious academic debate over the role of biodiversity in ecosystem functioning has polarized discussion, hampered important syntheses, and created skepticism towards this important work among the general public (Mooney 2002; Naeem et al. 2002). Happily, recent collaborative syntheses have reduced these tensions and identified important new directions of investigation (e.g., Loreau et al. 2001; Hooper et al. 2005). Second, there are different measures of ecosystem function relevant to human well-being, including primary and secondary productivity, stability, resistance to invasion, and resilience, and there is little reason to expect that these different characteristics will be affected by biodiversity losses in similar ways (Hooper et al. 2005; Loreau et al. 2001; Schwartz et al. 2000). Third, multiple mechanisms may be responsible for observed relationships between diversity and ecosystem function (Loreau et al. 2002), highlighting one of the frequent difficulties ecologists face in attempting to infer processes from patterns. Finally, much of the recent experimental work has focused on studying the effects of manipulation of small-scale systems with relatively low species richness (e.g., McGradySteed et al. 1997; Petchey et al. 1999; The´bault and Loreau 2003; Tilman 1999). As most applied conservationists are primarily concerned with complex, large-scale systems, the practical relevance of insights gained from the study of much simpler systems is debatable on several grounds (e.g., Aarson 1997; Carpenter 1996; Hooper and Vitousek 1997; Huston 1999; Huston and McBride 2002; Rosenfeld 2002; Strivastava and Vellend 2005). From a conservation standpoint, the key question is related to ecological redundancy (Lawton and Brown 1993; Rosenfeld 2002): are all species in an ecosystem necessary to sustain normal function, or can most ecosystem services be provided by a small subset of species (i.e., are many species functionally redundant)? It is unlikely that there is a universal relationship between diversity and ecosystem

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function across all ecosystem types and functions (Hooper et al. 2005; Naeem et al. 1994). Some studies indicate that there are relatively high degrees of ecological redundancy and that substantial losses in biodiversity may have limited effects on the provision of certain ecosystem services, especially at small temporal and spatial scales or when environmental variability is relatively low (Hooper et al. 2005; Loreau et al. 2001; Schwartz et al. 2000, but see Rosenfeld 2002). However it should be noted that these studies typically use a limited definition of ecosystem function (often restricted to the effects of biodiversity loss on plant biomass), and many restrict their analyses to examining the effects of biodiversity loss within one trophic level. At larger temporal and spatial scales and in changing environments the number of species required to maintain ecosystem services increases (Hooper et al. 2005; Loreau et al. 2001, 2002). Research has now largely shifted away from focusing on simple indices of species richness to attempting to identify key functional species or groups that have disproportionate effects on ecosystem services (Loreau et al. 2001; Naeem and Wright 2003; Rosenfeld 2002). There is debate over the extent to which biodiversity-ecosystem function studies have direct relevance for conservation biology (Hector et al. 2001; Lawler et al. 2001; Schwartz et al. 2000; Srivastava and Vellend 2005). The lack of universal support for a direct link between biodiversity and ecosystem function has led some to suggest that widespread use of this linkage as a justification for conservation goals is unwise (Krebs 2001; Lawler et al. 2001; Schwartz et al. 2000). Others acknowledge this point but argue that interactions between biodiversity and ecosystem services can provide useful additional arguments in support of conservation (Hector et al. 2001). It has also been suggested that although research on the relationship between biodiversity and ecosystem function has had limited conservation applications to date, this area promises to provide important insights into conservation policy in the foreseeable future (Lawler et al. 2001; Srivastava and Vellend 2005). While there is much debate in academic circles on how reduction in species richness or loss of key functional groups will effect the function and stability of ecosystems (and the pertinence of these debates to more applied conservation issues), the vast majority of ecologists agree that these losses will increase susceptibility to invasion by exotic species (and presumably also pathogens), reduce environmental services, and negatively impact the biosphere (Hooper et al. 2005; Loreau et al. 2001; Schla¨pfer et al. 1999). Therefore, as ecologists work to identify which species and functional groups are irreplaceable, a precautionary approach to biodiversity preservation should serve as a broad governing theme in conservation management in Papua.

the papuan protected areas network The Papuan protected areas network encompasses approximately 66,500 km2 of terrestrial habitats. The major ecosystems are not equally or proportionately represented within Papua’s protected area system (Table 5.1.3). The most well protected land cover classes are lower montane forests and subalpine forests, with over 45% of each ecosystem type found within formally protected areas. However, lowland

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9,298

Savanna 18,242

0

211

47

2,317

2,216

12,539

106

38

51

718

5.2

0.0

4.7

1.1

8.5

25.6

5.9

0.2

0.4

0.7

4.7

Nature Reserve % land km2 cover

27,581

1,237

276

355

2,171

974

11,309

7,171

1,963

617

1,508

7.9

13.3

6.2

8.3

7.9

11.2

5.3

14.3

18.6

8.3

10.0

Wildlife Reserve % land km2 cover

20,445

1,346

388

1,523

4,042

777

6,785

2,397

591

474

2,122

5.8

14.5

8.6

35.7

14.8

9.0

3.2

4.8

5.6

6.4

14.0

National Park % land km2 cover

298

0

0

0

0

0

296

1

0

0

1

0.1

0.0

0.0

0.0

0.0

0.0

0.1

0.0

0.0

0.0

0.0

Nature Recreation Reserve % land km2 cover

66,567

2,583

875

1,925

8,531

3,966

30,928

9,675

2,592

1,142

4,349

19.0

27.8

19.5

45.1

31.2

45.8

14.5

19.2

24.6

15.3

28.8

Total % land km2 cover

Unforested lands are excluded. GIS analysis of protected areas map overlaying Landsat 7 ETM imagery of Papua, using a combination of images acquired in 1999 and 2000. Source: Forest Watch Indonesia–Conservation International–Ministry of Forestry.

351,147

4,490

Total

4,266

Brush

27,373

Upper montane rainforest

Subalpine forest

8,658

Lower montane rainforest

213,627

50,288

Swamp forest

Lowland evergreen rainforest

10,559

7,465

15,124

Swamp brush

Swamp

Mangrove forest

Forest cover classes

Area (km2)

Table 5.1.3. Representation of Papuan land classes in protected areas The Diversity and Conservation of Papua’s Ecosystems / 763

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evergreen forest, by far the most dominant ecosystem type in Papua (61% of land area), is proportionately the least well represented in protected areas, with only 14.5% of this ecosystem type occurring within currently designated parks and reserves. This is a source of major concern as lowland forest is the ecosystem type most likely to suffer heavy degradation from uncontrolled human development, logging, and mining. In other parts of Indonesia almost all lowland forest found outside protected areas has disappeared or been severely degraded (Fuller et al. 2004; Holmes 2002; Jepson et al. 2001; van Schaik et al. 2001; World Bank 2001), with substantial losses even occurring within protected areas (Curran et al. 2004). Although this trajectory of habitat loss is far from inevitable in Papua, we would be well advised to consider the worst-case scenario that few forests in Papua will exist in their present state outside protected areas at the end of the 21st century. Under this scenario the current protected areas network in Papua is unlikely to be sufficient to protect the full complement of species, ecological processes, and ecosystem functions that is found there today. Assessment of this possibility will require careful consideration of the representation of ecosystem types within the current Papuan protected areas network, the prospects for maintaining connectivity between ecosystems, and the potential effects of global and regional climate change on the spatial distribution of ecosystems.

ecosystem-based conservation approaches Numerous strategies are currently employed and championed by scientists, conservation organizations, and government agencies involved in natural resource management. The widely-publicized ‘‘Biodiversity Hotspots’’ approach advocates prioritizing severely threatened areas of high species richness and endemism (Myers et al. 2000). Other strategies suggest that preservation of large wilderness areas that are ecologically intact and sparsely populated represent important opportunities for biodiversity conservation (Mittermeier et al. 2003). Some have argued that conservation efforts should focus almost exclusively on landscapes that are largely unaltered by humans (e.g., Myers 1980; Noss 1991), while others embrace the conservation potential of the careful management of lands that have already been substantially impacted (e.g., by development or logging, Fimbel 1994; Fimbel et al. 2001; Frumhoff 1995; Johns 1983; Marshall et al. 2006; Meijaard et al. 2005). Integrated conservation and development projects promise simultaneously to protect biodiversity and promote human well-being, health, and poverty alleviation (Goodwin and Swingland 1996; McShane and Wells 2004; Salafsky et al. 2001), while others suggest that the most effective way to conserve wildlife and habitats is strict protection and exclusion of most local people from protected areas (e.g., Terborgh 1999). Numerous campaigns have focused on the conservation of single species or specific taxonomic groups (e.g., Mittermeier et al. 2005; Stattersfield et al. 1998) and others work to preserve ecosystems, ecoregions, or functional landscapes (Hudson 1991; Noss 1996; Pressey et al. 1993; Woinarski et al. 1996). Each approach has strengths and limitations (Bonn and Gaston 2005; Kareiva and Marvier 2003; Kiss 2004; Young 1999; Orme et al. 2005; Possingham

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and Wilson 2005), and it is likely that the most favorable conservation outcomes will result from careful application of a broad portfolio of conservation tactics and strategies. Although a range of conservation strategies have applicability to Papua, the fact that Papua’s ecosystems provide services of local, regional, and global importance strongly suggests that ecosystem-level conservation approaches are particularly warranted. The specific goals of ecosystem-based conservation plans will need to be carefully considered within the Papuan context, but the five basic goals of ecosystem management proposed by Grumbine (1992, 1994) provide a useful point of departure. Ecosystem management should strive first to protect sufficient habitat to ensure the long-term viability of populations of all native species; second, to represent all native ecosystem types across their range of natural variation within protected areas; third, to manage ecosystems on spatial scales that are sufficiently large to maintain important ecological processes (e.g., disturbance regimes, hydrological processes, nutrient cycles); fourth, to create ecosystem management plans for sufficiently long time scales (e.g., centuries) to permit evolutionary change; and fifth, to allow for human use and occupancy at levels that do not result in ecological degradation (Grumbine 1992, 1994). Ecosystem-based conservation plans in Papua are likely to be complicated to devise and even more challenging to implement effectively. Political support will need to be generated at all levels of government, ecosystems will need to be legally defined and delineated, consensus among diverse ethnic groups will need to be reached, and effective mechanisms to monitor the success of conservation interventions will need to be implemented. Ultimately, conservation efforts in Papua will not be successful unless such large-scale conservation issues are tackled. Papua, and New Guinea more broadly, is a region of global biological significance. It includes the highest summit in Oceania, the only equatorial glaciers in the Pacific, the most extensive and diverse mangrove forests in Indonesia, and one of the world’s largest remaining tracts of lowland tropical forest. Human population density in Papua is low. Rates of forest loss and remaining forest cover in Papua are encouraging when compared with many other areas in the tropics. Papua also is home to extensive and highly-diverse reefs that remain largely undamaged, at least in comparison to those in western Indonesia and many other parts of the world. However, threats to these ecosystems exist and will likely increase over time. We should have no illusions that protection of Papua’s ecosystems will be easy or simple. Despite unprecedented investment in conservation, efforts to protect Indonesia’s other lowland forests have largely failed (Curran et al. 2004; Fuller et al. 2004; van Schaik et al. 2001; Whitten et al. 2001). Our current conservation strategies have proved inadequate in the face of the legitimate and pressing demands of Indonesia’s poorest citizens and the greed of illegal logging bosses. Papua presents one of the few remaining opportunities for proactive conservation action in Indonesia. Avoiding the fate of the rest of Indonesia’s oncevast tracts of lowland forest will require a level of political will that has thus far

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proved difficult to generate in other parts of the country. But the stakes are too high for us to let Papua quietly go the way of Sumatra and Borneo. The fate of Indonesia’s last great wilderness area, and the people who rely on it, hangs in the balance.

Acknowledgments I thank Hendi Sumantri for assistance with maps and spatial analysis, and Conservation International and the Arnold Arboretum of Harvard University for postdoctoral support. I also thank Peter Ashton, Bruce Beehler, Amy Dunham, Mark Leighton, Cam Webb, and Tony Whitten for useful comments on this chapter.

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5.2. The Ecology of Papuan Coral Reefs douglas fenner o r al r e e fs are structures in shallow ocean water built at least in part by corals, but often with major contributions from coralline algae and green calcareous algae. Coral reefs are composed of calcium carbonate (CaCO3) produced by living organisms. Reefs range in size from small patches a few meters in diameter to the largest structure on earth built by living organisms, the Great Barrier Reef of Australia, which is about 2,000 km long, covers an area of about 128,000 km2, and contains about 2,500 individual reefs. Coral reefs can also be very thick. At Enewetak Atoll in the Marshall Islands, a hole was drilled through over 1,400 m of coral rock before reaching the underlying volcanic rock. Thus, some reefs are major geological features of planet earth. The coral rock near the bottom of the coral at Enewetak was found to be 65 million years old, indicating that some coral reefs are also very old. Three of the most widely recognized coral reef shapes are the fringing reef, barrier reef, and atoll. A fringing reef grows along the shore of a landmass, much like a fringe along the edge of a coat. A barrier reef parallels a coastline, with a lagoon between the barrier reef and the shore. These lagoons are relatively shallow (about 10–50 m deep) and usually have a sandy bottom. Barrier reefs are named because if they are continuous and reach the surface, they are a barrier to navigation. An atoll is a ring of coral with no land other than some low sand islands within the ring of coral. The center of the atoll is a lagoon. Other reef shapes include bank or platform reefs where an offshore reef does not reach the surface, and patch reefs, which are small patches of coral in lagoons. There are a wide variety of other shapes and intermediates between categories as well (e.g., Andre´foue¨t 2004; Guilcher 1988; Hopley 1982; Tomascik et al. 1997).

C

Evolution of Reefs Charles Darwin proposed a theory of the evolution of coral reefs (Darwin 1842; Tomascik et al. 1997). He proposed that coral reefs begin as fringing reefs, then become barrier reefs, and finally become atolls. Darwin’s theory applied to volcanic islands. He suggested that after an oceanic volcano erupts and builds an island, fringing coral reefs will grow around the shoreline of the volcanic island and that the island will then slowly subside or sink. As the island sinks, the reef will grow upward, and if it can grow upward as fast as the island sinks, eventually the reef will be separated from the island by a lagoon. The reef will then be a barrier reef, and the reef will mark the location of the original shoreline of the island. Eventually the island will sink under water and out of sight under the lagoon sand. The result is an atoll (Figure 5.2.1). Darwin knew that the way to test his theory was to Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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Figure 5.2.1. Schematic diagram of the creation of a coral atoll. Charles Darwin proposed that fringing reefs growing on volcanic islands changed over time to produce barrier reefs and ultimately formed coral atolls. See text for details. drill into an atoll. He predicted that the drill would reach volcanic rock under the coral. The technology to drill such a deep hole was not available in his time, but eventually atolls such as Enewetak were drilled and volcanic rock was found underneath the coral rock, confirming Darwin’s hypothesis. The full sequence of evolution of coral reefs can be seen in Hawai’i, where the Big Island has an erupting volcano and fringing reefs. Small barrier reefs can be found on older islands such as Oahu and Kaui, and the oldest islands (which are found in the northwestern Hawai’ian Islands) are all atolls (Grigg 1982, 1997; Scott and Rontondon 1983)

Papuan Reefs Papua Province has many fringing reefs, some barrier reefs, and very few atolls. So, for instance, in the Raja Ampat Islands, reefs are mostly fringing or platform reefs with 36 fringing reefs and nine platform reefs reported in one study (McKenna, Boli, and Allen 2002). Several maps show coral reefs in the Raja Ampat Islands at the west end of Papua, around the islands in and along the western shore of Cenderawasih Bay, on the south shore roughly across from Cenderawasih Bay, and around the Aru Islands south of the main landmass (UNDP/FAO 1988; Spalding, Ravilious, and Green 2000; Burke, Selig, and Spalding 2002). The south coast of the Vogelkop Peninsula has narrow fringing reefs, with 450 km of coastline suitable for reefs. Tomascik et al. (1997) list nine barrier reefs in Papua, total-

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ing 601 km in length and having an area of 2,366 km2. They also list one atoll in Irian Bay, nine in the Halmahera Sea, and five in the Pacific Ocean. The extent of coral reefs in Papua may be underestimated due to the assumption that coral reefs cannot live along coasts that have mangroves. The muddy shorelines associated with mangroves are often not suitable habitat for coral reefs, yet in New Guinea there are places where coral reefs grow adjacent to mangroves, and mangroves even grow onto the reef platform (P. Dalzell, per. comm.). Some maps of Papua (e.g., Spalding, Ravilious, and Green 2000) show long stretches of coast containing some of the largest mangrove forests in the world. A long stretch of the north coast is normally depicted as being devoid of coral reefs, but fringing reefs are believed to stretch along much of the coast between Sarmi and the border with Papua New Guinea. Fringing reefs are reported on the northern coast of the Vogelkop Peninsula, as well as east from Jayapura to the Papua New Guinea border, and a fringing reef west of Jayapura for 100 km, possibly for an additional 160 km (Tomascik et al. 1977). The coral reefs of Papua New Guinea are much better studied, and yet about half of its coastline has not been explored for coral reefs (Yamuna and McClanahan 2001). Whitehouse (1973) claimed that on the north coast of Papua New Guinea there are no active coral reefs for 1,250 km, but Kojis, Quinn, and Claereboudt (1985) found fringing reefs with a high coral cover and diversity are common along this coast except near the mouths of rivers. The north coast of Papua probably has similar reefs. The total amount of coral reefs in Papua could be several times that presently known. Reefs in Papua are protected from strong wave action. Reefs at 45 sites in the Raja Ampat Islands off the western end of the Vogelkop Peninsula (described in McKenna, Boli, and Allen 2002) vary from those exposed to the open ocean, to those that are in sheltered bays, to one that was so enclosed it was virtually a saltwater lake (Mayalibit Bay within Waigeo Island). Strong currents were not encountered at most sites. The seas are very calm compared to those at oceanic mid-Pacific reefs or the Great Barrier Reef. The reefs do not feature a reef crest with large crashing waves and high cover of coralline algae, nor extensive reef flats and lagoons. Rather, the bottom usually slopes away directly, starting at the shoreline. This is typical of reefs in the region, as the author has seen in northern Sulawesi (Allen and McKenna 2001), eastern Papua New Guinea (Allen et al. 2003), Malaysia (Harborne et al. 2000), and 11 areas in the Philippines (Fenner, under review c; Werner and Allen 2000). Reefs on the southwest side of Cenderawasih Bay include patch reefs with seaward margins that are sheer drop-offs from the crest to a first ledge at 20–40 m depth. Sub-sea level patch reefs have a variable gradient fore reef. Fringing reefs have a variable gradient in bays and sheltered areas but are steeper elsewhere (UNDP/FAO 1982; UNEP/IUCN 1988). Coral cover has been used as a measure of coral health. Because damage to reefs reduces coral cover, reefs with higher coral cover have been presumed to be in better condition. Data on coral cover for 13 sites in the Padaido Islands in Cenderawasih Bay are available (Tomascik et al.1997). Most commonly they had cover of 25–50%. In 44 sites in the Raja Ampats, McKenna, Boli, and Allen (2002) found

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that the average coral cover was 28%. A qualitative scale for coral cover was devised by Gomez et al. (1994), where 75–100% cover was considered excellent, 50–75% good, 25–50% fair, and 0–25% poor. Thus, reefs in these two areas would be considered fair. Indonesia as a whole is reported to have 2.6% of its reefs as excellent, 24.2% good, 31.6% fair, and 41.6% poor condition (Ming et al.1994). Such a scale should be approached with caution, since it implies value judgments that are not based on empirical studies. Coral cover varies substantially depending on habitat and sediment dynamics. No clear generalization can be drawn on the relationship of coral cover to reef health (Maragos 1997). Further, reefs that are among the most pristine known rarely have coral cover in the excellent range, and frequently are in the fair range. Examples include the northwest Hawai’ian Islands (Grigg 1983) and the Great Barrier Reefs (AIMS monitoring, www.aims.gov.au). The scale was originally proposed as a measure of reef health and degradation, yet the natural baseline conditions of reefs are not known. Senior ASEAN scientists assessed eastern Indonesia as having 10% of its reefs degraded 50 years ago and 50% in 1993 (Ming et al. 1994), but figures for Papua were not given. The latest Status of Coral Reefs of the World: 2004 Report (Wilkinson 2004) indicates that the overall reef condition of Indonesia has been improving since 1999, with a shift from reefs with less than 25% cover to reefs with 25–50% cover.

sea level changes The area of eastern Indonesia and New Guinea is a geologically active area, due to the collision of the Indo-Australian, Eurasian, Caroline, Philippine, and Pacific plates. There are many areas of uplift in eastern Indonesia, and many places that have terraces on slopes above the waterline, including along the northwestern coast of Papua (Tomascik et al.1997). On the Huon Peninsula of northeastern Papua New Guinea, continuous uplift of the landmass along with oscillating changes in sea level have produced a stair step series of fossil coral reefs on land. During ice ages, huge amounts of water are withdrawn from the oceans and locked up in giant ice sheets on land in North America, Europe, and Asia, much like in Antartica today. In addition, lower temperatures in the oceans cause the water to contract. These two processes together cause sea levels to drop substantially during ice ages. The last ice age peaked at about 22,500 years ago, causing a drop in sea level of about 120 m. This is well below the lower limits (usually around 30 m) of all presently living coral reefs. Thus all presently living coral reefs were exposed to air at that time and died. The change in sea level happened sufficiently slowly that larvae of sessile organisms such as corals were able to attach farther down and remain alive in the water, and begin building reefs farther down the slopes of islands and continents. In an area like the Huon Peninsula where land is rising steadily, coral reefs build up along the shore when the water levels rise at about the same rate as the land rises. Then when the sea level drops, the reef is left out of the water and new corals attach farther down. There have been a whole series of ice ages, and so the hillsides on the coast of the Huon Peninsula have a series of benches made of coral

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reefs up and down their slopes (Figure 5.2.2). A living fringing coral reef is in the water along the shore, and the series of raised reefs on the hillside begins with the youngest at the bottom and progresses to the oldest at the top. There are nine fossil reefs, spanning a period of 95,000 years. A comparison of the species on the living reef and in the fossil reefs shows that the reefs have the same species composition even though they differ in age by up to nearly a hundred thousand years. Complex reef assemblages have been able to reconstitute themselves in the same form time after time over a very long period of time. Differences were actually greater between points along the coast than at the same reef over 95,000 years (Pandolfi 1996). Similar areas of rising land in Papua are likely to have experienced the same series of events, and have similar fossil reefs. In areas where the land is not rising or is even sinking, reefs produced during lower sea level stands are likely to be found underwater. Such reefs produce terraces or ledges, such as the one reported from Cenderawasih Bay.

zonation Coral reefs have several zones. The term zonation refers to situations where the type of organisms present (i.e., species composition) changes along some environmental gradient. Rocky intertidal zones in temperate climates have zonation, with some organisms living only high in the intertidal, others living in the middle intertidal, and still others in the lower intertidal. On coral reefs there are a series of zones encountered as one moves out from shore (Figure 5.2.3). On a barrier reef, the zone closest to shore is the lagoon, which has a sandy bottom and may have seagrasses and algae on the bottom, along with patches of coral. Farther out, a shallow, hard calcareous (composed of calcium carbonate) bottom is called the reef flat and may have scattered corals. The crest is where waves break on the reef, and is usually dominated by crustose coralline algae. This type of algae forms a smooth hard layer over underlying coral rubble or rock, cementing it together and

Figure 5.2.2. Coral reef benches on the north shore of Huon Peninsula, Papua New Guinea.

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Figure 5.2.3. Zones on a barrier reef. withstanding the force of breaking waves. From the crest, the reef slopes downward in what is called the forereef slope. On the reef slope there may be a series of ridges and gullies running down the slope called spur and groove, tongue and groove, or ridges and sand channels. On some reefs the slope ends in a vertical drop-off that can be called a wall. Walls commonly have overhangs that alternate with steeply sloping sections. Some reefs have caves on reef slopes or walls, and most reefs have small holes in the reef that may extend in a maze through the reef. At the base of a wall, there is usually a slope of sand and debris, but few or no reef-building corals. Each of these zones is a distinct habitat. The zones differ in exposure to waves and currents, with lagoons being the most protected from waves and sometimes having restricted circulation. Organisms living in lagoons need to be able to burrow in, or attach to, or live on sand, which is the substrate. The reef flat usually has wave action and strong currents from the waves breaking over the crest, pumping water into the lagoon. Organisms here are also subjected to intense solar radiation on clear days. On the crest, organisms are battered with powerful waves and exposed to intense solar radiation. On the reef slope, wave surge and solar radiation decrease with depth. Coral diversity is usually highest on the reef slope, moderate on the reef crest, and lowest on the reef flat (Karlson, Cornell, and Hughes 2004). On walls, wave surge is usually nonexistent, and solar radiation decreases rapidly with depth. There is enough sunlight on steep slopes on a wall for organisms that need light, like coral and algae, to grow. But overhangs do not have enough light for such organisms, and have a strikingly different community of organisms. Overhangs are usually dominated by sponges, soft corals, and coralline algae (which need light but can grow in lower light levels than most corals). Caves have increasingly lower light levels with distance from their opening. Water circulation decreases with distance into caves and holes, and yet is still sufficient for some types of organisms. Oxygen levels may also decrease with distance inside holes in reefs as organisms use up the oxygen coming in on a limited

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flow of water. The zonation with decreasing light in caves may parallel over short distances the zonation on the wall or slope below the reef over a larger depth range. Thus an organism (such as a black coral or sclerosponge) that is found in deep water below the reef may also be found in shallower water under overhangs or within caves. Corals grow most rapidly between depths of about 10 and 30 m. Walls often begin at depths of around 20 m, though in some places they can start in just a few meters (such as several sites in the Philippines) or well below 30 m depth (Discovery Bay, Jamaica). Coral cover is usually highest at depths around 5–20 m, but this does not hold at all locations. Coral cover decreases with depth on most reefs below a depth of around 30 m, though the depth at which this begins is variable. Corals become quite rare most places below about 50 m depth. The deepest corals that require light have been said to be at about 100 m depth, but in Hawai’i living coral has been found as deep as 187 m (Chave and Malahoff 1998). The lower depth of some reefs is determined by habitat, with the reef ending in a sandy slope. Such a sandy slope can be reached at virtually any depth, with some beginning at less than 10 m depth and a few beginning as shallow as 5 m. Many corals have relatively broad depth ranges, yet some are quite restricted in their depth ranges. I found Acropora aspera to be present only on reef flats less than one meter deep in American Samoa, and Acropora cf. pinguis in Malaysia to be totally restricted to depths of less than two meters. Acropora digitifera is common only in shallow water (about 0–3 m). A. robusta and A. pulchra are rare except in shallow water (about 0–7 m deep). Acropora nana, a species with very thin delicate branches, is restricted to shallow water (about 0–2 m) and is somewhat surprisingly most common in heavy surf zones. The genus Leptoseris is largely restricted to low light level areas such as deep water and overhangs, as are the black corals (Antipatharia). Giant clams, Tridacna sp. and Hippopus sp., are most common in shallow water and densities drop off quickly with depth. Coral communities in lagoons may be dominated by corals that are rare on reef slopes and vice versa.

sedimentation Coral reefs are found in warm, shallow, clear tropical saltwater. Corals can only live in saltwater; none are found in freshwater or even brackish water. They are rarely found near the mouths of rivers, and almost never found near large rivers. Most corals thrive best in clear water, and cannot survive in water containing large amounts of sediment or in mud. Most corals require a hard surface to attach to, though there are some corals that do not attach or only attach for a short period in their life cycle. Sediment in the water that settles on a coral can be cleared off by the action of tiny hair-like structures called cilia. The cilia can remove small amounts of sediment but not large amounts. Further, if sediment buildup occurs on the bottom, sediment will begin to cover and smother the coral because the coral is attached and cannot move upward to get above the surface of the sediment. As a result of these processes, coral reefs are not found near the mouths of

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rivers that release huge volumes of suspended fine sediment into coastal waters. In Papua reefs are found around small islands, and along the western mainland coast. However there are no reefs in the eastern part of the province, where the landmass is large and larger rivers such as the Mamberamo, Pulau, and Digul empty into the ocean. Because the island of New Guinea is a geologically young, very high island in an area of very high rainfall (over 3,000 mm/yr in Papua; Tomascik et al. 1997), runoff of fresh water and sediment is very high. If the temptation to reap large quick profits by cutting the rainforests of Papua is not resisted, the amount of sediment runoff will grow many-fold, and coral reefs will be rapidly killed. Reef building corals need light to live and grow. Suspended sediment in the water scatters and absorbs light, reducing its availability to coral. Although there are limits to how much sedimentation corals can tolerate, some are able to thrive in areas with moderate levels of sedimentation. A coral reef named Middle Reef about a kilometer offshore of Townsville, Queensland, Australia, survives with high coral cover in water that has a visibility of only about one to two meters. The coral species on this reef are quite different from those found on the nearby outer edge of the Great Barrier Reef, in clear oceanic waters. Further, sediment input to the nearshore waters of Queensland increased dramatically when sheep and cattle were introduced to the area about 100 years ago (McCulloch et al. 2003). This increased sedimentation stresses the surviving corals and renders them less resilient, perhaps to the point of being unable to recover from other types of disturbance.

temperature Coral reefs are restricted to warm waters where the minimum temperature is above about 18C. The world’s most northern coral reefs are at Kure Atoll in the northwest Hawai’ian Islands, and in Japan, and the most southern are at Lord Howe Island, off southeastern Australia. Some coral communities can be found at even higher latitudes, but they do not accumulate calcium carbonate, and therefore do not form coral reefs. An example is in the Solitary Islands off New South Wales, Australia, where occasionally storms with waves up to ten meters tall sweep most corals off into deep water. Newly settled coral recruits then grow on the non-carbonate rocks, rebuilding the coral community, but their skeletons do not accumulate (Harriott, Smith, and Harrison 1994). Such coral communities can also be found in the tropics near the equator in marginal environments, such as sandy, high sediment, or low circulation areas. In addition, at the extremes of latitude, coral reefs tend to be small, and to have low diversity (Yamano et al. 2001, discussed below). Papua is situated in an area close to the equator, where temperatures are warm year-round (27.5–28.5C in the Java Sea; Tomascik et al.1997) and nearly ideal for coral reefs. Most coral reefs are found in warm, clear tropical water. The water is clear because it contains few of the tiny drifting plants and animals, which together are known as plankton. Temperate and polar waters are often opaque with a green

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or brown color, due to masses of plankton. The tiny drifting plants are called phytoplankton, and they require nutrients such as nitrogen and phosphorous, just as do other plants. In the tropics, the hot sun heats only the surface water but does not penetrate deeper water. Hot water rises above cold water because it expands slightly when heated. The boundary between the warm surface water and colder deep water begins at a depth of about 50 m in Indonesia, and extends down to about 300 m depth (Tomascik et al. 1997). The fact that warm surface waters float on top of the cold deeper water means that these two bodies of water do not mix. Phytoplankton absorb nutrients from the surface water as they perform photosynthesis and grow. The phytoplankton are fed on by zooplankton (tiny drifting animals), which are in turn fed on by larger animals in a food chain or food web. When any of these organisms die, they sink slowly down into the deep cold water and to the bottom, taking nutrients with them. While currents called upwelling bring nutrients from the bottom back up to the surface fueling blooms of plankton in temperate and polar waters, upwelling is rare in tropical waters. The result is low nutrient levels in warm tropical surface waters, low densities of plankton, and hence clear water.

zooxanthellae The low nutrients in warm, clear, shallow tropical waters pose a paradox for coral reefs. Coral reefs have abundant organisms, and high rates of photosynthesis and growth. How can this occur in waters that are low in nutrients? How can this oasis flourish in such a biological desert? A variety of mechanisms probably contribute, but perhaps the most important is the algae living in corals. Corals are animals related to sea anemones and jellyfish. Corals have small polyps that are nearly identical to sea anemones. However, they contain within them the seeds of their success. These are the tiny single-celled algae known as zooxanthellae. The algae are members of a group called dinoflagellates. The algae and the coral animals live in a mutualistic symbiosis (i.e., mutually beneficial coexistence). The waste products of the animal contain the nitrogen and phosphorus that the algae need. The algae perform photosynthesis in the sunlight, and leak much of what they produce into the surrounding animal cells (around 80%). So the algae benefit from the nutrients the coral animal produces, and the coral benefits from the food that the algae produce. In effect, this is a tight recycling arrangement, with nutrients passed from one partner to the other, and then back to the other partner. As a result, the combination of these two partners needs a smaller input of nutrients from the outside, and can survive in nutrient-poor, warm, clear, tropical waters.

Plants on Coral Reefs On most coral reefs, animals are obvious and appear to be common, while plants are less obvious and appear to be less common. But only plants can produce food through photosynthesis. When animals eat plants, most of the food is used to produce energy to run the animal’s bodily functions, while only a small part is

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added to the material of the growing animal. As a rule of thumb, only about 10% of what an animal eats is incorporated into its body in growth. As a result, there must be about ten times as much biomass of plants as herbivores that eat them. And there must be about ten times as much herbivore mass as carnivore mass that eats them, and so on up the food chain. Thus there must be about 100 times as much plant biomass as carnivore biomass for carnivores that eat herbivores; 1,000 times as much to support carnivores that eat other carnivores. And yet on coral reefs animals are obvious and appear abundant, while plants are usually less obvious and appear less abundant. How can this be? First, some plants are hidden. Zooxanthellae are found in hard corals, soft corals, giant clams, and a few other animals on coral reefs. They provide a large part of the food production on a coral reef, and yet are not obvious. They actually provide much of the color in corals and giant clams, yet we normally don’t recognize them as plants. Second, the algae which can be seen on coral reefs include some species which are small and hard to see but are highly productive. Large fleshy algae grow slowly and put most of their growth into defenses such as woody cellulose that is hard to digest, calcium, and chemicals that are bad tasting or toxic. Defenses are necessary for a plant to grow large on a coral reef, since there are many hungry mouths of herbivores, such as fish, sea urchins, and snails. Fish alone bite algae about 40,000 to 156,000 times per square meter of reef per day! So herbivory is intense on coral reefs. A second group of algae is the filamentous algae. These algae are made of tiny strings of cells with little or no defense. Their main defense is their ability to grow very rapidly. Herbivores bite most of their growth off daily or hourly, but the base of each filament attached to rock rapidly grows the string back. So filamentous algae are highly productive fast growing algae, but have a very low standing biomass and are hard to see.

Species Diversity Coral reefs are not only geological structures, but also biological communities. Coral reefs are amazingly diverse and complex ecosystems. They are the most diverse marine ecosystem known (i.e., they are the most species-rich). Sometimes they are said to be the most complex ecosystem on the planet, but they actually have fewer species than tropical rainforests. Rainforests have large numbers of insect species, and insects are by far the most species-rich group of organisms on the planet. There are more insect species known than all other organisms combined, and there are more insects in tropical rainforests than anywhere else on earth. There are very few marine insects, and none known on coral reefs. The total number of species is not known for either coral reefs or rainforests. Around 1.8 million species have been described on earth, with a majority of those being insects. Estimates for the total number of species on earth range from about 3 to 120 million, with about 5–10 million being most likely. About 85% of all species are arthropods, and a majority of those are insects. On coral reefs, one estimate is that 93,000 species may have been described, but the total may be ten times higher

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(Reaka-Kudla 1995a,b). However, at the level of the largest groups of animals, known as phyla (such as Mollusca, Arthropoda, and Chordata) there are more phyla of multicellular animals (metazoa) in the oceans (about 29 out of 32) than on land (about 13) and in freshwater (about 16) (Rupert and Barnes 1994), and coral reefs probably have more phyla than any other ecosystem (about 26).

Relationships among Organisms Coral reefs have very large numbers of species living together and interacting in a very complex web of relationships. We have already spoken of the mutualistic relationship between corals and the zooxanthellae they host. There are many other examples of coral reef species that have mutualistic relationships with other species, such as the Anemonefish (Amphiprion) that live among the tentacles of sea anemones; cleaner fish, such as Labroides dimidiatus, and shrimp, such as Periclimenes, that clean parasites off of fish; snapping shrimp (Alpheids) that excavate burrows which they and guardian prawn gobies (Gobiidae), occupy, and the Guard Crabs (Trapezia) that live in the branches of corals (Pocillopora) and defend them. In another type of symbiosis, commensalism, one organism lives on another and benefits, while the host organism is neither helped nor hurt by the relationship. An example is the shrimp Periclimenes that live on sea anemones, starfish, nudibranchs (sea slugs) and other animals. A third type of symbiosis is parasitism, where one partner benefits at the expense of the other. Many small crustaceans and flatworms live on the skin of fishes, eating mucus and tissue off of the fish. Some crustaceans called isopods live attached on the outsides of fish, sucking tissue and blood. There are so many species of flatworms that are parasitic on snails in at least one stage of their life cycle that it is said that nearly every species of snail is parasitized by a species-specific parasitic flatworm. One parasitic flatworm, Plagioporus, lives in corals, Porites, at one stage in its life, causing the host polyp to expand and turn pink. The larger pink polyp stands out and is often eaten by butterflyfish (Chaetodontidae). This transfers the flatworm to its next host (Aeby 1991). New relationships like these are being discovered all the time. For example, the snail Dendropoma maxima produces an uncoiled shell in a coral. The snail, called a vermatid because its shell resembles a worm tube, secretes mucus, which it drapes over the coral surface. The snail pulls the mucus in, dragging along with it additional mucus produced by coral, and eats it. The snail is thus parasitic on the coral, and stunts the growth of the surrounding coral (Fenner, under review a). Infectious diseases are similar to parasites, except that the agents that cause the diseases are generally microorganisms such as protozoa, bacteria, and viruses. Coral diseases have increased in number and severity in recent years, both on Caribbean and Indo-Pacific reefs. Diseases were detected at 10 of 45 (22%) sites during a recent survey at the Raja Ampat Islands (McKenna, Boli, and Allen 2002). Nearly all animals on coral reefs are either predators or herbivores, the exceptions being those hosting endosymbiont algae such as corals. Predation may shape

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some communities, with apex predators dominating fish communities in pristine coral reefs such as the northwest Hawai’ian Islands. Predators may increase diversity by preying on the most abundant species. Two important predators of corals are the Crown-of-Thorns (COTS) starfish (Acanthaster planci) and snails of the genus Drupella. Crown-of-Thorns starfish came to the attention of scientists and the public in the mid-1960s when there were large outbreaks on the Great Barrier Reef in Australia, and nearly all corals were eaten and killed on some reefs in the midsection of the Great Barrier Reef. Outbreaks were reported on many reefs in the Indo-Pacific, with some of the worst around Okinawa, Japan (Moor 1989). Reports of outbreaks are much less frequent today, and reefs around Okinawa are now largely free of outbreaks. Outbreaks continue to occur on the Great Barrier Reef periodically. The first outbreak of COTS reported in Indonesia occurred in 1995, on the reefs of the Seribu Islands south of Papua (Tomascik et al.1997). In the Raja Ampats, only 3 of 45 (6.7%) sites had any COTS at all (McKenna, Boli, and Allen 2002). A few outbreaks of the snail Drupella have been reported in the Indo-Pacific (Moyer, Emerson, and Ross 1982), but they have not done as much damage as Crown-of-Thorns starfish. In the Raja Ampats study, Drupella were only observed at one site.

The Coral Triangle: The Peak of Diversity The amazing diversity of species is perhaps the most notable aspect of the coral reefs of Papua Province. Papua is situated within the area that has been called the ‘‘Coral Triangle’’ (e.g., Allen 2002a; Wells 2002). This is the area of the highest diversity coral reefs in the world (Figure 5.2.4). It includes the Philippines, central and eastern Indonesia, and Papua New Guinea. There are more coral species in this area than in anywhere else in the world (Hughes, Bellwood, and Connolly 2002; Stehli and Wells 1971; Veron 1995), and the same is true of fish (Allen 2002a; Hughes, Bellwood, and Connolly 2002; Chapter 4.8), mollusks (Gosliner 2002; Wells 2002), and sponges (van Soest 1997). A recent expedition to the Solomon Islands found that the diversity of corals, fishes, and mollusks there is equally high,

Figure 5.2.4. Map of coral species diversity. Coral diversity decreases in all directions from the Coral Triangle (indicated in the darkest shade), which contains 581 coral species. Source: Reproduced from Veron (2000) with kind permission from the author.

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indicating that it too is part of the Coral Triangle (Green, Veron, and Allen 2004; Wilkinson 2004). Further, the number of mollusks in the Coral Sea area (Solomons to Great Barrier Reef to Vanuatu) is nearly as high as in the Coral Triangle (Wells 2002). In addition, recent work has found that diversity levels on outer barrier reefs of the northern Great Barrier Reef were marginally below that typical in the Coral Triangle (Fenner, unpublished). Coral diversity on mid-shelf reefs is even higher (Done 1982), so it is likely that the northern Great Barrier Reef has coral diversities within the range of diversities of the Coral Triangle.

diversity gradients The number of species decreases in all directions from the Coral Triangle (Figure 5.2.4). The diversity gradient to the north and south from the Coral Triangle is called the Latitudinal Diversity Gradient. This is perhaps best illustrated by the gradient in species diversity in southern Japan, with 342 species of coral in the small islands just north of Taiwan, decreasing almost in a straight line to zero species around Tokyo (Figure 5.2.5). A similar diversity gradient extends eastward from the Coral Triangle, with the number of species decreasing to the east in the Pacific until in the eastern Pacific there are a total of only about 33 species over a very large area. This gradient is called the Longitudinal Gradient. Both the Latitudinal and Longitudinal gradients have been documented in several groups of organisms. For instance, Indonesia has 90 species of Crinoids (feather stars), and going north, Palau has 30 and Guam has six. Going east, the Marshall Islands have 14 species and Hawai’i has none. Similarly, there are 536 species of sea slugs (opisthobranchs) known from the north coast of Papua New Guinea, 410 in Guam, 244 in Hawai’i, and 183 in Pacific Panama (Gosliner 1992). The diversity gradient for corals and fish shows a less steep gradient going west from the Coral

Figure 5.2.5. Coral diversity in Japan decreases with increasing latitude (n  10, R2  0.96). Source: Redrawn from Veron (1992).

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Triangle in the Indian Ocean. Recent work in the Red Sea by Emre Turak, Lyndon Devantier, and J. E. N. Veron (reflected in the maps in Veron 2000) has doubled the number of species known there from 150 to 300, showing that the Indian Ocean Longitudinal Gradient is not as steep as previously thought. A recent study (Karlson, Cornell, and Hughes 2004) compared coral species richness on a longitudinal gradient, comparing the number of species in Indonesia, Papua New Guinea, the Solomon Islands, American Samoa, and the Society Islands of French Polynesia. There were three sites in each country, with one of the sites in Indonesia being an island site to the west of the Vogelkop Peninsula of Papua. They separated transects on reef slopes, reef crests, and reef flats, and had an equal number of transects at each location and each reef zone. They found that on reef slopes, there was a high diversity from Sulawesi to the Solomon Islands, with no gradient between these sites. American Samoa and the Society Islands had significantly lower diversity on reef slopes. On reef crests, the diversity was highest in Indonesia, decreasing significantly in Papua New Guinea, decreasing further in the Solomon Islands, and lowest in American Samoa and the Society Islands. Reef flat diversity decreased a lesser amount from Indonesia to the Solomon Islands and on to American Samoa and the Society Islands (Figure 5.2.6). Thus, while the reef slope data indicate that the reefs of Papua New Guinea and the Solomon Islands are as diverse as those in Indonesia, the reef flat data and especially the reef crest data indicate that Indonesia has the highest diversity of all these areas, and that the longitudinal diversity gradient begins between Papua and Papua New Guinea. The authors did not separate the data for Papua from the other two Indonesian sites on Sulawesi. Borel Best et al. (1989) proposed that western Indonesia is outside of the Coral Triangle. Allen (2002b) found a lower diversity of fishes at Weh Island off the western end of Sumatra, which is consistent with this

Figure 5.2.6. Coral species richness as a function of longitude in three reef zones. Leftmost points are for Indonesia, followed to the right by Papua New Guinea, the Solomon Islands, American Samoa, and the Society Islands. Source: Redrawn from Karlson et al. (2004).

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proposal. Thus, the best current data indicates that Papua is in the area of highest coral diversity, but diversity begins to decrease to the east of Papua.

Why Is Diversity So High? The causes of the peak of marine diversity in the Coral Triangle and the latitudinal and longitudinal diversity gradients have been much debated. There have been many proposals. An early idea was that the center of diversity was a center of species formation (e.g., Briggs 1994). A second view is that more rapid extinction in outlying areas reduces the number of species in those areas. During ice ages, for instance, the Coral Triangle area probably experienced a much smaller drop in sea surface temperature than high latitudes and the eastern Pacific, so more coral species survived in the Coral Triangle than in those other areas. Another proposal is that currents in the tropical Pacific flow westward, carrying new species with them and causing the accumulation of species in the Coral Triangle area (Jokiel and Martinelli 1992). The proponents constructed a model that showed just this effect. Yet another proposal is that many islands close together allow any local populations that might go extinct to be rapidly replenished by larvae from nearby islands. The classic theory of island biogeography predicts higher numbers of species when an island is closer to a source of additional species (MacArthur and Wilson 1967). In areas with few islands, a population could go extinct more often on an isolated island before larvae from distant islands could reach it by chance and replenish the local population. Local extinctions have indeed been documented among the corals on the widely separated reefs of the eastern Pacific (Glynn 1977). A model using different densities of islands but random currents produces higher diversities in areas with more islands (Blanco-Martin 2002). Connell (1978) proposed an ‘‘Intermediate Disturbance Hypothesis’’ to account for high diversity in rainforests and coral reefs. Disturbances of intermediate intensity and frequency open spaces where additional species can settle, whereas without disturbance, superior competitors drive out inferior competitors in a biological succession that ends in lowered diversity. If this theory were used to try to explain diversity gradients, it might suggest that the area of highest diversity is where disturbances are of intermediate intensity and frequency. However, sea surface temperatures in Indonesia and New Guinea are disturbed less by cold events than in areas farther from the equator. Similarly, there are no cyclones near the equator in Indonesia (including Papua) and New Guinea (Figure 5.2.7a,b; Fenner and Riolo, under review). On the other hand, the northern Philippines experiences a moderate to high level of disturbance from cyclones, and yet is part of the Coral Triangle. Cyclones are probably one of the most important natural disturbances on coral reefs (Rogers 1993). Thus, the Intermediate Disturbance Hypothesis is unlikely to explain coral diversity gradients. In ecosystems with low diversity, each species may be represented by large numbers of individuals. For example, in the tundra of northern Canada and Alaska, there are millions of Snow Geese (Chen hyperboreus) in the summer, and millions

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Figure 5.2.7. a. Tracks of tropical cyclone and severe storm tracks for the Indo-Pacific region. All storms that were classified above a tropical depression in strength (wind speeds ⬎30 mph for two or more 6-hour periods) from 1945 to 2003 are included. b. Severe storm density in the Indo-Pacific region. Storm density was computed for each 50  50 km cell by summing the number of tracks found within 200 km of the cell and dividing by the total area sampled in each cell. Source: Data assembled by Unisys Corporation and Joint Typhoon Warning Center (www.npmoc.navy.mil/ jtwc.html) and downloaded from the Pacific Disaster Center website (atlas.pdc.org). Maps by F. Riolo.

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of mosquitoes. There is only one large mammal, the Caribou (Rangifer caribou) and it is also present in large numbers. Northern forests are composed of large numbers of individuals of a small number of tree species. By contrast, in the tropics there are large numbers of species, most of which are rare. In a hectare of rainforest in New Guinea, there can be several hundred species of trees, but few individuals of each species. An example from coral reefs is sea slugs (opisthobranchs): in the western Pacific there are many species (over 500), most of which are very rare. In the tropics, functional groups of species, called guilds, usually have many more species than in the temperate or, especially, polar areas. Research on terrestrial systems indicates that the loss of individual species does not have a great impact on a high-diversity ecosystem, because there are many other members of most guilds that can continue to perform that guild’s functions (Grime 1997; Moffat 1996). The loss of a member of a guild in a low-diversity ecosystem may have a much larger impact on the ecosystem, particularly if there is only one member of that guild so that with its loss the guild function is no longer performed. Similarly, Bellwood et al. (2004) have argued that low diversity coral reefs are more vulnerable to the loss of individual species than diverse ecosystems for these reasons. For example, the loss of a single species of sea urchin, Diadema antillarum, in the Caribbean in 1983–1984 (Lessios 1988; Lessios, Robertson, Cubit 1984), led to major phase shifts on some reefs from coral-dominated reefs to algal-dominated hard grounds. The large numbers of a single species in lowdiversity ecosystems also makes them more vulnerable to diseases and specialized predators. A dense population of a single species, as was the case with D. antillarum, makes the transmission of disease easier. The die-off of D. antillarum was the largest marine epizootic ever recorded. Similarly, two of the most common coral species in the Caribbean were Acropora palmata and A. cervicornis. Both form large, dense, single-species thickets of genetically identical organisms, or clones. The lack of genetic diversity means that any disease that can kill one individual can kill the whole clone. Both of these species have been decimated in much of the Caribbean by White Band disease (Aronson, Precht, and Macintyre 1998), and were considered for Endangered Species status (Precht, Robbart, and Aronson 2004; Shinn 2004; Wilkinson 2004); they received protected status on 8 June 2006. On diverse coral reefs, most species are rare, so disease transmission is much more difficult, and the likelihood of epizootics is reduced. This probably contributes to the stability of high diversity coral reefs.

Effects of Fishing Fishing can remove fish that are important for reef health. The removal of herbivorous fish in Jamaica left it vulnerable, so when a disease killed the last herbivore (sea urchins), the reef was overcome with algae (Hughes et al. 1987). Large fish such as sharks, Humphead Wrasse (Cheilinus undulatus), and Bumphead Parrotfish (Bulbometopon muricatum) are particularly vulnerable. Bumphead Parrotfish have been extirpated from several places in the Indo-Pacific (Bellwood et al. 2003;

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Dulvy et al. 2003). Humphead Wrasses are under heavy pressure over a large area due to the live food fish trade. Populations are reduced to low levels in areas with higher fishing pressure (Sadovy et al. 2003). Areas of Fiji where fishing pressure is greatest are also the areas where Crown-of-Thorns starfish have outbreaks and eat the tissue off corals, killing the corals (Dulvy et al. 2004). Humphead Wrasses are known to eat toxic invertebrates like the Crown-of-Thorns starfish, so overfishing them may leave reefs vulnerable to Crown-of-Thorns attacks. Although the reefs of Papua are remote and under relatively little pressure from human populations (but see Birkeland 1982), fishing makes sharks and Humphead Wrasse rare, with just two adult Humphead Wrasse seen in 45 sites (Allen 2002b; McKenna et al. 2002). Few coral reef researchers or managers have seen what truly pristine coral reef fish populations are like, and the amazing dominance of apex predators. Each generation of scientists remembers what coral reefs were like when they first saw them, and tend to think of that as the standard of undisturbed ecosystems. We have lowered our standards in a process called ‘‘shifting baselines.’’ Coral reefs in the Caribbean have declined in three decades from about 50% coral cover to about 10% coral cover (Gardner et al. 2003). Archeological methods have been used to study the effects of pre-Columbian fishing in the Caribbean, and the studies have found that declines began even before the arrival of Europeans (Wing and Wing 2001). Paleontological methods along with archeological and historical methods have shown declines in 14 coral reef systems worldwide since pre-human times (Pandolfi et al. 2003). The 14 reef systems studied had declined between about 28% and 78% of the way from pristine toward ecologically extinct.

Endemism and Extinction Endemism is commonly used in terrestrial conservation programs as a measure of the need to conserve areas (Allen 2003). It is especially important to avoid the local extinction of endemic species, because their loss represents the loss of an entire species. Endemic species are more vulnerable to extinction partly because any local disturbance can cause global extinction, and also because endemic species usually have small populations. The rates of endemism on coral reefs are quite different in different groups of organisms. Endemism is uncommon in larger organisms, but may be high in some groups of small organisms, and low in the tiniest microscopic organisms. Most groups of larger coral reef organisms have wide dispersal and very few species are endemic. Many coral reef species are broadcast spawners, releasing tiny eggs into the water that are carried with the currents. Currents can carry the eggs considerable distances during the several days to weeks required for them develop to the stage where they are ready to settle. For example, one species of sea urchin, Echinothrix diadema, has been found to be genetically the same species in Hawai’i and the east Pacific, across the largest expanse of open water in the tropics anywhere in the world (Lessios et al.1998). Some coral species have been observed attached to floating objects and thus are

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RAPID ASSESSMENT TECHNIQUE CONFIRMS PAPUA IS IN THE CENTER OF DIVERSITY Species diversity comparisons among areas are often based on the total number of species that have been found in each area. However, the number of species found in an area is heavily dependent on the amount of time, effort, and area covered searching for species. Additional searching time, effort or area explored almost always leads to additional species being found. Larger areas contain larger numbers of species, which is called the ‘‘species-area effect.’’ The number of species commonly rises as a power function of the area, as it did in a study of coral reef fishes (Chittaro 2002). Although such curves may appear to approach an asymptote on linear scales, on log scales they can be seen not to approach an asymptote. The search for an asymptote has been reported at times to reach areas the size of continents without reaching an asymptote (Williamson et al. 2001). The total number of coral species known from countries in the western Pacific has approximately doubled in the last three decades (Fenner, in review c). The author has participated in several rapid assessment programs for coral reef areas, such as those sponsored by Conservation International. The goal of such programs is to rapidly assess diversity in an area. It is an attempt to use limited resources in a targeted fashion, to gain information about diversity of an area without spending the enormous resources necessary to get even a near-complete assessment. In the present study, one scuba dive of approximately 60 minutes was spent by the author at each site in a roving search for coral species. The search began at the bottom of the reef or at about 30 m depth, whichever was less, and progressed upward during the dive, ending in the shallowest area that was accessible to a scuba diver. Comparisons among areas were based on equal numbers of dives. A strong latitudinal gradient was found in the central Pacific, with diversity falling off from eastern Papua New Guinea to American Samoa and Hawai’i (Figure 5.2.8). The Raja Ampat Islands are in the area of highest diversity. Across Malaysia to Rodrigues in the southwestern Indian Ocean, there is also a latitudinal diversity gradient (Figure 5.2.8), though Rodrigues may be lower in diversity than Malaysia both due to latitude and longitude. Diversity gradients are well known in the Pacific for corals and other groups of reef organisms. This rapid technique detects diversity gradients in much the same way as more labor intensive techniques. The lack of a gradient in this reef slope data between central Indonesia and New Guinea is consistent with the report that coral diversity on reef slopes is constant across this area (Karlson et al. 2004).

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Figure 5.2.8. Longitudinal diversity gradients from rapid ecological assessments. Raja Ampat Islands, Papua, indicated by the open square, is among the areas of highest diversity. Points, in order from east to west, are for Rodrigues, Andaman Islands, Peninsular Malaysia, Sarawak, Sabah, Sulawesi, Raja Ampat Islands, Milne Bay (Papua New Guinea), Fiji, American Samoa, and Hawai’i.

probably able to ‘‘raft’’ over vast distances (Jokiel 1990). This is even true of species that brood their young, releasing larvae that quickly attach close to the parent. Reef fish have also been observed rafting by staying near floating debris (Mora 2001). This wide dispersal means that there are few endemic species on most coral reefs, especially in the western Pacific where there are many reefs close together. For instance, currently no endemic coral species are known in the Philippines or Indonesia, where 535 and 581 coral species are currently known, respectively (Fenner, under review c; Turak 2003; Veron 2002), and only one endemic species is known from Papua New Guinea, where 494 coral species are currently known. Levels of endemism in reef fish are also relatively low (Hughes, Bellwood, and Connolly 2002), and the proportion of reef fish that are endemic is lower in the Coral Triangle than in outlying areas (Randall, 1998). Endemism may be overestimated if recently described species are included, because species are often described from small areas and subsequently found in additional areas (Fenner, under review b). So it is likely that Papua has very few endemic large species on its coral reefs. Most coral reef organisms that have been studied have relatively large individuals. Marine invertebrate species with small individuals frequently brood their off-

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spring instead of broadcast spawning (Reaka-Kudla, 1995a,b). This may be because their small size restricts them to producing relatively small numbers of offspring, and broadcast spawning is a high-risk strategy in which most offspring die. If a small number of offspring are produced, a high-risk strategy increases the likelihood that all offspring will die. This may select for lower-risk reproductive strategies, where more is invested in each offspring by producing larger offspring, which do not disperse as far. This reduced dispersal ability increases the frequency of endemism (Reaka-Kudla, 1995a,b). A good example may be the amphipods, a large group of small-bodied crustaceans that produce relatively large eggs. Some species of amphipods raft on algae or have pelagic hosts such as jellyfish, and thus have wide ranges. But many amphipod species have very small ranges (Thomas 2000). Most species are small, such as insects on land. The view that most marine species have wide ranges is largely based on larger organisms like corals, fish, and echinoderms. Yet most coral reef species are likely to be small (Reaka-Kudla, 1995a,b), and not yet described, let alone have their reproductive mode or biogeographic range studied. Many or most of these species may turn out to be endemics. In addition, some groups of larger organisms that do not have a larval dispersal stage may have high rates of endemism. For example, many or most reef sponges produce negatively buoyant, sticky eggs that do not go far from their parents. An estimated 43% of the sponges recorded from Indonesia are endemic to the region (van Soest 1997). However, studies of Indonesian sponges and sponge biogeography are in early stages. The total number of sponge species is likely to increase considerably and endemism figures to change. Another example is colonial ascidians (sea squirts), which produce tadpole larvae that go only short distances from their parents. As with corals and fish, the actual ranges of these species may be largely determined by their ability to raft, because rafting can spread widely even species with no larval dispersal phase. A very different view is presented by Fenchel and Findlay (2004). They report that most microbial marine organisms are cosmopolitan, and that the percentage of species found at one temperate marine location that have large ranges decreases with increasing body size. If this should also prove true of coral reefs, then groups of small organisms that are highly endemic, like amphipods, may be unusual. It may be that only groups like amphipods, sponges, and ascidians have high rates of endemism on coral reefs. Or it may be that that the largest organisms and microbes have low endemism, but intermediate size (small) organisms have high rates of endemism. If there are large numbers of small, undescribed, and unstudied species on coral reefs that are likely to be endemic, it will not be practical to study each species to determine its range. We will not know the ranges of even a fraction of the small species on coral reefs any time in the near future. Long before we can know which species are endemic, coral reefs may be highly degraded, and endemic species lost before they are even discovered. The primary cause of species extinctions is loss of habitat. The best way to save large numbers of small endemic undescribed coral reef species is to protect the habitat itself, without taking the time to discover all

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the tiny endemic species. Further, the number of small species present at a site is almost certain to be proportional to the number of large species found there. Bellwood and Hughes (2001) found that there is a high correlation between the diversity of organisms in one size group with those of another size group. Thus, the diversity of large species such as corals and fish is likely to be a good indicator for the diversity of small species. Although we know that low diversity coral reefs have a higher proportion of endemic species among large organisms than high diversity reefs, high diversity reefs are likely to have much higher absolute numbers of small endemic species than low diversity reefs. Thus, the conservation of both low and high diversity coral reefs is important. Further, while large species (‘‘charismatic megafauna’’) may capture public support, small endemic species will not (‘‘save the amphipods’’?). Coral reefs, however, are highly charismatic, and have generated significant public support for conservation.

Threats to Papuan Reefs Coral reefs face threats from a wide variety of human sources. The ‘‘Reefs at Risk’’ program (Burke et al. 2002) identified six principle threats to coral reefs, and evaluated five of those. The five threats they evaluated were coastal development, marine-based pollution, sedimentation and pollution from inland sources, overfishing, and destructive fishing. The sixth threat was climate change and coral bleaching. Their method was to identify sources of human pressure that produce stress on coral reefs and represent these sources of stress on a map. They developed distance-based rules by which the level of threat declines with distance from the source of the stressor, such as the distance from a river mouth, city, and so on. For Indonesia, destructive fishing (i.e., blast fishing) turned out to be the biggest threat to coral reefs, followed by overfishing, sedimentation, marine-based pollution, and coastal development, in that order. Climate change was not evaluated because of the lack of data and inability to predict strong local variations in this relatively new threat. The Reefs at Risk program identified the reefs most at risk in eastern Indonesia (Burke et al. 2002). In Papua, reefs in the Raja Ampat area to the northwest of the western end of the Vogelkop Peninsula were shown as being under medium threat, while islands just to the south of that and straight west of the western end of the Vogelkop Peninsula, beginning with the Fam Islands and Batanta Island, were shown as being high or very highly threatened. Biak and Yapen Islands on the north side of Cenderawasih Bay were shown as experiencing medium threat, with reefs at the eastern end of Biak viewed as being under low threat. Reefs along the western side of Cenderawasih Bay were shown as having low threat, while those on the east side of the bay were shown as having high or very high threat. Reefs along the Onin Peninsula and just to the east of that on the south shore of Papua were shown as having medium to low threats. The Aru Islands south of the main landmass of Papua were shown as having high or very high threats. The primary

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and almost only immediate threat to the coral reefs of Papua was identified as destructive fishing. The two types of destructive fishing that were identified were poison fishing and blast fishing. Poison fishing today uses cyanide to stun and catch fish, and blast fishing utilizes homemade explosives made of fertilizer in bottles. The Conservation International 2002 Rapid Assessment of reefs in the Raja Ampat Islands found evidence of destructive fishing practices at 13% of the sites visited. Slight fishing pressure was evident at 32 of the 45 sites, and moderate pressure observed at one site. A total of only seven sharks, two Manta Rays (Manta birostris), and only one sea turtle (Hawksbill: Erectmochelys imbricata) were observed by the reef condition team on these 45 sites. Humphead Wrasses were much less common than on less heavily fished sites (Table 5.2.1). Sixteen sites had slight siltation and one site had moderate siltation, with seven of these sites having freshwater input as well. Slight evidence of eutrophication/pollution was observed at eight sites. Nickel mining has been proposed for Gag Island, and there has been some logging of forests. Other stressors such as coral diseases, coral predators, and bleaching were rarely observed (McKenna, Boli, and Allen 2002). In the Raja Ampats, 90% of the inhabitants lived in coastal areas and depended on marine resources for survival. Humphead Wrasse and groupers (Plectropomus leopardus and P. areolatus) were targeted for the live fish export trade, and cyanide was used to catch them. Shark fins were taken for export, and shark finning was the likely cause for the rarity of sharks. Small sea cucumber and lobster fisheries existed. Illegal fishing methods were used by the poorer communities (Amarmollo and Farid 2002). Although destructive fishing was the main threat to the coral reefs of Papua, we cannot assume that the other threats have not had effects on the coral reefs there. The study by Pandolfi et al. (2003) showed that all 14 coral reefs that they studied around the world showed degradation from human activities, and

Table 5.2.1. Frequency of Napoleon Wrasse (Cheilinus undulatus) for locations in the Indo-Pacific Location

No. sites where seen

% of total sites

Approx. no. seen

Phoenix Islands (2002)

47

83.92

412

Milne Bay, PNG (2000)

28

49.12

90

Milne Bay, PNG (1997)

28

52.83

85

Raja Ampat Islands (2002)

9

18.00

14

Raja Ampat Islands (2001)

7

15.55

7

Togian/Banggai Islands (1998)

6

12.76

8

Calamianes Is, Philippines (1998)

3

7.89

5

Weh Island, Sumatra (1999)

0

0.00

0

Source: Allen (2003).

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even the Great Barrier Reef, long thought to be relatively unaffected by human activities, was impacted. Fishing has been allowed on most of the Great Barrier Reef, and even though it is not intense, it has doubled since 1990 and has had an effect. Fish populations in small, strictly protected (i.e., no-take) areas on the Great Barrier Reef are higher than outside those areas. The Raja Ampat Islands to the northwest of the Vogelkop Peninsula have relatively good fish stocks. The average total biomass of fish in 2002 was 209 tons/km2, compared to 124 in Milne Bay Province, Papua New Guinea, 66 in the Togian-Banggai Islands of northern Sulawesi, Indonesia, and 17 in the Calamianes Islands, Philippines (La Tanda 2002). Further, the mean density of groupers was 5.45 per 1,000 m2, compared to 3, 2.7, and 2.9 for each of the other three areas, respectively. The average size of groupers was relatively small (about 25 cm), while at the other three sites they were 20 cm, 20 cm, and 30 cm, respectively. Humphead Wrasses were uncommon (Table 5.2.1), with most individuals under 30 cm length. This species is intensively harvested in this area for export in the live-fish restaurant trade. Further, the reef fish populations are not dominated by apex predators, such as jacks and sharks. The low populations of apex predators in the Raja Ampats indicates that fishing pressure has already made major changes to the structure of fish populations there, while the total biomass indicates that fishing has not yet caused drastic changes to fish populations such as have occurred in many other places in the region. The long-term future threats to the reefs of Papua are not restricted to destructive fishing and overfishing. If population growth and population transmigration to Papua from western Indonesia continue or intensify, stresses to the coral reefs of Papua will increase with the increasing population and development. Deforestation is currently proceeding rapidly in most other parts of Indonesia, with deliberate forest burning during dry summers causing huge smoke clouds covering large areas where there are intense clearing efforts, such as Kalimantan and Sumatra, with clouds of smoke so large they can be carried to neighboring countries. In the major fires of 1997, smoke from fires in Sumatra provided iron that helped lead to a red tide event that in turn caused the death of coral reefs by asphyxiation. There has been no analogous coral mortality in that area in the last 7,000 years (Abram et al. 2003). Growing world populations and demand for wood and agricultural land can make logging and burning so attractive that it can become common in spite of being illegal. Not long ago, Kalimantan (Indonesian Borneo) was a wild and remote area, but now it is being cleared at an alarming rate. A similar fate could await Papua, followed by massive sediment runoff and reef destruction (McKenna, Boli, and Allen 2002). The low population density and relative underdevelopment of Papua affords time to try to avert coral reef destruction, but complacency could result in much greater threats to reefs in Papua in the future. Saving the coral reefs of Papua will require the assistance of many people, but luckily Papua’s reefs remain in better condition than many reefs elsewhere in the world. This provides conservationists and managers with a rare opportunity to be

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proactive in conservation activities. It remains to be seen whether we will be able to take advantage of this opportunity and protect Papua coral reefs for future generations.

Acknowledgments I thank Conservation International for inviting me to participate in their expedition to Papua, Gerry Allen for inviting me to write this chapter, and Gerry Allen and the editors for helpful suggestions.

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798 / douglas fenner Ming, D.L., Wilkinson, C., Gomez, E., and S. Sudara. 1994. Status of coral reefs in the ASEAN region. Pp. 8–12 in Wilkinson, C.R. (ed.) Living Coastal Resources of Southeast Asia: Status and Management. Australian Institute of Marine Science, Townsville. Moffat, A.S. 1996. Biodiversity is a boon to ecosystems, not species. Science 271: 1497. Moor, R.J. 1989. A hit-and-run reef vandal: progress understanding the crown-of-thorns problem. Trends Ecol. Evol. 4: 36–37. Mora, C. 2001. Dispersal of reef fishes by rafting. Reef Encounter 29: 16–17. Moyer, J.T., W.K. Emerson, and M. Ross. 1982. Massive destruction of scleractinian corals by the muricid gastropod, Drupella in Japan and the Philippines. Nautilus 96: 69–82. Pandofi, J.M. 1996. Limited membership in Pleistocene reef assemblages from the Huon Peninsula, Papua New Guinea: constancy during global changes. Palaeo 22: 152–176. Pandolfi, J.M., R.H. Bradbury, E. Sala, T.P. Hughes, K.A. Bjorndal, R.G. Cooke, D. McArdle, L. McClenachan, M.J.H. Newman, G. Paredes, R.R. Warner, and J.B.C. Jackson. 2003. Global trajectories of the long-term decline of coral reef ecosystems. Science 301: 955–958. Precht, W.F., M.L. Robbart, M.L., and R.B. Aronson. 2004. The potential listing of Acropora species under the U.S. Endangered Species Act. Mar. Poll. Bull. 49 (7–8): 534–536. Randall, J.E. 1998. Zoogeography of shore fishes of the Indo-Pacific region. Zool. Stud. 37: 227–268. Reaka-Kudla, M.L. 1995a. An estimate of known and unknown biodiversity and potential for extinction on coral reefs. Reef Encounter 17: 8–12. Reaka-Kudla, M.L. 1995b. The global biodiversity of coral reefs: a comparison with rainforests. Pp. 83–108 in Reaka-Kudla, M.L., D.E. Wilson, and E.O. Wilson (eds.) Biodiversity II, Understanding and Protecting Our Biological Resources. Joseph Henry Press, Washington, D.C. Rogers, C.S. 1993. Hurricanes and coral reefs: the intermediate disturbance hypothesis revisited. Coral Reefs 12: 127–137. Ruppert, E.E., and R.D. Barnes. 1994. Invertebrate Zoology. 6th ed. Saunders, Philadelphia. Sadovy, Y., M. Kulbicki, P. Labrosse, Y. Letourneur, P. Lokani, and T.J. Donaldson. 2003. The Humphread Wrasse, Cheilinus undulatus: synopsis of a threatening and poorly known giant coral reef fish. Reviews in Fish Biology and Fisheries 13: 327–364. Scott, G.A.J., and F.M. Rotondon. 1983. A model for the development of types of atolls and volcanic islands of the Pacific lithospheric plate. Atoll Res. Bull. 260: 1–33. Shinn, E.A. 2004. The mixed value of environmental regulations: do acroporid corals deserve endangered species status? Mar. Poll. Bull. 49 (7–8): 531–533. Spalding, M.D., C. Ravilious, and E.P. Green. 2001. World Atlas of Coral Reefs. University of California Press, Berkeley. Stehli, G.G., and J.W. Wells. 1971. Diversity and age patterns in hermatypic corals. Syst. Zool. 2: 115–126. Thomas, J.D. 2000. A new model for identifying evolutionary diversity in coral reefs using marine invertebrates: a synthesis of geology and biodiversity. Proc. 9th Int. Coral Reef Symp. Abstr.: 223. Tomascik, T., A.J. Mah, A. Nontji, and M.K. Moosa. 1997. The Ecology of the Indonesian Seas, Parts I and II. The Ecology of Indonesia Series, Vol. VII, VIII. Periplus Editions, Hong Kong. Turak, E. 2003. Coral diversity and the status of coral reefs in the Raja Ampat Islands. in Donnally, 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, Bali.

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The Ecology of Papuan Coral Reefs / 799 UNDP/FAO. 1982. Survey of Coastal Areas in Irian Jia. FP/INS/78/061, Bogor. UNEP/IUCN. 1988. Coral Reefs of the World. Volume 2: Indian Ocean, Red Sea and Gulf. UNEP Regional Seas Directories and Bibliographies. IUCN, Gland, Switzerland, and Cambridge, U.K./UNEP, Nairobi, Kenya. van Soest, R.W.M. 1997. Indonesian sponges: ecology and biogeography. Pp. 1060–1063 in Tomascik, T., A.J. Mah, A. Nontji, and M.K. Moosa (eds.) The Ecology of the Indonesian Seas, Part II. The Ecology of Indonesia Series. Periplus Editions, Hong Kong. Veron, J.E.N. 1995. Corals in Space and Time; The Biogeography and Evolution of the Scleractinia. University of New South Wales Press, Sydney. Veron, J.E.N. 2000. Corals of the World. Volumes 1–3. Australian Institute of Marine Science, Townsville. Wells, F.E. 2002. Centres of species richness and endemism of shallow water marine molluscs in the tropical Indo-West Pacific. Proc. 9th Int. Coral Reef Symp. 2: 941–945. Werner, T.B., and G.R. Allen (eds.). 2000. A Rapid Marine Biodiversity Assessment of the Calamianes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Conservation International, Washington, D.C. Whitehouse, F.W. 1973. Coral reefs of the New Guinea region. Pp. 169–186 in Jones, O.A., and R. Endean (eds.) Biology and Geology of Coral Reefs 2. Academic Press, London. Wilkinson, C. (ed.). 2004. Status of Coral Reefs of the World: 2004, Vol 1–2. Australian Institute of Marine Science, Townsville. Williamson, M., K.J. Gaston, and W.M. Lonsdale. 2001. The species-area relationship does not have an asymptote! J. Biogeogr. 28: 827–830. Wing, S.R., and E.S. Wing. 2001. Prehistoric fisheries in the Caribbean. Coral Reefs 20: 1–8. Yamano, H., K. Hori, M. Yamagawa, and A. Ohmura. 2001. Highest-latitude coral reef at Iki Island, Japan. Coral Reefs 20: 9–12. Yamuna, R., and T. McClannahan. 2001. Coral reefs in Papua New Guinea. Reef Encounter 29: 23–30.

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5.3. Seagrass Ecosystems of Papua len mckenzie, rob coles, and paul erftemeijer e a gr a s s me a d o ws form a significant coastal habitat in the Papua coastal region, extending from intertidal to subtidal, along mangrove coastlines, estuaries, and shallow embayments, as well as coral-reef platforms, inter-reef seabeds and island locations. Seagrasses are a functional grouping of vascular flowering plants that have adapted to the nearshore soft bottom environments of most of the world’s continents. Most are entirely marine although some species cannot reproduce unless emergent at low tide. Seagrass are among the few plants that have migrated back to the seas roughly 100 million years ago during the Cretaceous (den Hartog 1970). Seagrasses probably evolved from a freshwater hydrophyte (a plant adapted to growing in water or inundated soil) or salt marsh-type primitive stock (den Hartog 1970). A well developed seagrass flora may have existed by the end of the Cretaceous period (Larkum and den Hartog 1989). The earliest fossil records from Malesia (Indonesia, Borneo, and New Guinea) are from well preserved fossils of Cymodocea serrulata described from Miocene deposits northeast of Makassar, South Sulawesi (Larkum and den Hartog 1989). Seagrasses can survive in a range of conditions including freshwater, estuarine, marine, or hypersaline. There are relatively few species globally (about 60) and these are grouped into just 13 genera and five families. The greatest diversity of seagrasses occurs in the Indo-Pacific region. Global seagrass distribution has been described for most species (den Hartog 1970; Phillips and Menez 1988; Spalding et al. 2003). There is now a broad understanding of the range of species and seagrass habitats although shallow subtidal and intertidal species distributions are better recorded than seagrasses in water greater than ten meters below mean sea level. Surveying deeper water seagrass is time consuming and expensive and it is likely that areas of deep water seagrass are still to be located (Lee Long, Coles and McKenzie 1996). Short, Coles, and Pergent-Martini (2001) in reviewing the world distribution of seagrasses identified the islands of the southwest Pacific and Indian Ocean, including Papua, as areas where knowledge of seagrass habitats are less well known. Papua is, however, included in the Indo-Pacific Region IX (Short, Coles, and Pergent-Martini 2001), which has the largest number of seagrass species worldwide and a high species diversity of associated flora and fauna. Short, Coles, and Pergent-Martini (2001) reported 13 species from Papua New Guinea, 16 species from the Philippines, and 16 species from neighboring northern Australia. Humoto and Moosa (2005) reported that eight genera and 13 species of seagrass inhabit Indonesian coastal waters. These include Cymodocea serrulata, Cymodocea rotundata, Enhalus acoroides, Syringodium isoetifolium, Halodule pinifolia, Halodule uninervis, Halophila spinulosa, Halophila decipiens, Halophila ovalis, Thalassia

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hemprichii, Halophila minor, Thalassodendron ciliatum, and Ruppia maritima. Halophila minor was originally reported as H. ovata, but taxonomists now regard H. ovata in the Indo-Western Pacific as only present in the South China Sea and Micronesia (Kuo 2000). The R. maritima record was based on a specimen at Herbarium Bogoriense collected from Jakarta Bay and has never been reported again. A 14th species, Halophila beccarii, although similarly known from a specimen at the Herbarium Bogoriense, was thought to exist in Indonesian waters, but has not been found in the field (Kuriandewa et al. 2003). Among the 13 species, besides R. maritima, T. ciliatum has a distribution limited to only in the eastern part of Indonesia, and H. spinulosa and H. decipiens have been recorded in only a few locations.

Importance of Seagrass Seagrasses rank as one of the major marine ecosystems in the world. In the last few decades, seagrass meadows have received greater attention with the recognition of their importance in stabilizing coastal sediments, providing food and shelter for diverse organisms, as a nursery ground for fish and invertebrates of commercial and artisanal fisheries importance, as carbon dioxide sinks and oxygen producers, and for nutrient trapping and recycling. Seagrass meadows are rated the third most valuable ecosystem globally (on a per hectare basis, behind estuaries and swamps/floodplains) and the average global value for their nutrient cycling services and the raw product they provide has been estimated at US$19,004 per hayr (in 1994 dollars) (Costanza et al. 1997). Seagrasses are also food for the endangered Green Sea Turtle (Chelonia mydas) and Dugong (Dugong dugon) (Lanyon, Limpus and Marsh 1989), which are found throughout the seas surrounding Papua, and used by traditional communities for food and ceremonial use. Tropical seagrasses are also important in their interactions with mangroves and coral reefs through fluxes of particulate and dissolved substances, physical interactions, and animal migrations. Along coastlines dominated by mangrove forests, seagrass communities often provide a functional link and a buffer between the seaward reefs and the inshore mangroves. Each of these systems exerts a stabilizing effect on the environment, resulting in important physical and biological support for the other communities. Seagrasses slow water movement, causing suspended sediment to fall out, and thereby benefit corals by reducing sediment loads in the water.

Factors That Influence Seagrass Distribution The distribution of seagrass in Indonesia is not completely known, with vast areas, including the Papua coast, unsurveyed. At least 30,000 km2 of seagrass meadows are known to occur throughout the Indonesian archipelago (Kuriandewa et al. 2003). Short, Coles, and Pergent-Martini (2001) identified nine factors that influence the distribution of seagrasses. These include: light, water depth, tide and

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water movement, salinity, temperature, human impacts, climate change, availability of propagules, and competition from other plants. In the tropics, key seagrass habitats occur on shallow fringing reef platforms and sheltered shallow bays where distribution is also driven by the physical microtopography of the location. Seagrass habitats in Papua are determined by factors that vary among regions and among seasons. It is likely that the distribution of seagrass on the muddy mangrove-lined southern coast of Papua is determined by different factors than that of seagrasses on the reef platforms surrounding coastal islands. Little is known about much of this region but it would be safe to assume that of the nine factors, human impacts, propagule availability, and climate change would have only limited influence, and that determinates of distribution would be suitable bottom type, light availability (depth and turbidity), temperature and exposure to drying, and tide and water movement (including protection from waves). Given the low population density of Papua, human impacts on seagrasses here are probably less severe than on other Indonesian islands, but there have been reports of destruction of seagrass meadows by trawl fishing in Cenderawasih Bay, and mass migration of dugongs into the Torres Strait prompted by a die-off of seagrass meadows in Papua (Marsh, Harris, and Lawler 1997; Putrawidjaja 2000). Growth and abundance of seagrasses is likely to be higher inshore due to higher nutrient levels rather than in nutrient poorer offshore waters (Kuriandewa et al. 2003). In the high rainfall tropics with a distinct monsoonal wet season, the seagrass distribution will be influenced by seasonal pulses of sediment laden, nutrient-rich freshwater discharges and run-off (Carruthers et al. 2002). Seasonal freshwater inputs will also determine which seagrass species can survive. On reef platforms and in lagoons the presence of pooling water at low tide prevents drying out and enables seagrass to survive tropical summer temperatures which would otherwise cause seagrasses to desiccate (Stapel, Manuntun, and Hemminga 1997). The sediments in these locations are often unstable and their depth can be very shallow, restricting seagrass growth and distribution. A complex set of interactions may impact a single region, including the type of habitat, the time of year, and the species growing. While little is known about long-term natural cycles in the abundance and distribution of seagrasses in Papua, seagrasses nearby in the Torres Strait and northern Queensland, where similar species occur, show abundance trends positively related to nutrient availability, and negatively correlated to sediment input and to years of high storm and freshwater disturbance (Carruthers et al. 2002). Most Papuan seagrasses are found in water less than ten meters deep and meadows can be monospecific or may consist of multispecies communities, with up to ten species present at a single location. It is generally agreed that of the 13–14 seagrass species in waters around Indonesia itself (Humoto and Moosa 2005), at least 11 are recorded from Papua (S. isoetifolium, C. serrulata, C. rotundata, H. pinifolia, H. uninervis, H. minor, H. spinulosa, H. ovalis, T. hemprichii, E. acoroides, and T. ciliatum); see Table 5.3.1. Unlike neighboring Australia, where structurally small species (e.g., members of

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Table 5.3.1. Seagrasses recorded in Papuan waters Taxon Notes Cymodoceaceae Taylor Cymodocea rotundata Ehrenb. & Hemp. Ex Aschers. Widespread and common on clear water intertidal reef flats; intermediate species that can survive a moderate level of disturbance; often grows in mixed species communities, but can form significant monospecific meadows; important nursery grounds for juvenile fish, prawns, and other invertebrates; relatively fast growing; known dugong and Green Turtle food in eastern Indonesia. Cymodocea serrulata (R. Br.) Aschers. & Magnus Widespread and common on muddy reef flats; frequently found in shallow subtidal areas, just seaward of mangroves; generally occurs with a mix of other common reef seagrasses; intermediate genera that can survive a moderate level of disturbance. Halodule uninervis (wide- & narrow-leaf) (Forsk.) Aschers. Occupies a wide range of habitats from muddy intertidal to reef tops; often forms monospecific meadows in disturbed inner reef flats and steep sediment slopes; can tolerate large fluctuations in salinity; ephemeral pioneering species with rapid turnover and high seed set, well adapted to high disturbance and high rates of grazing; important species for actively stabilizing sediments in areas of high disturbance; preferred dugong food. Halodule pinifolia (Miki) den Hartog Occurs in intertidal sandy or muddy substrates; found in sheltered bays, on coral platforms, and in high energy locations; often forms monospecific meadows; fast growing pioneering species with rapid turnover and high seed set, well adapted to high disturbance and high rates of grazing; preferred dugong food. Syringodium isoetifolium (Aschers.) Dandy Common in shallow subtidal sand/mud/silt substrates; highly variable in meadow formation, sometimes forming thick, tall monospecific stands, or sometimes a dispersed and diminutive component of mixed species meadows; intermediate genera that can survive moderate to high levels of disturbance; responds with rapid growth to excess nutrient availability; sometimes grazed by dugong when preferred species are unavailable. Thalassodendron ciliatum (Forsk.) den Hartog Typically found in rocky areas with strong currents, can tolerate considerable wave action; often dominates in the upper sublittoral in association with corals; depth range from reef crest to ca 4 m; intolerent of any freshwater inputs; common in atoll lagoons where it forms large monospecific meadows; heavily grazed by sea urchins. (continued)

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Table 5.3.1. (Continued) Taxon Notes Hydrocharitaceae Jussieu Enhalus acoroides (L.) Royle Structurally the largest tropical seagrass in stature; common in coastal lagoons, large sheltered bays, estuarine habitats, and often fringing mangrove forests; common on silty/muddy to medium and coarse sediments; forms monospecific meadows and dominates in mixed communities where it often grows with T. hemprichii; is a slow turnover, persistent species with low resistance to perturbation, suggesting that there are some coastal habitats that are quite stable over time; important shelter for juvenile fishes. Halophila decipiens Ostenfeld Only pan-tropical seagrass species; most common in subtidal waters ⬎10 m deep on fine to medium sands; often grows in large monospecific meadows or sometimes with H. spinulosa; tolerant to low light conditions; known dugong food. Halophila minor (Zollinger) den Hartog Found in sheltered areas and shallow lagoonal environments with sandy and muddy substrates; plants tolerate heavy sedimentation; usually forms monospecific meadows. Halophila ovalis (R. Br.) Hook f. Found in all habitats, but dominant species in the intertidal; has the widest environmental ranges of all species; occurs in low salinity to hypersaline; wide depth range from intertidal to ⬎30 m; exhibits significant leaf morphological plasticity; ephemeral pioneering species with rapid turnover and high seed set, well adapted to high disturbance and high rates of grazing; often the first species to recruit following disturbance; preferred food for dugong. Halophila spinulosa (R. Br.) Aschers. in Neumayer Subtidal, generally found in waters ⬎10 m deeper on fine to medium sands; often form extensive meadows mixed with H. ovalis/H. decipiens; important structural element of deepwater seabeds, providing significant habitat; able to tolerate variable and low light environments; known dugong food; often confused with two species of green algae, Caulerpa sertularioides and C. taxifolia. Thalassia hemprichii (Ehrennb.) Aschers. in Petermann The most abundant and widespread species in Papua; common on intertidal reef flats and shallow lagoons: often dominates in mixed communities; plays major role in stabilizing sediments on reef flats; grows on a variety of substrates such as silty sand, medium coarse sand, or coarse coral rubble; intolerent to large freshwater inputs; important food for turtles.

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the genera Halodule and Halophila) comprise the majority of the coastal nearshore seagrass meadows, Papuan seagrass are dominated by structurally large seagrasses (e.g., the genera Thalassia, Enhalus, and Cymodocea). Seagrasses have the ability to act as a biosink for nutrients, sometimes containing high levels of tissue nitrogen and phosphorous. Macro-grazers—Dugongs (Dugong dugon) and Green Sea Turtles (Chelonia mydas)—may also be an important feature in structuring seagrass communities in Papua. Seagrass habitats along the coastline of Papua and associated reefs can be generally categorized into four main habitats (Table 5.3.2), similar to those in tropical northern Australia (see Carruthers et al. 2002). These four broad groups of seagrass habitats are river estuary, coastal, reef, and deep water. In their natural state, these habitats are characterized by low nutrient concentrations, are primarily nitrogen limited, and are influenced by seasonal and episodic coastal runoff. All

Table 5.3.2. Seagrass habitats of Papua Habitat

Limiting factor

Seagrass species

Feature/threats

River estuaries (incl. large shallow lagoons)

Terrigenous runoff Water clarity

Cymodocea rotundata Cymodocea serrulata Halodule uninervis Enhalus acoroides Halophila minor Halophila ovalis

Highly productive High density, low diversity Often associated with mangroves Highly threatened

Coastal

Physical disturbance

Cymodocea rotundata Cymodocea serrulata Halodule uninervis Halodule pinifolia Syringodium isoetifolium Enhalus acoroides Halophila ovalis Thalassia hemprichii

Very diverse Highly productive Important for fisheries Supports dugongs Dynamic Threatened by development

Reef (incl. fringing, barrier or isolated)

Low nutrients

Cymodocea rotundata Halodule uninervis Syringodium isoetifolium Thalassodendron ciliatum Halophila ovalis Thalassia hemprichii

Support high biodiversity Shallow unstable sediment Variable physical environment Little studied Least threatened

Deep-water

Low light

Halophila decipiens Halophila spinulosa

⬎10m deep Monospecific High turnover Least known habitat Treats unknown

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seagrass habitats in Papua are influenced by high disturbance and are both spatially and temporally variable. However, the spatial and temporal dynamics of the different types of seagrass habitat are poorly understood. Each of these four habitat types has a number of dominant processes that influence seagrass growth, survival, and community biodiversity. River estuary habitats can be subtidal or intertidal, contain many seagrass species, and are often highly productive. In Papua, these habitats are closely associated with mangrove forests, characterized by fine sediments, and prone to high sedimentation and anoxic conditions. The dominant influence of river estuary habitats is terrigenous (from the land) runoff from wet-season rains. Increased river flow results in higher sediment loads that combine with reduced atmospheric light to create potential light limitation for seagrass (McKenzie 1994). Associated salinity fluctuations and scouring make river and inlet habitats a seasonally extreme environment for seagrass growth. Catchments to river estuary habitats often support a large range of land uses, including agricultural, mining, and forestry (logging). These land use practices result in increased sediment inputs (Spalding et al. 2003). In river estuary systems, differences in the life history strategies of seagrasses results in varying species assemblages. E. acoroides is a slow turnover, persistent species with low resistance to perturbation (Bridges, Phillips, and Young 1981; Walker, Dennison, and Edgar 1999), suggesting that there are some coastal habitats that are quite stable over time. However, E. acoroides is susceptible to disturbance and it is predicted that removal of a 1 m2 area from a meadow would take more than 10 years for full recovery (Rollon et al. 1998). In contrast, C. serrulata, H. uninervis, and H. ovalis are more ephemeral (Birch and Birch 1984). H. uninervis and H. ovalis are considered pioneer species growing rapidly and surviving well in unstable or depositional environments (Bridges, Phillips, and Young 1981; Birch and Birch 1984). C. serrulata grows in deeper sediments, and has been linked to increased sediment accretion (Birch and Birch 1984). Coastal habitats are both subtidal and intertidal and support the most diverse seagrass assemblage of all habitat types. Physical disturbance from waves and swell, associated sediment movement, and macro-grazers primarily control seagrass growing in coastal habitats. Episodic events such as cyclones or storms can have severe impacts at local scales, making this a dynamic and variable habitat. Sediment movement due to prevalent wave exposure creates an unstable environment where it is difficult for seagrass seedlings to establish or persist. Areas of seagrass that have been physically removed by a cyclone can take many years to regrow (Preen, Lee Long, and Coles 1995). Succession or recolonization after extreme loss has been suggested to be directional and modified by small-scale perturbations, resulting in patchiness in seagrass distributions (Birch and Birch 1984). Cymodocea and Syringodium are seen as intermediate genera that can survive a moderate level of disturbance, while Halophila and Halodule are described as ephemeral species with rapid turnover and high seed set, well adapted to high disturbance and high rates of grazing (Walker, Dennison, and Edgar 1999). The end result of this successional process, however, varies with geographic location.

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Reef habitats support seagrass communities of high biodiversity and can be highly productive. Fringing reef platforms are almost always intertidal. Shallow unstable sediment, fluctuating temperature, and variable salinity in intertidal regions characterize these habitats. Nutrient concentrations are generally low in reef habitats, however intermittent sources of nutrients are added by seasonal runoff and seabirds. The primary limiting nutrient for seagrass growth (either phosphate or nitrogen) in carbonate sediments can vary between geographic locations around the world (Short, Dennison, and Capone 1990; Fourqurean, Zieman and Powell 1992; Erftemeijer and Middelburg 1993; Udy et al. 1999). Tight nutrient recycling strategies of T. hemprichii (e.g., the location of nitrogen in the rhizomes), aids in survival in the nutrient-poor reef habitat when leaves are shed due to desiccation stress (Stapel, Manuntun, and Hemminga 1997). Reef seagrass communities also have unique faunal interactions. Bioturbation by shrimps can be so prevalent in some reef environments as to prevent seagrass growth (Ogden and Ogden 1982; Tomascik et al. 1997). A region of bare sand often separates coral heads from seagrass meadows; previous research suggests this is maintained by parrotfish and surgeonfish associated with the coral (Randall 1965). Deep water seagrasses occur at subtidal depths greater than 10 m, and are restricted to where high water clarity allows sufficient light penetration for photosynthesis (Lee Long, Mellors, and Coles 1993). Deep water seagrass areas can be extensive and dominated by Halophila species (Lee Long, Mellors, and Coles 1993; Lee Long, Coles, and McKenzie 1996). Large monospecific meadows of seagrass occur in this habitat (e.g., Halophila decipiens), which contrasts with coastal and reef habitats where the seagrass meadows are generally diverse and mixed (Coles et al. 1987). Halophila species display morphological, physiological, and life history adaptations to survive low light conditions. Halophila species have rapid growth rates and are considered opportunistic species (Birch and Birch 1984). H. decipiens has an open canopy structure with relatively little below ground biomass and high leaf turnover and rhizome elongation rates (Josselyn et al. 1986; Kenworthy et al. 1989). Halophila species also have high seed production. For example, Kuo and Kirkman (1995) reported H. decipiens seed banks of 176,880 seeds per m2. The distribution of deep water seagrasses, while mainly influenced by water clarity, is also modified by seed dispersal, nutrient supply, and current stress. Although the ecological role of deep water seagrasses is poorly understood, some deep water meadows are important dugong feeding habitat (Lee Long, Coles, and McKenzie 1996; Marsh and Saalfeld 1989; Anderson 1994). Unfortunately, deep water systems are the least understood seagrass community. The four broad groups of seagrass habitats in Papua contain a large range of life history strategies, which provides some insight into the dynamic but variable physical nature of Papuan seagrass habitats. The species present in the different habitats reflect the observed physical and biological impacts, suggesting that reef, deep water, and coastal environments are particularly variable and dynamic, while estuarine habitats have stable areas but are extremely harsh. Of these seagrass habitat types in Papua, both estuarine (including large shallow lagoons) and

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coastal seagrass habitats are of primary concern with respect to water quality due to their location immediately adjacent to catchment inputs.

Papuan Seagrasses Papua includes the most eastern province of Indonesia (formerly known as Irian Jaya) and extends west from the northeast border of Papua New Guinea to Halmahera (north Maluku Province). It encompasses the overall north and south coasts and northern offshore oceanic islands. Overall this is a region separated from the main Indonesian archipelago by relatively complex bathymetry, where waters are very deep, and even islands only a few tens of kilometers apart might be separated by depths of over 1,000 meters (Spalding, Raviolus, and Green 2001). The only areas of relatively extensive shallow water and true continental shelf are a platform west of the Vogelkop Peninsula and to the south where Papua shares a common continental shelf with northern Australia. Surface currents are somewhat mixed in this region, however a northward current flows between Papua and Halmahera and an eastward current flows along the north shore of Papua during the northeast monsoon. This pattern reverses during the southeast monsoon. The Raja Ampat Archipelago includes the four large islands of Waigeo, Batanta, Salawati, and Misool and hundreds of smaller islands. Ecological Rapid Assessments through the Raja Ampat Islands in 2001 and 2002 visited a total of 45 and 59 sites, respectively, surveying coral, mangrove, seagrass, other marine habitats, and turtle nesting beaches (McKenna, Allen, and Suryadi 2002; Donnelly, Neville, and Mous 2003). Substantial shallow seagrass meadows of T. hemprichii, C. rotundata, and H. uninervis were reported on Sayang Island, in the bays on the southeastern side of Kawe, on reefs off the northern Waigeo coast and at Deer Island off the northern coast of Kofiau (Donnelly, Neville, and Mous 2003; Hitipeuw 2003). In the south of the archipelago, extensive seagrass meadows of E. acoroides and S. isoetifolium have been reported on the reef flat of Batanta Island (Tomascik et al. 1997; Scheltze-Westrum 2001). Seagrasses were also recorded at Kri Island, Pef Island, Waigeo Island (Mayalibit Passage), Wruwarez Island, the northwest side of Batanta Island, North Fam Island, Batang Pele Island, Wofah Island and Yeben Kecil Island (McKenna, Allen, and Suryadi 2002). The seagrasses reported from northern coast of the Vogelkop, east of Sorong District, include S. isoetifolium, C. serrulata, C. rotundata, H. pinifolia, H. spinulosa, H. ovalis, T. hemprichii, and E. acoroides (Tomascik et al. 1997, Kuriandewa et al. 2003; see Table 5.3.3). There is however, little or no information describing the reef communities further east around Vogelkop Peninsula. An ecological Rapid Assessment of Biak and the Supiori Islands in 1996 found nine seagrass species (T. hemprichii, C. rotundata, C. serrulata, H. uninervis, H. pinifolia, E. acoroides, H. ovalis, H. minor, and S. isetofolium; MREP 1996), of which C. rotundata and T. hemprichii were the most widely distributed, creating high density monospecific meadows (1,276 shoots per m2; Kuriandewa et al. 2003).

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•

Halodule pinifolia

•

•

Halodule uninervis

•

•

Halophila spinulosa

•

•

•

•

•

• •

Aru Islands

Syringodium isoetifolium

•

•

Merauke

•

•

•

Mimika

•

•

Fakfak

Enhalus acoroides

•

Jayapura

•

Biak-Numfoor

•

Yapen-Waropen

•

Cymodocea rotundata

Nabire

Cymodocea serrulata

Manokwari

Species

Sorong (incl. Raja Ampat)

Table 5.3.3. Distribution of seagrass species in Papua, including Aru Islands, Maluku Province

•

• • •

Halophila decipiens Halophila ovalis

•

•

•

•

Thalassia hemprichii

•

•

•

•

• •

•

•

Halophila minor Thalassodendron ciliatum

•

•

•

•

A prominent feature of the reefs in the Padaido Islands (south of Biak Island) is the presence of extensive reef-top seagrass meadows (530 ha) dominated by Cymodocea spp., E. acoroides, and T. hemprichii (Tomascik et al. 1997). The western extremity of the Padaido Islands has a dense coverage (95–100%) of seagrass over 529 ha of shallow reef flat, consisting of seven species (T. hemprichii, C. rotundata, C. serrulata, H. uninervis, H. pinifolia, E. acoroides, and H. ovalis). Similarly, extensive reef top seagrass meadows (T. hemprichii, C. rotundata, H. ovalis, and H. pinifolia) have been reported on Numfoor Island (Tomascik et al. 1997) and along the southern coast of Yapen Island. Cenderawasih Bay National Park (established in 1994) is the largest marine park in Southeast Asia and the only marine park in the region. Extensive lagoonal seagrass meadows (T. hemprichii, C. rotundata, H. uninervis, E. acoroides, and H. ovalis) are present along the mainland coast of southwestern Cenderawasih Bay, particularly in Wandammen Bay (Nietschmann et al. 2000). The vast seagrass meadows in this bay are reported to harbor a large dugong population (Petocz, 1989). Maruanaya (2000) reported three species of seagrass in the same region

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covering an area of 24 ha on the seaward side of mangrove areas with average density of 56 shoots per m2. Fringing reefs surrounding the many small islands in the region are also covered with seagrass meadows, including Pepaya Island (near Nabire) (Chou et al. 2002). Virtually nothing is known of marine ecosystems along the north coast of Papua. The north coast is almost reef free and continues as such from Cenderawasih Bay to Sarmi, with only very occasional areas of fringing reef about some of the small islands (Spalding, Raviolus, and Green 2001). Further east, fringing reefs are believed to follow a large proportion of the coastline to the border with Papua New Guinea. For the most part these are poorly described, but reef flats are estimated to reach 300–400 meters wide in places. Tomascik et al. (1997) reported six seagrass species (C. rotundata, E. acoroides, H. ovalis, C. serrulata, T. ciliatum, and T. hemprichii) on the near continuous fringing reefs from Jayapura to the border with PNG. Relatively little is known about the seagrasses along the shores of southern Papua. This area of the coastline has extensive mangrove forests. Over half the area of mangroves in Indonesia are located in Papua (Spalding, Blasco, and Field 1997). Bintuni Bay, which contains more than 1.1 million acres of mangroves, is the world’s third largest mangrove area and the second largest in Asia. However, the only seagrass known in the bay is an anecdotal report from Berau Bay (the west side of Bintuni Bay) (Jamartin H. S. Sihite, pers. comm.). No seagrasses were located anywhere else within Bintuni Bay or within the open, deeper waters (up to 60 m) towards McCluer Gulf (Erftemeijer, Allen, and Zuwendra 1996). This is possibly a consequence of the high turbidity throughout Bintuni Bay: secchi depths (a parameter used to measure the clarity of surface waters) of 11 to 85 cm in the mangrove area (creeks and rivers) and maximum 157 cm in the open waters of the bay (Erftemeijer, Allen, and Zuwendra 1996). Unfortunately Bintuni Bay marine ecosystems are increasingly threatened by overharvesting, logging, and clearing to make way for coastal shrimp farm facilities. Although the Bintuni Bay Nature Reserve affords some protection, there are no seagrasses in the Reserve and economic development is increasing due to a new liquified natural gas field in the bay, and the human population is expanding rapidly. Although there are other significant areas of mangroves and wetland areas with sago palms in the gulf near Timika, Mimika district, around the Asmat region, and surrounding Yos Sudarso Island, Merauke (Spalding, Blasco, and Field 1997), the presence of seagrass communities is unknown. Large amounts of sediment are found along the southeastern coast (apparent in remote images; see http:// eosweb.larc.nasa.gov/; http://eol.jsc.nasa.gov/), which possibly prohibit reef development in this region. Significant land clearing, logging, and mine tailings may exacerbate sedimentation, further prohibiting seagrass growth in localized areas. For example, large tracts of mangrove were also cleared at the mouth of the Timika River to construct the Amamapare seaport and mine tailings are now polluting nearby coral reefs. As part of the Freeport-McMoRan Copper & Gold Inc. mine Long-Term Environmental Monitoring Program, monitoring of benthos occurs at

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14 sites in the estuaries (e.g., Minajerwi River) and 40 sites in the Arafura Sea (http://www.fcx.com/envir/wtsd/2004/env-perform.htm). The monitoring indicates no impact of tailings on the marine benthos in the Arafura Sea outside of the tailings management area. It is likely however, that seagrasses are present throughout this region as significant seagrass meadows surround the nearby Aru Islands, a group of about 95 low-lying islands (8,563 km2) in the Arafura Sea (Moluccas), southwest of Papua. These meadows are predominately E. acoroides, T. hemprichii, C. rotundata, and S. isoetifolium. H. decipiens has also been reported from the deeper waters in the north of the islands (Nietschmann et al. 2000) and possibly extends northward to the Papuan mainland coast. Expansive seagrass meadows, which support significant Green Sea Turtle populations, also surround the adjacent Kai Islands, Kai Kecil and Kai Besar (Sua´rez 2001). South of this region are the expansive seagrass meadows of the Torres Strait. The Torres Strait is a shallow (mostly 10–20 m depth) body of water formed by a drowned land ridge extending from Cape York to southwestern Papua New Guinea. Seagrass communities occur across the open sea floor, on reef flats and subtidally adjacent to continental islands. The large expanses of open water bottom are covered with either sparsely distributed Halophila or mixed species (Halodule, Thalassia, and Syringodium) communities (Coles, McKenzie, and Campbell 2003). It is likely that these meadows may extend northward to the Merauke coast, but surveys (e.g., Long et al. 1995) have not included this region. Johnstone (1982) considered that seagrass zonation, where it occurs, was fairly similar across New Guinea and seems to be determined by comparable biotic and abiotic parameters. It can be safely assumed that such zonation would also be relevant in Papua, where species and habitats are similar. From intertidal to subtidal, the zonation pattern of seagrasses generally begins with a zone of one or two species (mostly H. uninervis, H. pinifolia, or H. minor). Subsequently, in the lower eulittoral zone, other seagrass species join in a mixed seagrass meadow generally dominated by C. rotundata, H. uninervis, and T. hemprichii, with isolated patches of H. ovalis. In the upper sublittoral zone, the mixed seagrass meadow is dominated by T. hemprichii and E. acoroides, with isolated patches of S. isoetifolium, C. serrulata, and H. uninervis. The lower edge of the meadow consists of a combination or two to four species when a reef plateau is present, or monospecific H. decipiens or H. spinulosa at the deepest depths on the sublittoral sandy slopes. The remaining species are less common and not widely distributed. Monospecific patches of T. ciliatum have been reported to occur on coral rubble banks in 6–8 meters depth on the deeper edges of the reef slopes (e.g., Jayapura; Johnstone 1982). Monospecific patches of T. ciliatum are also common on reef edges in the nearby Torres Strait between Papua and northern Australia (Coles, McKenzie, and Campbell 2003). Local conditions may determine which seagrass species are present. Extensive mixed seagrass meadows are the dominant community type in the bays, harbors, and sheltered capes along the coasts of the Papuan mainland and large continental

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islands. These extensive seagrass meadows are dominated by T. hemprichii and/or E. acoroides, with up to nine other species present to varying degrees. H. decipiens meadows sometimes occur in the deeper areas and meadows of E. acoroides occur in shallow lagoons and border the gentle sloping mangrove fringes in the more protected bays. This species is common in sheltered bays and on reef platforms throughout the tropics in water depth less than two meters at low tide. This is a species that must be able to reach the surface to pollinate and so is restricted to shallow and sheltered waters. Throughout the rest of Papua most seagrass occurs in shallow lagoons or on the reef platforms and leeward shores of small vegetated islands. These communities are dominated by colonizing and intermediate species, such as T. hemprichii, C. rotundata, and H. uninervis, which can survive a moderate level of disturbance. E. acoroides occurs in small protected bays or behind the reef crest on the sublittoral reef flat, as it has low resistance to perturbation (Walker, Dennison, and Edgar 1999). Smaller islands are generally characterized by relatively small fringing reef platforms, where seagrass communities dominated by C. rotundata and T. hemprichii, with small quantities of H. ovalis, are restricted to locations with shallow lagoons (0–2 m depth). Heijs and Brouns (1986) studied the Wewak coastline of northern Papua New Guinea, which consists of several bays separated by headlands (capes) with extensive mixed species seagrass communities generally located on the fringing reef platforms from Wewak to Vanimo near the Papuan border. These mixed meadows are dominated by T. hemprichii with E. acoroides, S. isoetifolium, and C. rotundata. Other species such as H. uninervis, H. ovalis, and C. serrulata occur occasionally. On the landward side, seagrass meadows are dominated by a narrow band of H. uninervis mixed with C. rotundata. The seaward side of the meadows are generally of combination of two to four species, which become monospecific H. decipiens in the deeper areas. The distribution of E. acoroides is either interspersed or forming small isolated patches behind the reef crest.

flora and fauna associated with seagrass Although few studies have examined the macro- and mega-fauna in seagrass meadows in Papua, some general remarks can be made. The most conspicuous macrofauna is often the abundance of holothurians (sea cucumbers), the most common is the black sea cucumber Holothuria atra. Echinoids (sea urchins and sand dollars) are also common in the mid-and lower eulittoral areas, and the genus Tripneustes is abundant. Asteroidea (true starfish) are abundant, particularly in seagrass meadows with sandy substrate. Reef platform seagrass meadows support a wide range of mollusks, fish, holothurians, and decapods (shrimp, lobster, and crabs). Common gastropods (snails) found associated with seagrasses include Strombidae, Cypraidae, and Conidae. Most of these occur in the eulittoral and sublittoral areas. Other mollusks such as the trochus shell Trochus niloticus found in seagrass meadows are collected as a source of cash income. Similarly the Holothurians have been a valuable source of cash income although now heavily overfished (Uthicke and Conand 2005).

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The existence of productive commercial shrimp fisheries in the coastal waters of the Aru Islands, Moluccas, is largely due to the presence of extensive seagrass meadows in the area. An average shrimp catch of about 490 tons per year has been reported from commercial trawling grounds south of the Aru Islands (Tomascik et al. 1997). Megafauna such as Green Sea Turtles and dugong depend on the seagrass meadows present throughout Papua which are recognized as significant foraging grounds. Such areas include Cenderawasih Bay, northwest of Biak Island, and Sahul Shelf (Arafura Sea) near the Aru Islands. Many of these locations are adjacent to important nesting beaches for Hawksbill and Green Sea Turtles, such as Ingressau Beach on the northeastern coast of Yapen Island. Many of the dugong and turtle populations supported by the seagrass meadows are also traditionally hunted. Last but by no means least, an abundant array of fishes uses seagrass meadows’ different tide regimes during parts of their life history. Some fish are herbivorous, feeding either on the seagrass leaves or the epiphytes, such as Siganids. Maruanaya (2000) studied seagrass associated fish in Cenderawasih Bay and recorded 55 seagrass fish species dominated by sardines (Stolephorus bucanieri), rabbitfish (Siganus canaliculatus), and Gerres kapas. Some indication of the likely use of tropical Pacific seagrass meadows are reports that 154 species of tropical invertebrates and fish feed directly on seagrasses (Klump, Howard, and Pollard 1989), and that Coles et al. (1993) listed and classified 134 taxa of fish and 20 shrimp species found in tropical Australian seagrass meadows. Other fish such as the Lutjanidae (snappers) use the seagrass as shelter when they are juveniles, and some Syngnathids (seahorses and pipefishes) permanently reside or shelter in seagrass meadows. Pyle (1999) lists at least 3,392 fish described as reef and shore fish from the Pacific Islands but it is not possible to distinguish which are from seagrass meadows. However, Allen (2003) reported from the ecological Rapid Assessment conducted of the Raja Ampat Islands, that although the region has one of the world’s richest coral reef fish faunas, other habitats such as silty bays, mangroves, seagrass meadows, and pure sand-rubble areas were consistently the poorest areas for fish diversity. Seagrass meadows throughout Papua are of significant importance to subsistence fisheries for Siganids (rabbitfish), Hemirhamphidae (garfish), holothurian species, and shellfish.

losses and threats Tropical seagrass meadows are known to fluctuate in size seasonally and across years (Erftemeijer and Herman 1993; Mellors, Marsh, and Coles 1993; McKenzie 1994; McKenzie et al. 1996), and losses have been reported from most parts of the world, sometimes from natural causes such as cyclones and floods (Poiner, Walker, and Coles 1989; Campbell and McKenzie 2004). More commonly, loss has resulted from human activities such as dredging, land reclamation, industrial runoff, oil spills, or changes in land use and agricultural runoff (Short and WyllieEcheverria 1996).

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The major changes in Papuan seagrass meadows have occurred since World War II and are related to coastal development, agricultural land use, and population growth. However there is insufficient information and no long-term studies from which to draw direct conclusions about historic trends. Munro (1999) reported that 2,000 year old mollusk shell middens in Papua New Guinea have essentially the same species composition as present day harvests, suggesting indirectly that the habitats, including seagrass habits and their faunal communities, are stable and that any changes occurring are either short-term or the result of localized impacts. Dependence on coastal marine ecosystems for protein remains high and subsistence fishing is widespread. Localized impacts are likely to occur from sedimentation, that increases turbidity of marine waters, and is related to coastal agriculture (palm oil plantations), land clearing (upland logging and mining), bush fires, and from the discharge of mine tailings (e.g., from Freeport-McMoRan Copper & Gold Inc., in Timika; Coles and McKenzie 2005). Inappropriate coastal development or construction often result in beach erosion. Major impacts result from collecting beach sand for construction materials, construction of airports, hotels, and other structures too close to beaches or in offshore waters, and sand mining. In other locations in Indonesia (e.g., Seribu Islands and the coast of Bali), heavy coral mining and collection from reef flats have resulted in the deterioration of seagrass meadows (Humoto and Moosa 2005). Other factors that negatively impact seagrass ecosystems include sewage discharge, industrial pollution, and overfishing. For example, there have been reports that dugong are disappearing from Cenderawasih Bay National Park because the shallow water seagrass meadows are being destroyed by trawl fishing as well as sedimentation resulting from deforestation (Putrawidjaja 2000). Most of these impacts remain localized and relatively small and can be managed with appropriate environmental guidelines. However, in the future climate change and associated increase in storm activity, water temperature, or sea level rise has the potential to damage seagrasses in the region or to influence their distribution. All identified seagrass habitats have high ecological or economic value, whether supporting fisheries or biodiversity. Estuary/lagoonal and coastal habitats are considered to be the most threatened, due to extensive coastal development. However, the limited knowledge of deeper water seagrass habitats suggests that impacts on these habitats are extremely difficult to assess.

conservation Currently there is no legislation in Indonesia that specifically stipulates that the function of seagrass ecosystems should be maintained (Indonesian Seagrass Committee 2002). However, seagrasses do not exist in nature as a separate ecological component and are often closely linked to other community types. Associations are likely to be complex interactions with mangrove communities, algae beds, salt marshes, and coral reef systems. Worldwide, many management activities to protect seagrasses have their origins in the protection of wider ecological systems or

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are designed to protect the overall biodiversity of the marine environment. The protection of seagrass habitat for species listed as threatened or vulnerable to extinction (e.g., Dugong and Green Sea Turtle), and their importance as habitat for juvenile fish and crustaceans which form the basis of economically valuable subsistence and commercial fisheries, have become motivating factors for the protection of seagrasses. In Indonesia existing legislation relevant, directly or indirectly, to the management of seagrass ecosystems is considered sufficient for the adequate protection of seagrass ecosystems in the near future. However, there is an urgent need to reach common understanding regarding the vision and mission required to implement these laws in the field (Indonesian Seagrass Committee 2002). Law enforcement is still weak and ineffective; hence pollution and degradation of seagrass ecosystem continue to occur (Indonesian Seagrass Committee 2002). The Indonesian Seagrass Committee in 2002 assessed many of the problems of legal aspects relevant to seagrass management in Indonesia and made a number of recommendations. These recommendations suggest that the following five steps be taken. First, an institution must be assigned specific authority to coordinate the campaign against pollution and degradation of the sea. Second, many legal acts (including Fisheries, Management of Living Environment, and acts concerning natural resources and their ecosystems) must be revised. Third, terrestrial spatial planning should be integrated with that of coastal areas and the sea on the basis of integrated ecosystems; Provincial and District/Municipality governments should designate new conservation areas in accordance with the land-use plan. Fourth, the division of authority between Provincial and District/Municipality governments in administrative aspects be publicized so that the management of ecosystems are assessed holistically and in integrated manner, and free of bureaucratic complications. Fifth, forest cutting, which directly affects coastal and marine ecosystems (including river banks, greenbelts of dams, lakes, rivers, and coast lines) be prohibited. Implementation of such recommendations may require several approaches. Coles and Fortes (2001) separated these into three components: a prescriptive legal approach; a non-prescriptive broad-based approach ranging from planning processes to education; and a reactive approach designed to respond to specific issues, such as a development proposal. These approaches may overlap and be used simultaneously in many cases. Prescriptive management of seagrass issues might range from local laws to a Presidential Decree. In Southeast Asian countries such as Indonesia and in the Pacific Island countries, protection is often strongest at the village or district level by government-supported agreements or through local level management (Coles and Fortes 2001). At the village level, successful enforcement is heavily dependent on community support. While no international legislation specifically protects seagrass, there are international conventions that recognize the importance of wetlands and coastal areas, such as the Ramsar Convention on Wetlands. In some cases, seagrass meadows

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have been inadvertently protected because they are located within protected areas, such as Cenderawasih Bay National Park and Kamiali Wildlife Management Area. It is hoped that recognition of the global significance of areas such as Raja Ampat, which also include seagrasses, will also provide some degree of future protection. Prescriptive management can include establishment of Marine Protected Areas (MPAs). A Marine Protected Area is an area of sea that is dedicated to the protection and maintenance of biological diversity and of natural and associated cultural resources, and is managed through legal and other effective means (IUCN 1994). A Marine Protected Area may be a ‘‘no-take’’ area like a terrestrial national park or it may comprise a multiple-use area, zoned in such a way to minimize conflicts and allow extractive activities to occur in specific areas. Establishing even a small Marine Protected Area is a complex process and includes a needs assessment and requires the involvement of all stakeholders and government agencies in defining a border and specifying permitted uses. An alternate and complementary non-prescriptive approach is a Locally-managed Marine Area (LMMA). A Locally-managed Marine Area is an area of nearshore waters (including include coral reefs, seagrass meadows, mudflats, mangrove, and other areas) that is actively being managed by local communities or land-owning groups, or is being collaboratively managed by local communities together with local government and other partners based in the immediate vicinity. For example, Yayasan Rumsram, a nongovernmental conservation organization in Biak and the Padaido Islands, is pioneering the use of Locally-managed Marine Areas (LMMAs) in Indonesia through traditional marine resource management and customary prohibition (sasizen) practices. Non-prescriptive methods of protecting seagrasses generally have a strong extension or educational focus. Providing information is important because it encourages and enables individuals to act voluntarily act in ways that reduce impacts on seagrasses. Actions in response to such information could range from being more aware of the downstream effect of poor agricultural practices to lobbying politicians for stronger sanctions against decisions that lead to seagrass loss. Nonprescriptive methods range from simple explanatory guides to complex industry codes of practice developed in negotiation with the industry in question (Coles and Fortes 2001). Reactive processes generally occur in response to a perceived operational threat, such as a coastal development proposal. Reactive processes can also include risk management plans that identify areas of seagrass to be protected in the event of an impact (e.g., oil spill or ship grounding). Reactive processes are generally identified in environmental impact statements, which also propose strategies (e.g., redesign, response, or by reducing future risk) to minimize the effects of a development or structure on the coastal environment, including seagrasses. The combination of project redesign in response to environmental impact statements and reactive environment management systems can provide enormous improvements to coastal seagrass protection.

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Discussion A key step in protecting seagrasses in this region will be to obtain better distributional and abundance data and to develop a better understanding of seasonal changes and local ecosystems. At the present time information is patchy at best and it is quite likely that large areas of seagrass could be lost without any formal record. Seagrass dieback has been recorded in nearby waters of the Torres Strait (Long et al. 1995) and is considered of sufficient concern to be a major focus of the Australian Cooperative Research Centre for the Torres Strait. A survey of 3,000 kilometers of the northern Australia coastline has just been completed (Roelofs, Coles, and Smit 2005) using a helicopter as a cost-effective way of estimating seagrass area, abundance, and species over a large and remote area. Similar methods could provide an up-to-date map of Papuan seagrass with a precision suitable for quantifying future gains and losses. What is recorded for Papua suggests distribution patterns of seagrasses are comparable to that found in other parts of the Indonesian archipelago, Papua New Guinea, the western Pacific islands, the Philippines, and northern Australia (Coles and Lee Long 1999; Coles et al. 2003; Green and Short 2003). Subsets of the same suite of tropical species occur and the zonation patterns described can be found in similar locations in all the adjoining countries and islands. The threats to seagrasses are also relatively generic to the region, with local land clearing and resulting sediment run off, mine tailings, inappropriate fishing methods, and nutrients from sewage likely to be the major problems at a local scale. Population and development levels in Papua are generally low at the present time, but as they increase, transport infrastructure development issues will affect coastal seagrasses as they have elsewhere. Climate change is likely to be the major variable in the medium to long term. Climate change is predicted to raise sea level and seawater temperatures, and to increase carbon dioxide concentrations in seawater. Rising sea levels could increase the distribution of seagrass because more inland areas will be covered by seawater. However, the sediment erosion that is likely to be associated with sea level rise could destabilize the marine environment and cause seagrass losses. Increasing concentrations of carbon dioxide in seawater could increase the area of seagrass because seagrasses will have more carbon available for growth and could increase photosynthetic rates. Increased seawater temperatures might raise the photosynthetic rate of seagrasses. However, in some places, seagrasses are close to their thermal limit and rising temperatures could cause ‘‘burning’’ and tissue death. To provide an early warning of change, long-term monitoring and community engagement programs have been established as part of the Global Seagrass Monitoring Network (www.SeagrassNet.org, Short et al. 2002; www.seagrasswatch.org, McKenzie, Campbell, and Roder 2001). Establishing a network of monitoring sites in Papua would provide valuable information on temporal trends in the health status of seagrass meadows in this region and provide a tool for decision makers

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in adopting protective measures. Monitoring encourages local communities to become involved in seagrass management and protection. For example, one of the recommendations for conservation action after the 2002 ecological Rapid Assessment in Raja Ampat was the establishment of monitoring programs, including seagrass monitoring (Donnelly, Neville, and Mous 2003). Working with both scientists and local communities, this approach is designed to draw attention to the many local anthropogenic impacts on seagrass meadows that degrade coastal ecosystems and decrease their yield of natural resources.

Acknowledgments We thank the participants of the University of New Hampshire and the David and Lucile Packer Foundation–funded Seagrass 3M Workshop: Mapping, Monitoring and Management of Seagrass Resources in the Indo-Pacific, held at The Nature Conservancy, Southeast Asia Center for Marine Protected Areas, Sanur, Bali, 9th to 12th May 2005, for their assistance. We also thank Yayu La Nafie and the members of the Indonesian Seagrass Association ([email protected]) for their encouragement and support.

Literature Cited 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, held October 30–November 22, 2002. The Nature Conservancy–Southeast Asia Center for Marine Protected Areas, Sanur, Bali, Indonesia. Anderson, P.K. 1994. Dugong distribution, the seagrass Halophila spinulosa, and thermal environment in winter in deeper waters of eastern Shark Bay, Western Australia. Wildlife Research 21: 381–388. Birch, W.R., and M. Birch. 1984. Succession and pattern of tropical intertidal seagrasses in Cockle Bay, Queensland, Australia: a decade of observations. Aquatic Botany 19: 343–367. Bridges, K.W., R.C. Phillips, and P.C. Young. 1981. Patterns of some seagrass distribution in the Torres Strait, Queensland. Australian Journal of Marine and Freshwater Research 33: 273–283. Campbell, S.J., and L.J. McKenzie. 2004. Flood related loss and recovery of intertidal seagrass meadows in southern Queensland, Australia. Estuarine, Coastal and Shelf Science 60: 477–490. Carruthers, T.J.B., W.C. Dennison, B.J. Longstaff, M. Waycott, E.G. Abal, L.J. McKenzie, and W.J. Lee Long. 2002. Seagrass habitats of northeast Australia: models of key processes and controls. Bulletin of Marine Science 71 (3): 1153–1169. Chou, L.M., V.S. Tuan, Philreefs, T. Yeemin, A. Cabanban, Suharsono, and I. Kessna. 2002. Status of Southeast Asia coral reefs. Pp. 123–152 in Wilkinson, C.R. (ed.) Status of Coral Reefs of the World: 2002. GCRMN Report, Australian Institute of Marine Science, Townsville. Coles, R.G., and M. Fortes. 2001. Protecting seagrass–approaches and methods. Pp.

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Seagrass Ecosystems of Papua / 819 445–463 in Short, F.T., and R.G. Coles (eds) Global Seagrass Research Methods. Elsevier Science BV, Amsterdam. Coles, R.G., and W.J. Lee Long. 1999. Seagrass. Pp. 21–46 in Eldredge, L.G., J.E. Maragos, P.F. Holthuis, and H.F. Takeuchi (eds.) Marine and Coastal Biodiversity in the Tropical Islands PacificRregion. Vol 2: Population, Development and Conservation Priorities. Proceedings of two workshops held at the East-West Center, Honolulu, November 1994. East-West Center, Honolulu. Coles, R.G., W.J. Lee Long, B.A. Squire, L.C. Squire, and J.M. Bibby. 1987. Distribution of seagrasses and associated juvenile commercial penaeid prawns in northeastern Queensland waters (Australia). Australian Journal of Marine and Freshwater Research 38: 103–120. Coles, R.G., W.J. Lee Long, R.A. Watson, and K.J. Derbyshire. 1993. Distribution of seagrasses, and their fish and penaeid prawn communities, in Cairns Harbour, a tropical estuary, northern Queensland, Australia. Australian Journal of Marine and Freshwater Research 44: 193–210. Coles, R.G., and L.J. McKenzie. 2005. Seagrass 3M workshop: mapping, monitoring and management of seagrass resources in the Indo-Pacific. Report to UNH on workshop held at The Nature Conservancy, Southeast Asia Center for Marine Protected Areas, Sanur, Bali, 9–12 May, 2005. DPI&F, Cairns. Coles, R.G., L.J. McKenzie, and S.J. Campbell. 2003. The seagrasses of eastern Australia. Pp. 119–133 in Green, E.P., and F.T. Short (eds.) World Atlas of Seagrasses. Prepared by the UNEP World Conservation Monitoring Centre. University of California Press, Berkeley. Coles, R.G., L.J. McKenzie, S.J. Campbell, M. Fortes, and F.T. Short. 2003. The seagrasses of the western Pacific Islands. Pp. 161–170 in Green, E.P., and F.T. Short (eds.) World Atlas of Seagrasses. Prepared by the UNEP World Conservation Monitoring Centre. University of California Press, Berkeley. Costanza, R., R. d’Arge, R. de Groot, S. Farber, M. Grasso, B. Hannon, K. Limburg, S. Naeem, R.V. O’Neil, J. Paruelo, R.G. Raskin, P. Sutton, and M. van der Belt. 1997. The value of the world’s ecosystem services and natural capital. Nature 387 (15): 253–260. den Hartog, C. 1970. The Seagrasses of the World. North-Holland Publishing, Amsterdam. Donnelly, R., D. Neville, and P.J. Mous (eds.). 2003. Report on a Rapid Ecological Assessment of the Raja Ampat Islands, Papua, Eastern Indonesia, held October 30–November 22, 2002. Report by The Nature Conservancy–Southeast Asia Center for Marine Protected Areas, Sanur, Bali, Indonesia. Erftemeijer, P.L.A., G. 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 No. 8, Bogor. Erftemeijer, P.L.A., and P.M.J. Herman. 1993. Seasonal changes in environmental variables, biomass, production and nutrient contents in two contrasting tropical intertidal seagrass beds in South Sulawesi, Indonesia. Oecologia 99: 45–59. Erftemeijer, P.L.A., and J.J. Middelburg. 1993. Sediment-nutrient interactions in tropical seagrass beds: a comparison between a terrigenous and a carbonate sedimentary environment in South Sulawesi (Indonesia). Marine Ecology Progress Series 102: 187–198. Fourqurean, J.W., J.C. Zieman, and G.V.N. Powell. 1992. Relationship between porewater nutrients and seagrass in a subtropical carbonate environment. Marine Biology 114: 57–65.

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820 / len mckenzie , rob c oles , & p a ul erf tem ei jer Green, E.P., and F.T. Short (eds.). 2003. World Atlas of Seagrasses. Prepared by the UNEP World Conservation Monitoring Centre. University of California Press, Berkeley. Heijs, F.M.L., and J.J.W.M. Brouns. 1986. A survey of seagrass communities around the Bismarck Sea, Papua New Guinea. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen. Series C, 89 (1): 11–44. Hitipeuw, C. 2003. Status of sea turtle populations in the Raja Ampat Islands. Pp. 85–96 in Donnelly, R., D. Neville, and P.J. Mous (eds.) Report on a Rapid Ecological Assessment of the Raja Ampat Islands, Papua, Eastern Indonesia, held October 30–November 22, 2002. Report by The Nature Conservancy–Southeast Asia Center for Marine Protected Areas, Sanur, Bali, Indonesia. Humoto, M., and M.K. Moosa. 2005. Indonesian marine and coastal biodiversity: present status. Indian Journal of Marine Sciences 34: 88–97. IUCN. 1994. Guidelines for Protected Area Management Categories. Commission on National Parks and Protected Areas with the assistance of the World Conservation Monitoring Centre, Gland, Switzerland. Indonesian Seagrass Committee. 2002. Assessment of institution and legal aspect relevant to management of seagrass ecosystem. The 1st Six Months Progress Report of Indonesian Seagrass Committee. UNEP-GEF Project: ‘‘Reversing Environmental Degradation Trends in South China Sea and the Gulf of Thailand.’’ UNEP, Jakarta. Johnstone, I.M. 1982. Ecology and distribution of seagrasses. Pp. 497–512 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Monographiae Biologicae Vol 42. Dr. W. Junk Publishers, The Hague. Josselyn, M., M. Fonseca, T. Niesen, and R. Larson. 1986. Biomass, production and decomposition of a deep water seagrass, Halophila decipiens Ostenf. Aquatic Botany 25: 47–61. Kenworthy, W.J., C.A. Currin, M.S. Fonseca, and G. Smith. 1989. Production, decomposition and heterotrophic utilization of the seagrass Halophila decipiens in a submarine canyon. Marine Ecology Progress Series 51: 277–290. Klump, D.W., R.K. Howard, and D.A. Pollard. 1989. Thropodynamics and nutritional ecology of seagrass communities. Pp. 394–457 in Larkum, A.W.D., A.J. McComb, and S.A. Shepherd (eds.) Biology of Seagrasses: A Treatise on the Biology of Seagrasses with Special Reference to the Australian Region. Elsevier, Amsterdam. Kuo, J. 2000. Taxonomic notes on Halophila minor and H. ovata. Biol. Mar. Medit. 7 (2): 79–82. Kuo, J., and H. Kirkman. 1995. Halophila decipiens Ostenfeld in estuaries of southwestern Australia. Aquatic Botany 51: 335–340. Kuriandewa, T.E., W. Kiswara, M. Hutomo, and S. Soemodihardjo. 2003. The seagrasses of Indonesia. Pp. 171–182 in Green, E.P., and F.T. Short (eds.) World Atlas of Seagrasses. Prepared by the UNEP World Conservation Monitoring Centre. University of California Press, Berkeley. Lanyon, J.M., C.J. Limpus, and H. Marsh. 1989. Dugongs and turtles: grazers in the seagrass system. Pp. 610–34 in Larkum, A.W.D., A.J. McComb, and S.A. Shepherd (eds.) Biology of Seagrasses: A Treatise on the Biology of Seagrasses with Special Reference to the Australian Region. Elsevier, Amsterdam. Larkum, A.W.D., and C. den Hartog. 1989. Evolution and biogeography of seagrasses. Pp. 112–156 in Larkum, A.W.D., A.J. McComb, and S.A. Shepherd (eds.) Biology of Seagrasses: A Treatise on the Biology of Seagrasses with Special Reference to the Australian Region. Elsevier, Amsterdam.

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Seagrass Ecosystems of Papua / 821 Lee Long, W.J., R.G. Coles, and L.J. McKenzie. 1996. Deepwater seagrasses in Northeastern Australia—How deep? How meaningful? Pp. 41–50 in Kuo, J., R.C. Phillips, D.I. Walker, and H. Kirkman (eds.) Seagrass Biology. Proceedings of an International Workshop. University of Western Australia, Perth. Lee Long, W.J., J.E. Mellors, and R.G. Coles. 1993. Seagrasses between Cape York and Hervey Bay, Queensland, Australia. Australian Journal of Marine and Freshwater Research 44: 19–31. Long, B.G., T.D. Skewes, I.R. Poiner, and C.R. Pitcher. 1995. Torres Strait seagrass survey, November 1993. Final report to the Torres Strait Scientific Advisory Committee. Marsh, H., A.N.M. Harris, and I.R. Lawler. 1997. The sustainability of the indigenous dugong fishery in Torres Strait, Australia/Papua New Guinea. Conservation Biology 11 (6): 1–13. Marsh, H., and W.K. Saalfeld. 1989. Distribution and abundance of dugongs in the northern Great Barrier Reef Marine Park (Australia). Australian Wildlife Research 16: 429–440. Maruanaya, Y. 2000. Studi komunitas lamun dan ikan di taman nasional teluk Cendrawasih Irian Jaya. Master’s thesis, Universitas Hasanudin, Makassar. McKenna, S.A., G.R. Allen, and S. Suryadi (eds.). 2002. A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. Bulletin of the Rapid Assessment Program 22, Conservation International, Washington, D.C. McKenzie, L.J. 1994. Seasonal changes in biomass and shoot characteristics of a Zostera capricorni Aschers. dominant meadow in Cairns harbour, northern Queensland. Australian Journal of Marine and Freshwater Research 45: 1337–52. McKenzie, L.J., S.J. Campbell, and C.A. Roder. 2001. Seagrass-Watch: Manual for Mapping and Monitoring Seagrass Resources by Community (Citizen) Volunteers. DPI&F, Cairns. McKenzie, L.J., M.A. Rasheed, W.J. Lee Long, and R.G. Coles. 1996. Port of Mourilyan seagrass monitoring, baseline surveys—summer (December) 1993 and winter (July) 1994. EcoPorts Monograph Series No. 2. PCQ, Brisbane. Mellors, J.E., H. Marsh, and R.G. Coles. 1993. Intra-annual changes in seagrass standing crop, Green Island, northern Queensland. Australian Journal of Marine and Freshwater Research 44: 33–42. MREP 1996. PSL Unpatti and Direktorat jenderal pembangunan daerah–1996. Studi social ekonomi, budaya dan lingkungan. Munro, J.L. 1999. Utilization of coastal molluscan resources in the tropical Insular Pacific and its impacts on biodiversity. Pp. 127–144 in Maragos, J.E., M.N.A. Peterson, L.G. Eldredge, J.E. Bardach, and H.F. Takeuchi (eds.) Marine/Coastal Biodiversity in the Tropical Island Pacific Region: Vol 2: Population, Development and Conservation Priorities. Workshop proceedings, Pacific Science Association. East-West Center, Honolulu. Nietschmann, B.Q., T.B. Norris, R.S. Rose, and J.M. Roswell. 2000. Coral world map (scale 1:28,510,000) and virtual reefscape poster. In Williams, R.S., J. Steele, H.J. deBlij, and B.S. Nietschmann (eds.) National Geographic Society: Committee for Research and Exploration, Washington D.C. Ogden, J.C., and N.B. Ogden. 1982. A preliminary study of two representative seagrass communities in Palau, Western Caroline Islands (Micronesia). Aquatic Botany 12: 229–244. Petocz, R.G. 1989. Conservation and Development in Irian Jaya. A Strategy for Rational Resource Utilization. E.J. Brill, Leiden.

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822 / len mckenzie , rob c oles , & p a ul erf tem ei jer Phillips, R.C., and E.G. Menez. 1988. Seagrasses. Smithsonian Institution Press, Washington, D.C. Poiner, I.R., D.I. Walker, and R.G. Coles. 1989. Regional studies—seagrasses of tropical Australia. Pp. 279–296 in Larkum, A.W.D., A.J. McComb, and S.A. Shepherd (eds.) Biology of Seagrasses. Elsevier, New York. Preen, A.R., W.J. Lee Long, and R.G. Coles. 1995. Flood and cyclone related loss, and partial recovery, of more than 1000 km2 of seagrasses in Hervey Bay, Queensland, Australia. Aquatic Botany 52: 3–17. Putrawidjaja, M. 2000. Marine species trade in Irian Jaya. Cendrawasih Bay National Park case (report). Conservation Science, WWF-Indonesia-Sahul Bioregion, Jayapura. Pyle, R.L. 1999. Patterns of Pacific reef and shore fish biodiversity. Pp. 157–175 in Eldredge, L.G., J.E. Maragos, P. Holthuis, and H.R. Takeuchi (eds.) Marine and Coastal Biodiversity in the Tropical Island Pacific Region. Volume 2. Population, Development, and Conservation. East-West Center, Honolulu, and Pacific Science Association, Honolulu. Randall, J.E. 1965. Grazing effects on seagrass by herbivorous reef fishes in the West Indies. Ecology 46: 255–260. Roelofs, A., R.G. Coles, and N. Smit. 2005. A survey of intertidal seagrass from Van Deimen Gulf to Castlereagh Bay, Northern Territory, and from Gove to Horn Island. Report to the Australian National Oceans Office, March 2005. DPI&F, Cairns. Rollon, R.N., E.D.D. van Steveninck, W. van Vierssen, and M.D. Fortes. 1998. Contrasting recolonization strategies in multi-species seagrass meadows. Marine Pollution Bulletin 37: 450–459. Scheltze-Westrum, T. 2001. West-Papua: only the village people can save their reefs and rainforests. Biodiversity 2 (1): 15–19. Short, F.T., and S. Wyllie-Echeverria. 1996. Natural and human-induced disturbance of seagrasses. Environmental Conservation 23 (1): 17–27. Short, F.T., R.G. Coles, and C. Pergent-Martini. 2001. Global seagrass distribution. Pp. 5–30 in Short, F.T., and R.G. Coles (eds.) Global Seagrass Research Methods. Elsevier Science, Amsterdam. Short, F.T., W.C. Dennison, and D.G. Capone. 1990. Phosphorus-limited growth of the tropical seagrass Syringodium filiforme in carbonate sediments. Marine Ecology Progress Series 62: 169–174. Short, F.T., L.J. McKenzie, R.G. Coles, and K.P. Vidler. 2002. SeagrassNet Manual for Scientific Monitoring of Seagrass Habitat. DPI&F, Cairns. Spalding, M.D., F. Blasco, and C.D. Field. 1997. World Mangrove Atlas. The International Society for Mangrove Ecosystems, Okinawa, Japan. Spalding, M.D., C. Raviolus, and E.P. Green. 2001. World Atlas of Coral Reefs. Prepared at the UNEP World Conservation Monitoring Centre. University of California Press, Berkley. Spalding, M., M. Taylor, C. Ravilious, F.T. Short, and E.P. Green. 2003. Global overview: the distribution and status of seagrasses. Pp. 5–26 in Green, E.P., and F.T. Short (eds.) World Atlas of Seagrasses. University of California Press, Berkeley. Stapel, J., R. Manuntun, and M.A. Hemminga. 1997. Biomass loss and nutrient redistribution in an Indonesian Thalassia hemprichii seagrass bed following seasonal low tide exposure during daylight. Marine Ecology Progress Series 148: 251–262. Sua´rez, A. 2001. The sea turtle harvest in the Kai Islands, Indonesia. www.arbec.com.my/ sea-turtles/art1julysept01.htm

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Seagrass Ecosystems of Papua / 823 Tomascik, T., A.J. Mah, A. Nontji, and M.K. Moosa. 1997. Seagrasses. Pp. 829–906 in Tomascik, T., A.J. Mah, A. Nontji, and M.K. Moosa. The Ecology of the Indonesian Seas VIII, Part II. Oxford University Press, Oxford. Udy, J., W.C. Dennison, W.J. Lee Long, and L.J. McKenzie. 1999. Responses of seagrass to nutrients in the Great Barrier Reef, Australia. Marine Ecology Progress Series 185: 257–271. Uthicke, S., and C. Conand. 2005. Local examples of beche-de-mer overfishing: an initial summary and request for information. SPC Beche-de-mer Information Bulletin 21: 98–114. Walker, D.I., W.C. Dennison, and G. Edgar. 1999. Status of Australian seagrass research and knowledge. Pp. 1–24 in Butler, A., and P. Jernakoff (eds.) Seagrass in Australia: Strategic Review and Development of an R & D Plan. CSIRO, Collingwood.

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5.4. Mangrove Forests of Papua daniel m. alongi a n gr o v e fo r e s ts are one of the major ecosystems within the coastal zone of Indonesia. They develop best where low wave energy and shelter foster the deposition of fine sediments, and are the only woody plants living at the confluence of land and sea. Evidence of their success is the fact that the standing crop of mangrove forests is, on average, greater than any other aquatic ecosystem. Unlike other tropical forests, mangroves are architecturally simple, being composed of relatively few tree species and often lacking an understory of shrubs and ferns (Figure 5.4.1). Mangrove trees possess morphological and physiological characteristics that make them uniquely adapted to the tidal zone, including aerial roots, salt-excreting leaves, and viviparous water-dispersed young seedlings (i.e., seeds that germinate while still on the parent tree). Mangroves forests are heavily used traditionally for food, shelter, timber, fuel, and medicine. These tidal forests occupy a crucial niche along the Indonesian coast, as they are a valuable ecological and economic resource. Mangroves provide important nursery grounds and breeding sites for fishes, reptiles, birds, crusta-

M

Figure 5.4.1. A mature, mixed Rhizophora-Bruguiera forest in the Fly Delta, Papua New Guinea. Photo: P. Dixon.

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

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Mangrove Forests of Papua / 825

ceans, shellfish, and mammals; accumulation sites for sediment, contaminants, carbon, and nutrients; protection against coastal erosion; and a renewable source of wood (Alongi 2002). This chapter describes the mangrove forests and associated ecosystems in Papua. As the mangroves of Papua are not structurally and functionally different from those in Papua New Guinea, the mangroves of the entire (800,000 km2) oceanic island of New Guinea will be reviewed here. Information about mangroves and their ecology on the other islands of Indonesia can be found in chapters of the other volumes of this series (e.g., Chapter 19 in Tomascik et al. 1997).

Distribution The island of New Guinea has large tracts of mangrove forest with the greatest species diversity of mangroves in the world due to its location bordering the Australasian and Indo-Malesian centers of diversity (Duke 1992). There may be as many as 43 species in New Guinea (Table 5.4.1) with fewer species on the north coast than on the south coast. This disparity of species richness is indicative of a floral discontinuity between the northern and southern sides of the island. Duke (1992) maintains that this is convincing evidence of a fusion of boundaries between two previously isolated and different mangrove floras. Similar floristic discontinuities have also been described for upland plants on the island (Heads 2001). These discontinuities are associated with tectonic events of the collision between the Pacific and Australian plates. The southern coast of New Guinea is a part of the stable Australian plate and has been subjected to alternating episodes of submergence and emergence as a result of glaciation that last took place around 18,000 years ago, when sea level was about 100–150 m lower than at present. In contrast, the north coast of New Guinea lies at the northern edge of the Australian Plate, which has remained submerged. Saenger (2002) notes that the mangroves along the northern shore of the island represent more ancient forests than those along the southern coast; the northern flora is derived from the Indo-Malesian mangroves but the southern flora is largely derived from northern Australia. The geographical isolation of the mangrove flora of the southern and northern coasts of the island is maintained by the high mountain ranges which form the backbone of New Guinea (Milliman 1995). The island of New Guinea contains approximately 34,739 km2 of mangrove forest, of which 13,820 km2 are in Papua and 5,399 km2 are in Papua New Guinea (Darsidi 1984; Soemodihardjo 1986; Soemodihardjo et al. 1993; Spalding, Blasco, and Field 1997; Tomascik et al. 1997). The area of mangroves on the island is subject to considerable uncertainty; the area of Papua mangroves is a crude estimate only, as figures range from 13,000–26,000 km2 (Tomascik et al. 1997). What it is certain is that the mangroves of Papua constitute by far the largest area of mangroves (69–80%) in Indonesia. Mangrove forests in New Guinea are situated in the deltas of large rivers and

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Table 5.4.1. Mangrove plant species and mangrove associates recorded on New Guinea Mangrove species

Distribution

Acrostichum aureum

North coast

Acrostichum speciosum

Ubiquitous

Aegialitis annulata

South coast

Camptostemon schultzii

South coast

Heritiera littoralis

Ubiquitous

Diospyros ferrea

North coast

Aegiceras corniculatum

Ubiquitous

Cynometra iripa

Ubiquitous

1

Lumnitzera racemosa

Ubiquitous

Lumnitzera rosea

South coast

Lumnitzera littorea

Ubiquitous

Pemphis acidula

Ubiquitous

1

Osbornia octodonta

South coast

Sonneratia ovata

South coast

Sonneratia alba

Ubiquitous

Sonneratia gulgai

North coast

Sonneratia caseolaris

North coast

Sonneratia merauke

South coast

Sonneratia lanceolata

South coast

Sonneratia xurama

South coast

Bruguiera gymnorrhiza

Ubiquitous

Bruguiera sexangula

Ubiquitous

Bruguiera exaristata

South coast

Bruguiera hainesii

South coast

Bruguiera parviflora

Ubiquitous

Bruguiera cylindrica

Ubiquitous

Ceriops tagal

Ubiquitous

Ceriops decandra

Ubiquitous

Ceriops australis

South coast

Rhizophora stylosa

Ubiquitous

Rhizophora lamarckii

South coast

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Mangrove Forests of Papua / 827 Rhizophora apiculata

Ubiquitous

Rhizophora mucronata

Ubiquitous

Excoecaria agallocha

Ubiquitous

Xylocarpus granatum

Ubiquitous

Xylocarpus mekongensis

Ubiquitous

Avicennia marina var eucalyptifolia

South coast

Avicennia alba

North coast

Avicennia rumphiana ( A. lanata)

Ubiquitous

Avicennia officinalis

South coast

Acanthus ebracteatus

South coast

Acanthus ilicifolius

Ubiquitous

1

1

Dolichandrone spathacea1

Ubiquitous

Scyphiphora hydrophyllacea

Ubiquitous

Nypa fruticans

Ubiquitous

1

Note: 1denotes a mangrove associate species. Source: Fosberg (1975); Frodin and Huxley (1975); Percival and Womersley (1975); Floyd (1977); Kartawinata et al. (1979); Johnstone and Frodin (1982); Duke and Jackes (1987); Duke (1991, 1992, 1994); Soemodihardjo et al. (1993); Long and McLeod (1997).

along the banks of 253 small and medium rivers (Figure 5.4.2). The large rivers on the island are the Mamberamo, Sepik, Ramu, Markham, Purari, Kikori, Bamu, Fly, Digul, and Palau-Palau rivers, which cumulatively discharge 1.7 billion metric tons of sediment to the adjacent coastal ocean (Milliman 1995). This high fluvial discharge is a result of high rainfall on the island and facilitates the development of large river deltas colonized by extensive inland freshwater and estuarine mangrove forests that can often penetrate quite deeply inland. For example, Sonneratia caseolaris occurs 75 m above sea level and several kilometers inland in southern New Guinea; in the Fly Delta, mangroves can be found 500 km upstream (Saenger 2002) and in Bintuni Bay on the west coast of Papua, mangroves can be found 30 km inland. Often there is little or no discontinuity from the sea to upland forests; the coastal vegetation progressively changes from mangroves to inland freshwater swamp and terrestrial forest (Taylor 1959). Generally, the mangrove forests along the southern and western coasts of New Guinea are more expansive than on the northern and eastern coastlines. Mangroves thus develop best in areas associated with high rainfall. It is in the largest river deltas, where high rainfall and subsequent runoff transports and deposits mud, that the most luxuriant mangrove forests develop. In dry regions of the island, such as near Port Moresby, mangrove forests are reduced in height and are of lower species diversity (Frodin and Huxley 1975).

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Figure 5.4.2. Map of New Guinea showing the major river systems and mangrove forests (blackened areas).

Forest Structure and Zonation The physical settings of mangrove forests are based on the dominance of key physical characteristics: rivers, tides, waves, and sediment type and origin (Woodruffe 1992). The majority of mangrove forests in New Guinea inhabit river- and tide-dominated settings (Cragg 1987), but a great variety of composites of these settings are known (Johnstone and Frodin 1982). Mangroves are typically distributed from mean sea level to highest spring tide with the most conspicuous feature being the sequential change in species either perpendicular or parallel to shore. Mangroves in New Guinea often consist of narrow crowned trees that can attain 30–40 meters in height, although emergents are common in Nypa palm stands. Rhizophora stylosa and Rhizophora apiculata are emergents in Bruguiera cylindrica and Bruguiera exaristata forests (Figure 5.4.3), with a dense ground layer or understory of Nypa proximate to the river bank. Similarly, Avicennia species are emergents in Ceriops tagal forests. The understory, when present, most often consists of Acanthus ilicifolius, Acrostichum speciosum, Excoecaria agallocha, Dalbergia candenatensis, and Maytenus emarginata, and various lianas and scrambling vines (Johnstone and Frodin 1982). There are subtle and complex patterns of species distribution across the intertidal seascape and upstream-downstream, relating to individual species tolerances to abiotic factors (e.g., soil salinity, nutrient status, degree of anoxia [lack of oxygen], degree of soil wetness) and to biotic factors (e.g., competition, predation). Some of these factors come into play over different temporal and spatial scales to control the distribution of tree species, prohibiting generalizations about the mechanisms governing zonation. Many such physical and ecological variations are often expressed within a single estuary (Duke, Ball, and Ellison 1998). For an

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Figure 5.4.3. A mature (⬎ 30 m tall) mixed Bruguiera forest with a dense understory close to the river bank, Fly Delta, Papua New Guinea. Photo: D. M. Alongi.

individual tree, several factors operate to regulate tree growth, including temperature, nutrients, solar radiation, oxygen, and water. Mangrove forests in New Guinea are often naturally disturbed by storms, lightning, tidal surges, and floods, and may take decades to recover (Johns 1986). For instance, many river deltas in Papua and in Papua New Guinea experience tidal bores which are powerful tidal surges that can sweep up a river to destroy entire forests (Figure 5.4.4). Other natural events, such as disease and pests, may not immediately kill trees but can cause stunted growth, slow death, or the replacement of a species. Dieback of mangrove stands has been observed in Papua New Guinea (Arentz 1988) and attributed to either lightning strikes or periods of drought. Johns (1986) similarly reported the death of stands of mangrove forest in New Guinea, attributed to lightning strikes. Regeneration of mangrove seedlings was recorded, but analysis of aerial photographs suggested that mangrove forests affected by previous events had required over 200–300 years to recover fully. In the river deltas of New Guinea, the zonation and distribution of some mangrove forests corresponds to the ‘‘classical’’ zonation parallel to shore, but most do not, as various zonation schemes for mangroves have been overemphasized. Brass (1938), Percival and Womersley (1975), Floyd (1977), Paijmans (1976), Paijmans and Rollet (1977), Green and Sander (1979), Spenceley (1981), Johnstone (1983), and Gylstra (1996) have described zonation of mangroves in various sites around New Guinea and adjacent islands (e.g., the Aru Archipelago). For the

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Figure 5.4.4. The destructive power of tidal bores in the Fly Delta, Papua New Guinea. Photo: D. M. Alongi.

mangroves in New Guinea, Johnstone and Frodin (1982) presented a more realistic depiction of patterns of zonation based on the following factors: tidal range and inundation frequency, degree of wave action, drainage, salinity, substrate type, and composition of biota. Zonation patterns are inconspicuous or absent in flat areas, but become more obvious with increasing ground slope, as water depth and frequency of tidal inundation control the seaward limit of mangroves. A good example of the importance of this factor is the mangrove flora of Galley Reach on the southern coast of New Guinea (Paijmans and Rollet 1977) where there are two large-scale zones of ‘‘true’’ mangroves and ‘‘transitional’’ mangroves. Many species occur in both zones, but some species are restricted to either zone: Bruguiera cylindrica, Bruguiera gymnorrhiza, Rhizophora mucronata, Sonneratia alba, Sonneratia caseolaris, and Xylocarpus granatum in the true mangrove zone and Avicennia rumphiana, Exocoecaria agallocha, Heritiera littoralis, Lumnitzera racemosa, Acrostichum aureum, and Acanthus ilicifolius in the transitional zone. The transition is very distinct, suggesting that floral discontinuity is related to the tides. Drainage and substrate type appear to also be important factors controlling mangrove distribution. The well-drained banks of mangrove creeks are often inhabited by Rhizophora mucronata and Avicennia officinalis, whereas Sonneratia caseolaris is common on poorly drained banks. On coarse-grained and rocky substrates, Aegialtis annulata and Osbornia octodonta are common. Heritiera littoralis, Acrostichum speciosum, and Acanthus ilicifolius frequently occur on biogenic

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structures such as callianassid lobster (Thalassina anomala) mounds that can approach one to two meters in height. In sandier habitats, Avicennia marina, Heritiera littoralis, Ceriops tagal, Ceriops decandra, Lumnitzera racemosa, Lumnitzera littorea, Avicennia rumphiana, and Xylocarpus mekongensis are frequently found, although the latter species often occurs in mud. Salinity is one of the major factors regulating community composition of Papuan mangroves. Along vast expanses of river banks of low salinity, the mangrove palm Nypa fructicans and, to a lesser extent, Sonneratia caseolaris, dominate the vegetation. In high salinity areas where rainfall is low (e.g., near Port Moresby), Ceriops tagal is usually the last mangrove species to be found before the transition to open ground (Frodin and Huxley 1975). Despite that fact that no single factor or simple set of factors regulate the distribution and zonation of Papuan mangroves, large-scale patterns have been defined in specific locations around New Guinea. An ‘‘open coast’’ pattern (where wave action is significant) has been described for the northwest side of Hood Lagoon (Johnstone and Frodin 1982) and in the Raja Ampat Islands in far western Papua (Takeuchi 2003). From the sea to the land, the discernible assemblages in Hood Lagoon are: a beach fringe of Avicennia marina and Sonneratia alba followed by denser stands of Rhizophora stylosa, Rhizophora apiculata, Bruguiera cylindrica, and Bruguiera gymnorrhiza. Further inland, Ceriops tagal and stunted A. marina are common. In the Raja Ampat Islands, mangroves are sparse and species-poor compared with mangroves on the main island of New Guinea, and consist of Bruguiera gymnorrhiza-Rhizophora mucronata associations along the banks of the Gam and Kasin rivers, and a well-developed upstream sequence of Rhizophora mucronataCeriops tagal, Bruguiera gymnorrhiza, and Nypa fruticans with a brackish-freshwater zone composed of Xylocarpus granatum, Dolichandrone spathacea, and Heritiera littoralis. A ‘‘deltaic’’ pattern (where muddy soils and quiescent conditions predominate) has been described for a variety of river deltas and sheltered embayments, such as the Purari and Fly deltas discharging into the Gulf of Papua (Cragg 1983; Robertson, Daniel, and Dixon 1991), Bintuni Bay on the sheltered west coast of Papua (Erftemeijer et al. 1989) and on the banks of the Ajkwa and Tipoeka estuaries in southwestern Papua (Ellison 2005). The mangroves of Bintuni Bay are the most developed and extensive mangrove forests of Papua, covering an area of 618,500 hectares. The most seaward stands are dominated by seedlings and saplings of Avicennia marina and Sonneratia alba. Further upstream, the vegetation is dominated by stands of Rhizophora apiculata, Bruguiera parviflora, and Bruguiera gymnorrhiza. Overwash islands, colonized mostly by Rhizophora apiculata and, to a lesser extent, by Bruguiera parviflora and Bruguiera gymnorrhiza, abound within the embayment. In the Ajkwa and Tipoeka estuaries, Ellison (2005) identified five major mangrove forest types, recording extensive Bruguiera-dominated forests, consisting of Bruguiera cylindrica, Bruguiera parviflora, and Xylocarpus mekongensis, mostly north of the main Ajkwa River mouth. Nypa fruticans and mixed mangrove-floodplain forest dominated areas

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landward in both lower salinities and at higher elevation. At the seaward margins were found Rhizophora-dominated forests, mostly composed of R. stylosa, R. apiculata, and R. mucronata, whereas accreting mudbanks were colonized by pioneering stands of Avicennia marina and Sonneratia caseolaris. Within river channels, high structural diversity was found, apparently in relation to microscale topography. Ellison (2005) noted that tree heights for Bruguiera and Rhizophora often exceeded 25 m. In the Fly Delta of Papua New Guinea, mangroves cover 87,400 hectares mostly on the delta islands (Robertson, Daniel, and Dixon 1991). Twenty-three species of mangroves were recorded, classified into three major forest types: Rhizophora apiculata-Bruguiera parviflora (salinities ⬎ 10); Nypa fruticans (salinities 1–10); and Sonneratia lanceolata-Avicenna marina (accreting banks). On accreting banks in very low salinity areas, S. lanceolata was found in large monospecific stands. In the Purari delta further east of the Fly delta, Cragg (1983) recognized three major types of mangrove forest: fringing, main, and transitional. He also classified mangrove associations for the southern coast of New Guinea and identified groups related primarily to salinity regime (Table 5.4.2). Sonneratia lanceolata is the dominant mangrove found in fringing stands, ranging from the seaward edge to many kilometers inland. In lower salinity, Sonneratia alba, Avicennia eucalyptifolia, and Aegiceras corniculatum are major members of fringing forests, while palms (Pandanus sp. and Nypa fruticans) dominate fringes below a salinity of 2. The main mangrove species is Rhizophora apiculata, followed closely by Bruguiera parviflora and Bruguiera sexangula. Greatest diversity is encountered in the transitional areas

Table 5.4.2. Major species groups of mangroves recorded on the south coast of New Guinea Forest type

Salinity

Species groups

Fringing

⬎ 30

Avicennia eucalyptifolia, Sonneratia alba

10–30

Sonneratia lanceolata, Avicennia alba, Aegiceras corniculatum, S. alba, A. eucalyptifolia

2–10

A. alba, A. corniculatum

⬎30

Rhizophora stylosa, Rhizophora mucronata

10–30

Bruguiera parviflora, Camptostemon schultzii, Rhizophora apiculata, Bruguiera cylindrica, Bruguiera sexangula, Bruguiera gymnorrhiza

2–10

Nypa fruticans, C. schultzii, B. parviflora, Xylocarpus moluccensis

⬍2–10

Excoecaria agallocha, B. gymnorrhiza, Heritiera littoralis, Hibiscus tiliaceus, N. fruticans, X. moluccensis, B. sexangula

Main

Transitional

Source: Modified from Cragg (1983).

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between zones, where true mangrove species, mangrove associates, terrestrial intruders, epiphytes, and climbing plants coexist. The most common mangrove and mangrove associates in this zone are Bruguiera sexangula, Camptostemon schultzii, Dolichandrone spathacea, Diospyros spp., Excoecaria agallocha, Heritiera littoralis, R. apiculata, and Xylocarpus granatum. Several freshwater swamp species invade this zone and frequently develop root structures similar to mangroves. These species include Calophyllum sp., Intsia bijuga, Myristica hollrungii, and Amoora cucullata. In this zone, an understory of Barringtonia, Brownlowia, Inocarpus, Hibiscus, and Cerbera with scattered small palms Areca, Arenga, Metroxylon, and Nypa is often formed. There are marine macroalgae associated with mangroves, particularly with stilt roots of Rhizophora and pneumatophores of Avicennia and Sonneratia (Coppejans and Meinesz 1988; King 1990). In Bintuni Bay, the red alga Gracillaria crassa is very common on the pneumatophores of Sonnertia alba. In the Madang region of Papua New Guinea, 25 species of macroalgae have been recorded, including a ‘‘Bostrychia-Caloglossa’’ association and the genera Caulerpa, Halimeda, Neomeris, Chnoospora, Cutleria, Dictyota, Padina, Catenella, Laurencia, Murrayella, Peyssonnelia, Polysiphonia, and Stictosiphonia (King 1990). Further information is fragmentary, but it appears that macroalgae associated with mangroves in New Guinea are derived from inshore reefs (Tanaka and Chihara 1988).

Forest Biomass and Production The mangrove forests of New Guinea are among the largest on earth, rivaling the height and mass of even the largest tropical rainforests. Figure 5.4.5 provides best estimates of the above-ground biomass of the world’s mangrove forests, including the few data from New Guinea (nearly all of the values between 2 and 8 S Latitude). Mangrove forest biomass ranges from 48 to 580 metric tons dry weight per hectare, with most mature forests being between 100–400 metric tons/ha in weight. The mean weight of all New Guinea mangroves is 285 metric tons/ha. Arguably the New Guinea mangroves are the largest stands yet recorded. Critical to our ability to estimate the role of mangroves in fisheries and wood yield is an accurate estimation of net primary production. This is because primary producers and the carbon they fix via photosynthesis are the crux of mangrove food chains. About 2% of the radiant energy reaching the earth’s surface is used by plants to assimilate atmospheric CO2 into organic compounds used to construct new leaf, stem, branches, and root tissue, as well as to maintain existing tissue, create storage reserves, and provide chemical defense against insects, pathogens and herbivores. Net production is the balance between gross photosynthesis and leaf dark respiration, and represents the amount of carbon available for growth and tissue maintenance. Photosynthesis varies with many factors, especially light intensity, temperature, nutrient and water availability, salinity, tidal range, stand age, species composition, wave energy, and weather. Five methods have been used to measure

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Figure 5.4.5. Mangrove forest biomass as a function of latitude. Nearly all data points between 2–8 S Latitude are from New Guinea. Source: Modified from Alongi (2006).

mangrove forest primary production: litter fall and incremental growth of the stem, harvesting, gas exchange of leaves, light attenuation/gas exchange under the canopy; and demographic/allometric measurements of trees. Litter fall is by far the most common method used because it is inexpensive and easy to measure, but it only measures leaf production and not growth of the remainder of the tree. Two studies have measured mangrove litter fall in New Guinea, but unfortunately, both took place near Port Moresby where rainfall is less than on the rest of the island (Leach and Burgin 1985; Bunt 1995). Both sets of values indicate very high rates of litter fall (⬎ 1,000 g dry weight per m2 per yr). Seasonally, as in other places, maximum litter fall is cued to the onset of the summer wet season (January–March), although different species flower at different times of the year. Harvesting is labor intensive and slow, and accounts for only above-ground production. It often does not account for leaf production. Gas exchange is precise and rapid, although subject to error due to the problem of extrapolating from an individual tree to an entire stand. Moreover, relying solely on gas exchange measurements overestimates net production as it does not account for most tree respiration. Combining measurements offers the best hope of accounting for production of

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all, or most, tree parts. Measuring litter fall and incremental growth of the trunk accounts for all above-ground production, but not below-ground production. Arguably one of the best methods is to measure light attenuation. The method relies on relating the amount of light absorbed by the mangrove canopy to the total canopy chlorophyll content. The early efforts (e.g., Bunt, Boto, and Boto 1979) provided rapid and relatively easy estimates of potential net primary production. The method, however, suffers from lack of actual photosynthesis measurements and a number of untested assumptions based on light attenuation models from temperate forests. Four workers subsequently modified the light attenuation method, combining measurement of light attenuation with a more robust method of calculation of photon flux density at the bottom of the canopy and empirical measurements of leaf photosynthesis (Gong, Ong, and Wong 1991; Gong, Ong, and Clough 1992; Clough 1997; Clough, Ong, and Gong 1997). Litter fall underestimates, and gas exchange overestimates, net primary production, but the modified light attenuation method gives the most reasonable estimate of total production, while litter fall plus incremental growth can give reasonable estimates of above-ground production (and excluding below-ground root production). The modified light attenuation method is most reasonable because it measures total net fixed carbon production and incorporates the most robust assumptions based on tree physiology and carbon balance. A number of recent studies have measured above-ground production using allometry (relationships of tree weight to stem diameter) coupled with litter fall or leaf turnover (Duarte et al. 1999; Coulter et al. 2001; Ross et al. 2001; Sherman, Fahey, and Martinez 2003). The estimates of net primary production made using the modified light attenuation method include below-ground production but there is currently no clear understanding of how carbon is allocated to different parts of the tree. As pointed out by Clough (1998), it is not yet possible to construct a robust model of carbon balance for mangrove trees because of the lack of empirical data and the difficulty of measuring root processes and respiration of woody parts. However, some preliminary carbon data for mangroves suggest that roughly half of carbon incorporated into the tree is respired, an estimate that is in agreement with similar estimates for terrestrial trees (Barnes et al. 1998). If we accept the data obtained using the modified light attenuation method as the most comprehensive estimate of net primary productivity of mangroves, the average rate of net primary production in New Guinea and surroundings (Table 5.4.3) is 51 tons dry weight per ha per yr. There is considerable range between values, but the figures do suggest that mangroves are significant primary producers. This is supported by empirical measurements of rates of leaf photosynthesis in mangroves (Clough and Sim 1989). Measuring gas exchange characteristics and water use efficiency for various mangrove species located on the Era, Wapo, and Ivi rivers along the Gulf of Papua, Galley Reach, and Motupore Island, Clough and Sim (1989) measured the most rapid rates of carbon dioxide uptake, stomatal conductance, and water-use efficiency yet measured. These high rates of CO2 as-

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Indonesia Indonesia

R. stylosa, S. alba

R. apiculata

74.3

63.7

109.4

106.1

103.2

96.9

104.6

24.4

30.1

30.5

29.2

30.6

40.5

23.4

Net Primary Production1

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attenuation

Light attentuation

Light attenuation

Light attenuation

Litter fall/incremental

Method

1

Note: Net primary production given in tons dry weight per hectare per year. Source: Data from Boto, Bunt, and Wellington (1984); Atmadja and Soerojo (1991); Robertson, Daniel, and Dixon (1991); Amarasinghe and Balasubramaniam (1992); Sukardjo and Yamada (1992); Sukardjo (1995); Clough (1998); Alongi, Tirendi, and Clough (2000); Alongi et al. (2004).

Indonesia

C. tagal, R. apiculata

Indonesia

Papua New Guinea

A. marina, Sonneratia lanceolata

C. tagal, R. apiculata

Papua New Guinea

Nypa fruticans

Indonesia

Papua New Guinea

R. apiculata, B. parviflora

A. officinalis, A. marina

Australia

Mixed Rhizophora spp.

Indonesia

Australia

A. marina

Indonesia

Australia

R. stylosa

R. apiculata, A. marina

Indonesia

R. mucronata

R. apiculata, A. marina

Location

Species

Table 5.4.3. Estimated net primary production of mangrove forests of Indonesian islands and tropical Australia

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Table 5.4.4. Rates of CO2 assimilation, stomatal conductance, and water-use efficiency for mangroves in Papua New Guinea Rate of CO2 assimilation (A)

Stomatal conductance (g)

Water-use efficiency (W)

3 7

13.570.36 13.191.11

0.1110.016 0.1310.007

0.550.06 0.460.04

Galley Reach A. corniculatum A. marina B. gymnorrhiza C. decandra H. littoralis R. apiculata X. granatum

4 9 3 12 6 12 6

18.571.48 22.041.02 9.530.94 7.910.56 8.780.51 13.771.06 6.450.46

0.2660.024 0.2700.017 0.1150.018 0.1050.004 0.0920.010 0.1940.011 0.0750.021

0.360.02 0.430.02 0.380.07 0.340.03 0.440.04 0.350.03 0.480.08

Gulf of Papua A. officinalis B. gymnorrhiza B. sexangula H. littoralis R. apiculata R. mucronata R. stylosa X. granatum

5 7 2 2 3 8 3 6

13.191.25 8.910.85 5.781.33 8.570.14 9.840.60 11.260.22 19.131.24 9.941.15

0.2170.027 0.1170.018 0.0790.007 0.1130.018 0.1590.009 0.1860.010 0.2710.005 0.1960.016

0.310.03 0.360.02 0.310.05 0.340.04 0.290.00 0.300.02 0.370.03 0.250.03

Location/species Motupore Is. R. apiculata R. stylosa

Number of leaves (n)

Rates of CO2 assimilation (A) given in ␮mol per m2 per second; stomatal conductance (g) given in mol per m2 per second: and water-use efficiency (W ) given as grams of above ground dry matter produced per kilogram water used. Each value is the average of n leaves exposed to light levels ⬎ 800 ␮mol per m2 per second. Source: Data from Clough and Sim (1989).

similation and other physiological attributes are a reflection of favorable climatic conditions. Plotting all available data on mangrove productivity against latitude (Figure 5.4.6) gives a significant negative relationship, indicating that mangrove production declines away from the equator, mirroring the latitudinal decline in mangrove biomass (Figure 5.4.5) and litter fall (Saenger and Snedaker 1993). These graphs show that the mangroves of Papua are well placed geographically and climatically to grow to immense size and are as productive as any other tropical forests in the world.

Fauna Mangrove forests in Papua support a wide diversity of biota, ranging in size from bacteria to crocodiles, and like most mangroves, house species originating from both land and sea. The fauna of the Papuan mangroves is poorly known. Most information comes from faunal surveys along the southern coast of Papua New

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Figure 5.4.6. Latitudinal changes in net primary production measured using a modified light interception method. Source: Data from Gong, Ong, and Wong (1991); Gong, Ong, and Clough (1992); Atmadja and Soerojo (1991); Robertson, Daniel, and Dixon (1991); Sukardjo (1995); Clough, Ong, and Gong (1997); Clough (1998); Alongi and Dixon (2000); Alongi, Tirendi, and Clough (2000); Alongi et al. (2004).

Guinea and the west coast of Papua. Generally, there is a high level of similarity between the northern Australian and Papuan faunas (Macnae 1968); differences in species recorded are likely due to the lack of surveys in New Guinea rather than any real biogeographical anomalies. The most comprehensive studies of the mangrove fauna of the island have been conducted in the Purari and Fly river deltas bordering the Gulf of Papua (White and White 1976; Liem and Haines 1977; Bayley 1980; Cragg 1983; Liem 1983; Pernetta 1983). The mangrove-associated fish fauna of Bintuni Bay in Papua has also been surveyed (Ecology Team 1984; Erftemeijer et al. 1989), but most of the species lists reflect attention to species of commercial or subsistence use and must be considered incomplete. Of the mangrove vertebrates, 30 species of reptiles, 12 species of amphibians, 250 species of birds, 50 species of mammals, and 195 species of fish have been recorded on the island thus far (Appendix 8.3). Mangrove invertebrates have not been as well investigated, with the exception of commercially valuable groups, such as crabs and shrimps. The best described groups include the mollusks and insects, the most conspicuous of the latter being the Anopheles mosquito which is the vector for malaria and filariasis.

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The mangrove forests of Papua support a rich molluscan and crustacean fauna consisting of approximately 95 and 80 species, respectively (Kartawinata et al. 1979; Sabar, Djajasamita, and Budiman 1979; Kastoro et al. 1991). Numerically, gastropods are the dominant group of mollusks, with Littorina scabra frequently found at the seaward margin in large numbers, with Monodonta labio a co-occurring species (Soemodihardjo 1987). In comparison, bivalves are represented by only a few species, with the genus Enigmonia being dominant in many intertidal regions of the island. At Tatawori estuary in Bintuni Bay, the mangrove fauna is dominated by gastropods and crabs with densities of ⬎120 individuals/m2 of each group, with biomass averaging 10 g DW/m2 (Erftemeijer et al. 1989). The gastropods on the seaward forest edge are dominated by Melampus, but Nerita spp. and Littorina spp. dominate the forest, foraging on algae and detritus on mangrove roots and tree trunks. As in mangroves throughout the Indo-Pacific, the cosmopolitan species Telescopium telescopium dominates the high intertidal areas, scraping detritus and algae off substrata. Ocypodid and grapsid crabs dominate the crustacean fauna in the bay with Uca species being most ubiquitous, followed by species of Sesarma. About one-half of the world’s 60 Uca species are found in Indonesia, with these species exhibiting complex niche patterns to maintain high species diversity compared to other invertebrates (Susetiono 1989). These zonation patterns are the result of niche partitioning of sediment by grain size and organic content. The crabs feed on detritus and microbes attached to the sediment particles using specialized feeding maxillipeds that are unique to each species. In Papua, most of the mangrove species found throughout the rest of Indonesia probably occur, but only Uca dussumieri dussumieri, U. vocans vocans, U. coarctata coarctata, U. lactea annulipes, and U. seismeela have been recorded in Papuan mangrove forests thus far (Moosa and Aswandy 1983). The most commercially important crab, Scylla serrata, is a large carnivorous scavenger that lives in deep burrows within the forest floor and along river banks throughout Indonesia. It exhibits some color plasticity, being dark green in the western archipelago and dark brown in Papua. This species inhabits mangroves throughout its adult life, but females migrate to spawn in waters offshore. Megalopa (crab larvae) move into mangrove waters and by post-larval stage they are sedentary and grow to adulthood in mangrove environments. No quantitative ecological studies are available for the benthic fauna of Papua. Extensive species lists of gastropods, scaphopods, bivalves, and crabs exist for other Indonesian forests and can be found in Tomascik et al. (1997, Chapter 19). Table 5.4.5 summarizes the molluscan species recorded thus far in Papua. The faunal distribution of the mangroves may be considered within four categories (Cragg 1983): permanent residents; animals that also occur in adjacent forest; animals that are strictly estuarine and marine; and animals that spend their early stages in mangroves. Among the permanent residents are the mudskipper (Periophthalmus spp.), the Mud Lobster (Thalassina anomala), the Mud Crab (Scylla serrata), as well as numerous species of isopods (Ceratolana papuae, Bruce 1995)

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Table 5.4.5. Mollusk species recorded from mangroves of Papua and neighboring islands of Indonesia Assiminea corpulenta

M. granifer

A. lentula

M. luteus

A. navigorum

M. nucleolus

A. nitida

Ellobium auris-judae

A. radiata

E. auris-midae

A. sinensis

E. helvaceum

A. sorocula

Cassidula auris-felis

Cerithidea ornata

C. lutescens

Terebralia sulcata

C. mustellina

T. palustris

C. sowerbyana

Telescopium mauritsi

C. sulculosa

T. telescopium

C. triparietalis

Melampus caffra

C. turgida

M. cristatus

Auriculastra helvacea

M. fasciatus

A. oparica

M. obtusus

A. subula

M. ornatus

Nerita planospira

Pythia chrysotoma

Clithon qualaniensis

P. crassidens

Neritodryas cornea

P. imperforata

N. dubba

P. latidentata

Littorina scabra

P. obesula

L. undulata

P. obscura

Crassotrea cucullata

P. pantheriana

Polymedosa cyprinoids

P. pollex

P. divaricate

P. proxima

P. expansa

P. pyramidata

P. kochi

P. scarabaeus

P. nitida

P. striata

P. papua

P. undata

P. subtriangulata

P. variabilis

P. viridescens

P. verreauxi Source: Data from Budiman (1991).

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and brachyuran and sesarmid crabs (Paracleistostoma laciniatum, Baruna trigranulum, Rahayu and Ng 2003; Perisesarma foresti and Perisesarma cricotus, Rahayu and Davie 2002) endemic to New Guinea and the other islands of Indonesia. There are also many other as yet described species that occupy restricted niches in the mangroves of Papua. A variety of wood-boring bivalves (family Teredinidae) are also specialized for life in mangrove wood. Specialist mangrove fauna generally exhibit clear zonation patterns usually in relation to frequency of tidal inundation and salinity (Cook, Currey, and Sarsam 1985; Cragg and Aruga 1987). Animals that occur in both mangrove and neighboring terrestrial forests and swamps are within the second category, and this fauna includes mostly insects, birds, and mammals. These include water rats, bandicoots, bush pigs, wallabies, sugar gliders, possums, bats, and birds such as the Magpie Goose, cassowary, and Brush Turkey (Appendix 8.3). One of the most complex ecological relationships in this category is that between Philidris ants and epiphytic myrmecophytes, Hydnophytum moseleyanum, in mangrove forests in northern New Guinea (Maeyama and Matsumoto 2000). Both Philidris ants and the epiphyte are common throughout the forests of New Guinea, also occupying mangrove trees. The relationship is mutualistic. The epiphyte gets sustenance by absorbing nutrients from the detritus stored inside tree cavities by the ants, and the ants obtain honeydew secreted by scale insects attracted to the shoot tips of the host mangrove tree. The third category, strictly estuarine and marine organisms, enter mangrove creeks and waterways on the rising tide and depart as tides recede. Fish are the predominant animals in this category. As Haines (1983) describes for the Purari delta, few fish species are confined to any one zone but many species are confined to a range of zones with species of the same or closely related genera replacing each other along salinity gradients. Some species are wide ranging, such as the Archer Fish (Toxotes chatareus). The Saltwater Crocodile (Crocodilus porosus) lives only in the seaward limits within waterways, while the Freshwater Crocodile (Crocodilus novaeguineae) occurs in river waters upstream. Some marine invertebrates, such as the penaeid prawns (Table 5.4.6) and the Giant Freshwater Prawn (Macrobrachium rosenbergii), use the mangroves as nursery grounds during their early life stages. The most abundant and widespread species of commercial importance is the Banana Prawn (Penaeus merguiensis). This species is more abundant in regions of high salinity but it exhibits the life cycle typical of all penaeids. During the first stage, planktonic post-larvae settle in estuaries where they feed and grow until adolescence. An oceanic stage begins when the prawns emigrate from the estuaries to coastal waters where, after a period of growth to adulthood, they move into deeper waters to spawn. Penaeus merguiensis, Metapenaeus demani, and Metapenaeus eborancensis spawn in waters 10–20 meters deep, whereas other species such as Metapenaeus ensis and Penaeus semisulcatus spawn in waters up to 60 m deep. The larvae then move shoreward via currents, initially settling in the headwaters of mangrove-lined creeks and waterways. Juveniles and adults congregate in these river mouths before migrating out to sea. The cycle is continuous throughout the year.

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Table 5.4.6. Penaeid prawn species recorded in mangroves of Papua and Papua New Guinea Penaeus merguiensis

Metapenaeus affinis

Penaeus monodon

Metapenaeus lysianassa

Penaeus indicus

Metapenaeus intermedius

Penaeus chinensis

Parapenaeopsis coromandelica

Penaeus penicillatus

Parapenaeopsis coromandelica

Penaeus semisulcans

Parapenaeopsis hardwickii

Penaeus escualentus

Parapenaeopsis hungerfordi

Metapenaeus ensis

Parapenaeopsis maxillipedo

Metapenaeus brevicornis

Parapenaeopsis tenella

Metapenaeus elegans

Trachypenaeus fulvis

Metapenaeus dobsoni

Metapenaeopsis palmensis

Source: Data from Tomascik et al. (1997).

Food Web Dynamics Although little if any information exists on the trophic ecology of New Guinea mangroves, it is assumed that the dynamics of food webs in Papuan mangroves are similar to those in other tropical mangroves. Mangrove links with coastal fisheries have received a lot of attention, but the food webs of mangroves are mostly detritus-based, with most trophic activity focused on interactions among fauna either directly or indirectly through consumption of tree material such as leaves, flowers, propagules, wood, bark, and roots (Robertson, Alongi, and Boto 1992). Direct grazing on mangrove tissue, mainly by insects and arboreal crabs, generally constitutes a small proportion of energy flow. More recently, evidence of the trophic importance of algal food resources in mangrove ecosystems has emerged, demonstrating that a number of key faunal groups depend on phytoplankton, benthic microalgae, or macroalgae growing on above-ground roots and other tree parts, for food. From a nutritional perspective, algae are a better food than detritus derived from mangrove trees because of easier digestion and relatively higher nitrogen content. Various species of mammals, insects, and birds permanently or temporarily reside in some mangrove canopies. The feeding ecology of mangrove-associated birds is fairly well understood. Bird communities can be spatially and trophically complex and include up to eight feeding guilds: granivores, frugivores, piscivores, aerial hawkers, and hovering, gleaning, fly catching, and bark-foraging insectivores (Kathiresan and Bingham 2001). On and beneath the forest floor, crabs are generally the keystone group driving food webs. Sesarmid crabs (Grapsidae) are the most conspicuous organisms, but

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fiddler crabs (Ocypodidae) are also abundant, being highly efficient consumers of benthic microalgae. Recent work throughout the world has shown that large proportions of leaf and other litter deposited on the forest floor is consumed or hidden underground by crabs (Kathiresan and Bingham 2001). This pathway has profound effects on energy and carbon flow within mangrove forests, as the quantities of material available for export from forests are reduced, and the cycling of nutrients to support forest primary production is enhanced. Material that is consumed or hidden by crabs underground must eventually be decomposed by microbial communities in the sediments. The major pathway of trophic dynamics in mangrove sediments is via detritus/ algae to microbe to crab. This is, of course, an overly simplistic depiction of fairly complex interrelations among bacteria, fungi, protozoa, nematodes and other worms, algae, detritus, crabs, and other invertebrates (Figure 5.4.7). In mangrove waters, large swimming organisms, such as fish and prawns, are at the apex of a fairly complex food web in which ‘‘the microbial loop’’ forms a crucial part (Figure 5.4.7). The ‘‘microbial loop’’ is fuelled by the dissolved exudates of phytoplankton, especially from those algal cells broken up by ‘‘sloppy feeding’’ zooplankton. The rates of microbial activity in mangrove waters are thus tightly linked to rates of phytoplankton production. In New Guinea waters, rates of primary production vary greatly depending on the extent to which suspended particulate loads and tides affect turbidity and the availability of light. In the Fly delta, phytoplankton production is highly variable, with rates depending greatly on water clarity (Robertson et al. 1993; Robertson, Dixon, and Alongi 1998). Inside the delta where waters are most turbid, rates are

Figure 5.4.7. A conceptual model of food webs within mangrove forests and in adjacent waterways, dominated by trees, crabs, and ‘‘the microbial loop.’’

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low, ranging from 22 to 95 mg C per m2 per day. At the delta mouth where waters are deeper and less turbid, rates were considerably higher, ranging from 188 to 693 mg C per m2 per day. Rates of bacterial production mirror those of the phytoplankton, suggesting a close trophic link. Zooplankton biomass can be highly variable, weakly correlating with phytoplankton biomass but most often associated with large pieces of mangrove debris floating down river. A similar trophic connection exists in the Purari delta, where phytoplankton production is low in turbid waters but microbial activity is high (Pearl and Kellar 1980). In Indonesian mangrove waters, rates of phytoplankton production are most often light-limited (Soemodihardjo 1987). A complex consortium of microbes is responsible for colonizing and decomposing organic particles, including algal cells, and being the food for many larger planktonic organisms, such as larval invertebrates. Unfortunately, actual rates of trophic transfer from microbes to zooplankton are unknown for Papuan and Papua New Guinea waters. Larger animals such as birds and crocodiles, although highly conspicuous, generally do not play a major role of mangrove energy flow. In the Indo-West Pacific region, most mangrove forests occur in estuaries or as dense forests with intersecting tidal waterways in relatively protected embayments, and have a high proportion of forest to open water. Within such habitats, mangrove vegetation is likely to be the dominant contributor to food webs. Work using stable isotopes confirms that many consumers in mangrove habitats have an isotope signal close to that of mangrove tissue (e.g., Rodelli et al. 1984). In more open mangrove habitats, such as fringing mangroves with open canopies, algae appear to be more important as a food source (Bouillon et al. 2002). Mangrove waterways are often dominated by zooplankton and fish, with densities usually greater than in adjacent habitats. It is generally believed that the higher numbers of organisms in mangroves compared with adjacent habitats is a reflection of greater availability of food, as well as the increased availability of refugia from large predators. In one of the more detailed surveys of fish and their feeding relationships, Haines (1983) found that the fish fauna of the Purari delta is deficient in herbivores and plankton-feeders compared to the fauna in offshore waters. This implies that most fish in mangrove deltas feed primarily on detritus, insects and other invertebrates, and other fish, rather than on algal foods (Table 5.4.7). Prawns are a particularly important prey item for mangrove fish in the Purari, and are the largest contributors to fish biomass. A similar demersal fish fauna is found off the Mamberamo River in northern Papua (Muchtar 2004), suggesting a similar trophic function for the fish species off the northwestern coast of Papua. The same is true for the Markham delta off the coast of East Kalimantan (Dutrieux 1991), implying that there is a consistent fish community structure and function in coastal Indonesia. Although crabs and other invertebrates process large amounts of mangrove detritus (Wada and Wowor 1989), most of the decomposition of this material is mediated by fungi (Ulken 1981) and bacterial assemblages, especially those that are anaerobic (i.e., do not require oxygen). In mangrove and adjacent intertidal

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Table 5.4.7. Fish fauna in the Purari Delta, by trophic zone and feeding category Freshwater zone

Nypa zone

Estuarine mangrove zone

Trophic type

Coastal inshore zone

Omnivores

5 (2)

3 (2)

3 (2)

5 (3)

Piscivores

20 (8)

17 (10)

21 (15)

23 (14)

Mollusk feeders

10 (4)

7 (4)

6 (4)

10 (6)

Prawn feeders

37 (15)

37 (20)

29 (20)

24 (15)

Crab feeders

0

13 (8)

16 (11)

10 (6)

Insectivores

10 (4)

7 (4)

6 (4)

0

Frugivores

5 (2)

3 (2)

4 (3)

5 (3)

Other plant eaters

2 (1)

2 (1)

3 (2)

3 (2)

Detritivores

13 (5)

15 (9)

13 (9)

21 (13)

Note: Numbers in columns are percentages of total fish fauna. Numbers in parentheses are numbers of species. Source: Modified from Haines (1983).

muds of the Fly delta, Alongi (1991) and Alongi, Christoffersen, and Tirendi (1993) found that anaerobic bacterial assemblages were highly active, to the extent that these sediment deposits take up rather than release nutrients that may support food webs in the adjacent Gulf of Papua. Rapid growth of bacteria may be partially maintained by the decomposition and release of nutrients of mangrove roots and rhizomes. A close bacteria-nutrient-plant connection conserves scarce nutrients necessary for growth of the large mangrove forests in the delta (Alongi et al. 1993; Alongi and Robertson 1995).

Links to the Coastal Zone

fisheries Mangrove forests are functionally linked to the biota and abiotic processes (sediment and nutrient flow, water circulation) of the adjacent coastal zone. Nearly all of the evidence of these linkages in New Guinea comes from the mangroves bordering the Gulf of Papua (Alongi and Robertson 1995; Robertson, Dixon, and Alongi 1998). Three types of mangrove-associated fishing practices occur in the Gulf of Papua: gill netting for Barramundi, trawling for prawns; and spearfishing for lobsters. These practices may, to some extent, be exemplary of other coastal zones of Papua and Papua New Guinea. Barramundi (Lates calcarifer) spawn along the coastal strip of the western gulf, in salinities of 30 (Moore 1982; Moore and Reynolds 1982; Reynolds and Moore 1982). Most of the spawning aggregation occurs with the onset of summer, probably triggered by the coincidence of peak spring tides and strong onshore winds. Rich detrital material derived from mangroves and marshes carried offshore by

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ebb tides may be the trigger for spawning offshore. Eggs hatch and incoming tides carry larvae into shallow wetlands. Most predators are excluded by shallow tidal waters, and food is abundant to promote rapid growth of the young. From these nursery areas, most Barramundi migrate eastward to access mangroves inhabiting the river deltas to feed on the abundant prawn populations (Haines 1979). Localized fisheries take advantage of these predictable movements, especially during the summer spawning (Mobiha 1995). From 1971 to 1984, commercial landings declined dramatically from 394 tons/yr to 139 tons/yr. Annual catch from 1993– 1994 was seven tons in the western gulf while the total catch was 58 tons in the eastern gulf (Opnai and Tenakanai 1986; Dalzell, Adams, and Polunin 1996; Kare 1996). This decline can be partly attributed to overfishing and poor management practices. Two prawn fisheries operate along the southern coast of Papua New Guinea. The Torres Strait fishery is dominated by Metapeneaus endeavouri (50%), Penaeus esculentus (40%), and Penaeus longistylus (10%). The fishery in the Gulf of Papua is dominated by Penaeus merguiensis and, to a lesser extent, Penaeus monodon. The prime nursery grounds for the species are seagrass meadows; as the prawns mature they move eastward to deeper waters. For Penaeus merguiensis, the larvae migrate inshore and settle to the bottom as they reach post-larval stage. The principal nursery area for this fishery is the mangrove-fringed islands and channels between the Purari and Kikori rivers (MacFarlane 1980; Evans and Kare 1996). Prawn postlarvae settle in the mangroves in November, grow and recruit to the fishery in February (Evans, Opnai, and Kare 1995). The trawler fleet is one of the largest in the South Pacific, with the annual catch generating up to 1,300 tons/yr (Dalzell et al. 1996). The Gulf of Papua annual average prawn landing was estimated to be 523 tons/yr for 1974–1993 for Penaeus merguinsis, and 844 tons/yr for all the other prawn species (Evans, Opnai, and Kare 1995). The total fish catch off the west coast of Papua in 1997 was 151,133 tons, with most of the catch being tuna, skipjack, and prawns. Rock Lobsters (Panulirus ornatus) are a small, but important, fishery species in the region. Lobster larvae develop in the open ocean, taking about six months to grow to juvenile size, and then settle as post-larvae into the seabed of the Torres Strait (Pitcher 1991). When they are about two and one-half years old they begin a mass migration in August from the strait (MacFarlane and Moore 1986) to coastal reefs in the eastern gulf near Yule Island. Most of the lobsters arrive in poor condition and most apparently die after the breeding season (Dennis et al. 1992). However, it is suspected that some lobsters move to other, unknown, breeding grounds such as deep reef habitats on the edge of the continental shelf (Evans 1996). The lobster exhibit great plasticity of habitat use due to their complex reproductive, migratory, and settlement processes. They are found in a wide range of environments from sheltered, turbid waters to very silty areas near rivers and mangroves. Their diet consists mostly of mollusks and crustaceans (Joll and Phillips 1986). The lobsters are fished at both ends of the migration route and, until

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recently, along the route as well. Because this species is susceptible to being caught by trawlers, lobster by-catch was marketed at up to 200 tons/yr, but concerns about the reduction of spawning stocks and subsequent effects on recruitment led to a ban on keeping the by-catch (Pitcher 1991). In the Daru area, the lobster catch peaked at 92 tons in 1994 with a minimum catch of 57 tons in 1987 (Evans and Polon 1995). The annual catch at the Yule Island end of the migration route is usually two to three metric tons (Dennis et al. 1992). There is a Beˆche-de-Mer (Holothuria scabra) fishery in the Daru area which in 1995 had a total yield of 55 tons, but there is little known about holothurian biology or the sustainability of this fishery. According to estimates of fish resources in the Arafura Sea along the southern coast of Papua (Dalzell and Pauly 1989), potential yields for small pelagic and demersal fisheries are equivalent to those in the Gulf of Papua, with small pelagic yields on the order of 2.8 and 2.5 tons per km2 per yr and demersal yields averaging 1.1 and 1.5 tons per km2 per yr for the Gulf of Papua and Arafura Sea, respectively.

nutrient and sediment fluxes Mangroves are often considered to be accumulation sites for particulate nutrients and sediments, and this appears to be the case for the mangroves of New Guinea. Geological studies of mangroves bordering the Gulf of Papua and the Ajkwa and Tipoeka Rivers in west Papua (Thom and Wright 1983; Barham 1999; Brunskill et al. 2004; Walsh and Nittrouer 2004; Ellison 2005) have indicated that the mangroves are accreting. In west Papuan mangroves, Ellison (2005) estimated sedimentation rates in the range of 0.6–1.5 mm/yr, a rate in the same range as that measured by Thom and Wright (1983) in the Purari mangroves. In a series of contiguous cores, Brunskill et al. (2004) measured sedimentation rates in the Ajkwa mangroves on the order of 4.5–13 kg sediment per m2 per yr, which are well within the range of rates measured in other mangroves. Walsh and Nittrouer (2004) examined sedimentary history of the mangroves bordering the Gulf of Papua and measured accumulation rates ranging from 1.3–7.5 cm/yr (11–65 kg sediment per m2 per yr). Greatest rates of accumulation were observed on accreting banks in the mid-tidal zone, with lower rates above and below this tidal horizon. These figures are higher than those measured in the Ajkwa estuary, but remote sensing indicates that areas of accretion co-exist with areas of erosion in many mangrove/estuarine regions of the Gulf of Papua. Nevertheless, the net accumulation of coastal mangroves of the western Gulf of Papua is estimated to account for 2–14% of the total sediment load of the gulf. All of these sedimentation rates are testimony to the large volume of river sediment discharged from the land to the coastal zone of New Guinea. The discharge of water and sediment from the New Guinea highlands to the coastal plain translates not only into the deposition of particulate material in mangroves, but also into the export of nutrients to the adjacent coastal zone. It is believed that this export of dissolved and particulate material stimulates pelagic and benthic food webs in coastal waters, and supports the fisheries described above.

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In the Gulf of Papua region, there is sufficient evidence to show that mangroves utilize significant quantities of dissolved riverine materials to sustain their high rates of primary production (Liebezeit and Rau 1987; Alongi, Christoffersen, and Tirendi 1993). A budget of carbon gains and losses in the Fly delta (Table 5.4.8) indicates that of the approximately 22.1  1011 g C that comes into the delta from both river discharge and production by mangrove forests, roughly half is consumed in the delta and the other half is exported to the Gulf of Papua (Robertson and Alongi 1995). Roughly 25% of the organic carbon that is exported is mangrove-derived. Most of this material is low-quality detritus, such as leaves, roots, and bark. A study of the carbon-isotope composition of the sediments in the Gulf of Papua (Bird, Brunskill, and Chivas 1995) confirms that this material is exported from the Fly delta, but is limited to within a few kilometers of the coastline. The rapid decline of dissolved nutrient concentrations from the rivers to the adjacent coastal waters off the Fly, Purari, and Mamberamo deltas (Viner 1979; Robertson et al. 1993; Muchtar 2004) suggest similarly important, but geographically limited, export of mangrove material to the coastal ocean bordering the entire island. Extrapolating the sediment and carbon discharge rates for rivers draining into the Gulf of Papua to the rest of New Guinea suggests total sediment and carbon discharge rates for the island similar to those of the Amazon River (Milliman 1995). There is circumstantial evidence that mangrove litter reaches the deep Coral Sea. Considering the much narrower distance from the rivers to the deep sea along the north coast of New Guinea, it is likely that proportionally more mangrovederived matter reaches the deep ocean in the north (Kuehl et al. 2004).

Table 5.4.8. Organic carbon budget for the Fly Delta, PNG Organic carbon budget (X  1011 g C/yr) Inputs River DOC  POC Delta Mangrove POC Delta Water-column TOC

16.6 3.1 2.5

Total inputs

22.2

Outputs Water-column respiration Benthic respiration Sedimentation

3.1 7.0 2.1

Total outputs

12.2

Available for Export (Inputs minus outputs)

10.0

Note: DOC indicates Dissolved Organic Carbon, TOC indicates Total Organic Carbon (a measure of carbon dioxide released by chemical oxidation of organic carbon) and POC indicates Purgeable Organic Carbon (a measure of organic carbon in a sample that can be removed by gas stripping in laboratory analysis). Source: Modified from Robertson and Alongi (1995) using sedimentation data in Walsh and Nittrouer (2004).

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Human Impacts Most of the mangrove forests of Papua are still relatively pristine as human population density is low, and most human use is on a small scale. The only mangroves that have been subjected to a substantial degree of human impact are those near development projects and industries, such as copper mining, capture fisheries, wood chip extraction, and several oilfield projects. Mangrove losses in Papua have been small (⬍10%) but for the islands fringing the Timor Sea, the losses of mangrove forest range from less than 5 to 50% (unpublished references cited in Morrison and Delaney 1996). Table 5.4.9 lists examples of the various human uses of mangrove forest in Papua. The best-documented areas of human impact are the Ajkwa River estuary and Bintuni Bay. The main source of impact in the Ajkwa River estuary is the tailings from a copper-gold mine located some 3,700 m above sea level in the Moake Range. This mine is operated by PT Freeport Indonesia (PRFI) and has been open since 1972. Mine tailings are discharged into the estuary at a rate of about 125,000 tons/day. These tailings consist of sand and small pieces of ground rock, and deposit in a 130 km2 area contained by levees above the salt wedge of the estuary. Despite the levees, recent geochemical measurements (Brunskill et al. 2004) have found that copper accumulation rates have been enhanced 40-fold in mangrove sediments since the introduction of mining. The biological impact of these copper concentrations is unknown. In Bintuni Bay, the mangroves have been increasing affected by wood extraction, fisheries, and oil and gas development (Brotoisworo 1991; Ruitenbeek 1992,

Table 5.4.9. Current human uses of mangroves in Papua Potentially sustainable uses Food Tannins and resins Medicines and other bioproducts Timber Artisanal fishing Charcoal Cage aquaculture Other traditional uses Unsustainable uses Habitat modification/destruction/alteration (pond aquaculture, salt ponds) Damming Release of toxins Introduction of exotic species (ship ballast) Fouling by litter Release of petroleum hydrocarbons Shoreline erosion/siltation Mine tailings

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1994). Most of the primary forest bordering the bay has been allocated for timber concessions (Petocz 1987) with at least seven companies holding concessions of about 300,000 ha of the total mangrove area of 618,500 ha (Erftemeijer et al. 1989; Ruitenbeek 1994). An economic analysis of mangrove management options for Bintuni Bay (Ruitenbeek 1992, 1994) indicates that traditional non-commercial uses of mangroves have an estimated value of US$10 million/yr; commercial prawn fisheries are valued at US$35 million/yr and mangrove wood extraction are worth US$20 million/yr. Ruitenbeek (1994) suggested that the optimal management strategy was selective cutting of 25% of the harvestable mangrove for a total return of US$35 million/yr. Oil concessions to four companies have resulted in extensive oil and gas production, with an estimated recoverable reserve of about 12.2 million barrels (Brotoisworo 1991). With the extensive logging concessions and the fact that the penaeid prawn production of Bintuni Bay was 1,375 tons/yr or about 20% of the total prawn production for Papua, there is significant overlap in resource use in the bay. At present, there does not appear to be effective reconciliation between the need for development and the need for conservation.

Literature Cited Alongi, D.M. 1991. The role of intertidal mudbanks in the diagenesis and export of dissolved and particulate materials from the Fly Delta, Papua New Guinea. J Exp Mar Biol Ecol 149: 81–107. Alongi, D.M. 2002. Present state and future of the world’s mangrove forests. Environ Conserv 29: 331–349. Alongi, D.M. 2007. The Dynamics of Tropical Mangrove Forests. Springer. Alongi, D.M., P. Christoffersen, and F. Tirendi. 1993. The influence of forest type on microbial-nutrient relationships in tropical mangrove sediments. J Exp Mar Biol Ecol 171: 201–223. Alongi, D.M., P. Christoffersen, F. Tirendi, and A.I. Robertson. 1992. The influence of freshwater and material export on sedimentary facies and benthic processes within the Fly Delta and adjacent Gulf of Papua (Papua New Guinea). Cont Shelf Res 12: 287–326. Alongi, D.M., and P. Dixon. 2000. Mangrove primary production and above-and belowground biomass in Sawi Bay, southern Thailand. Phuket Biol Cent Spec Publ 22: 31–38. Alongi, D.M., and A.I. Robertson. 1995. Factors regulating benthic food chains in tropical river deltas and adjacent shelf areas. Geo-Mar Lett 15: 145–152. Alongi, D.M., F. Tirendi, and B.F. Clough. 2000. Below-ground decomposition of organic matter in forests of the mangroves Rhizophora stylosa and Avicennia marina along the arid coast of Western Australia. Aq Bot 68: 97–122. Amarasinghe, M.D., and S. Balasubramaniam. 1992. Net primary productivity of two mangrove forest stands on the northwestern coast of Sri Lanka. Hydrobiologia 247: 37–47. Arentz, F. 1988. Stand-level dieback etiology and its consequences in the forests of Papua New Guinea. GeoJournal 17: 209–215. Atmadja, W.S., and Soerojo. 1991. Structure and potential net primary production of mangrove forests at Grajagan and Ujung Kulon, Indonesia. Pp. 441–451 in Alcala, A.C.

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5.5. Inland Water Ecosystems in Papua: Classification, Biota, and Threats dan a. polhemus and gerald r. allen u e to i t s la r g e si z e, broad elevational range, and great topographic complexity, the island of New Guinea supports a diverse array of inland water ecosystems. All of the major aquatic ecosystem types are represented in Papua, ranging from obvious features such as the Mamberamo and Digul rivers and major lakes such as Paniai and Yamur, to thousands of smaller streams and rivers, a variety of other lakes occupying both lowland and upland basins, innumerable small seeps and springs, and vast coastal wetlands. This extensive array of aquatic ecosystems has in turn developed a rich and highly endemic biota.

D

Aquatic Ecosystem Classification Aquatic ecosystems are in many ways more amenable to classification than terrestrial ecosystems because they possess discrete boundaries and can be unambiguously defined by the presence of water. Even so, terminology has presented a persistent problem in aquatic ecosystem classification schemes, since different authors have often employed ecological terms such as ‘‘habitat’’ and ‘‘ecosystem’’ in different contexts and then discussed these terms without providing the necessary definitions. As defined by Polhemus et al. (1992), individual aquatic ecosystems consist of a watermass with relatively sharp, delineable boundaries, enclosing resident organisms, and possessing discrete physiochemical features. Commonly encountered examples of such ecosystems include lakes, marshes, streams, and estuaries. Two basic components of such ecosystems are the biota, or totality of living matter, and the environment, which represents the nonliving physiochemical components of the ecosystem, including spatial dimensions. Although the term ‘‘habitat’’ has often been used more or less interchangeably with ‘‘ecosystem,’’ it is more properly applicable in an autecological sense to designate all ecosystem requirements of a resident species, including space. Habitats are thus not spatially exclusive subdivisions of an ecosystem, in contrast to divisions such as zones, strata, and reaches. The various types of aquatic ecosystems present in Papua may be assigned to major divisions or classes, for example lotic (flowing) versus lentic (standing). Such classes are based on descriptions of both broad-scale environmental features, such as hydrological regime, water depth, salinity, and so on, and characteristic major taxa of the biota (with the fauna being generally more distinctive and useful in this regard than the flora). Within these broad classes, discrete ecosystem types may be defined by criteria that include altitude, topography, water character (e.g., Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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temperature, turbidity), cultural influences (environmental and biological), and the presence of individual genera or species. In general, oxygen content and pH are usually not adequate descriptors at the ecosystem level because in many waters, particularly those of low ionic content and abundant flora, strong photosynthesis and respiration can diurnally change the levels of such characteristics significantly (pH sometimes by more than two units). Utilizing the system outlined above and further explained in Polhemus et al. (1992), at least 15 types of inland aquatic ecosystems may be recognized in Papua, occurring across a wide range of elevations. These ecosystems may be split into two major divisions: surface and subterranean. The latter ecosystem types are poorly investigated in Papua, and are not dealt with in great detail here. Surface water ecosystems, by contrast, have received considerable attention, and may in turn be divided into two major ecosystem classes, lotic and lentic, within which there are many individual ecosystem types, discussed in greater detail below. See Definitions of Limnological Terms and Units, on the next page, for specific terms employed in discussion of these ecosystem types.

Lotic Ecosystems Lotic ecosystems may be technically defined as limnetic surface waters flowing unidirectionally down altitudinal gradients, and may be divided into four types (Polhemus et al. 1992): perennial streams, intermittent streams, rheocrenes, and artificial ditches and flumes. New Guinea lotic ecosystems are distinguished biologically by a flora consisting mainly of mosses, filamentous algae, and epilithic diatoms, a diverse and largely endemic aquatic insect biota including numerous species of Diptera, Trichoptera (Neboiss 1986a,b,c, 1987, 1989, 1994; Wells 1990, 1991), Ephemeroptera (Demoulin 1954; Grant, 1985; Edmunds and Polhemus 1990), Odonata (Lieftinck 1932, 1933, 1935, 1937, 1938, 1949a,b, 1955a,b, 1956a,b, 1957, 1958, 1959a,b, 1960, 1963), Coleoptera (Ochs 1925, 1955, 1960; Brinck 1976, 1981, 1983, 1984; Gentili 1980, 1981, 1989; Balke 1995, 1999, 2001; Balke and Hendrich 1992a,b; Balke et al. 1992, 1997, 2000; Bistrom et al. 1993), and Heteroptera (Andersen 1975; Baehr 1990; Brooks 1951; Hungerford and Matsuda 1958; Kormilev 1971; Lansbury 1962, 1963, 1965, 1966, 1968a,b,c, 1969, 1972, 1973, 1974, 1975, 1993, 1996; D. Polhemus 2002; D. Polhemus and J. Polhemus 1985, 1986a,b, 1989a,b, 1997, 1998, 2000a,b,c,d, 2001; J. Polhemus and Lansbury 1997; J. Polhemus and D. Polhemus 1987, 1990, 1991, 1993, 1994a,b, 1995, 2000, 2001, 2002; Todd 1955, 1959), but with very limited representation of Plecoptera and Megaloptera. In addition, such streams support a rich non-insect macrofauna of fishes (Allen 1991, 1996a, 2003a,b; Allen et al. 2000; Chapter 4.8), crustaceans (Bott 1974; Holthuis 1939, 1950, 1956, 1958, 1982, 1986), and mollusks (Haynes 2001), many of which are diadromous, with marine larval development. Comprehensive faunal surveys have been undertaken for only a few of the major river basins in Papua, notably the Ajkwa River and portions of immediately

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DEFINITIONS OF LIMNOLOGICAL TERMS AND UNITS Physiochemical Measurements o/oo: parts per thousand, a measure of salinity % gradient: relative slope measured as the unit of elevational change per 100 horizontal units (as in m/100 m) ␮mhos: reciprocal megohms, a measure of water conductivity Water Regime Lacustrine (lake-like): deeper open standing waters occupying distinct basins; lakes and ponds Lentic (standing): water not subject to direct gravitational movement, although internal currents may occur Limnocrene (spring pool): a pond or pool having a noticeable, discrete, subterranean water source (cf. rheocrene) Lotic (flowing): water moving unidirectionally in response to substrate altitudinal (elevational) gradient; excludes waters moving in response to wind currents, waves, and tides Palustrine (marsh-like): shallow standing water visually dominated by emergent vegetation such as mosses, sedges, rushes, trees, etc. Rheocrene (flowing spring): lotic water from a subterranean source but not in a well-developed channel, and flowing in relatively low and constant volume Dissolved Minerals Qualitative aspects Haline (halinity): brackish or salty water condition wherein dissolved ions are derived from seawater Saline (salinity): general term for water with noticeable salt content Quantitative aspects Limnetic: freshwater, salt content ⬍0.5 o/oo Mixohaline: brackish water, salt content 0.5–30 o/oo Euhaline: seawater, salt content 30–40 o/oo Hyperhaline: brine-like water, salt content ⬎40 o/oo Concentration vs. time Homiohaline: salt concentration stable or fluctuating only over a narrow range Poikilohaline: salt concentration fluctuating widely Ecological Qualifiers Migration and movement Amphidromous: type of diadromous animal (see below) that migrates between fresh and marine waters but not for breeding (e.g., sicydiine gobies) Catadromous: type of diadromous animal that inhabits freshwater but breeds in the ocean (e.g., anguillid eels)

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Inland Water Ecosystems: Classification, Biota, and Threats / 861 Diadromous: broadly referring to animals (e.g., certain fishes) that obligately migrate between fresh and marine waters during their life cycle Itinerant: refers to animals that may irregularly or opportunistically migrate between fresh and marine waters (e.g., haline marine fishes sometimes found in streams) Salt tolerance of biota Euryhaline/saline: occurs over a wide range of total dissolved solids Stenohaline/saline: occurs in a narrower range of total dissolved solids Substrate relationship Benthic: living on or in the bottom of a water body Epigeal: living on or above the earth’s surface Hypogeal: living beneath the earth’s surface ( subterranean)

adjacent systems (the Minajerwi and Iweka) draining the southern flank of the central ranges in the Timika region (D. Polhemus and J. Polhemus 2000d; Allen et al. 2000); the Wapoga River draining the western section of the central mountain ranges (D. Polhemus 1998; Allen and Renyaan 2000); and the lower reaches of the Idenburg (Taritatu) River (the major eastern branch of the Mamberamo River system) plus various tributaries (the Furu, Doorman, and Tiri) in the vicinity of Dabra (Polhemus 2002; Allen et al. 2002).

perennial streams Perennial streams (Figures 5.5.1–5.5.5) support continuous year-round flow and form the most widely distributed type of lotic ecosystem in Papua. Although the majority are continuous, discharging steadily to the ocean in their natural state, there are certain karst areas, particularly in the central mountain ranges and on the Vogelkop and Bomberai peninsulas, where such streams may be naturally interrupted, with their flow becoming subsurface in their middle or lower sections, although occasionally appearing as scattered pools in areas of bedrock exposure. In larger towns or near industrial developments, streams may also be artificially interrupted via partial or total diversion. Such human-made diversions are generally accompanied by channel alterations that in many instances modify or eliminate the native ecosystem character, particularly in urban areas. Altered streams of this type in lowland areas are often favored habitats for invasive aquatic species. Because of the relatively intact nature of Papuan forests, the water clarity of undisturbed streams, at least in the smaller order streams of a given network, is generally high except during spates, and dissolved oxygen is normally near saturation throughout most watercourses. Papuan perennial streams exhibit prominent altitudinal zonation of environmental conditions and biota (Allen et al. 2000; D. Polhemus and J. Polhemus 2000d). This longitudinal continuum may be divided into three broad zones: the headwater, mid-, and terminal reaches, described in greater detail below. Naturally interrupted streams also exhibit a similar zonation, except that the amount of

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available habitat in their mid- and terminal reaches is often significantly reduced. As such, their diadromous macrofauna, although similar to that of continuous perennial streams, is generally less diverse. Despite this, such streams may contain certain genera of diadromous gobioid fishes that access the upper reaches during intermittent spates that provide temporarily continuous water connections to the sea, and then hold over in the upper reaches until the next flood.

Headwater Reach Headwater streams (Figure 5.5.1) drain first and second order catchments lying at elevations above 800 m or possessing gradients in excess of 30%; streams of this type in montane regions possess both such attributes. The substratum is usually

Fig. 5.5.1. This cascading tributary of the Kikori River, on Mt Bosavi in south-central New Guinea, is a typical high gradient headwater reach of a perennial stream, with extensive bedrock exposures in the stream channel. Photo: D. A. Polhemus.

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bedrock or coarse alluvium such as large rocks and boulders; current speeds are generally high; water temperature is less than 18 C (most typically 12–15); conductivity is less than 50 ␮mhos (with dissolved solids less than 40 mg/l); and pH is often slightly acidic. The fauna of such small, steeply dropping streams is dominated by insects, particularly Ephemeroptera in the family Baetidae, Trichoptera in the families Glossosomatidae, Hydrobiosidae (Apsilochorema), Hydropsychidae (Hydropsyche), Hydroptilidae (Hydroptila), and Leptoceridae (Mystacides, Triaenodes), Diptera in the families Dolichopodidae and Ephydridae, Odonata in the families Platystictidae (Drepanosticta), Megapodagrionidae (Argiolestes), and Platycnemididae (Idiocnemis), Coleoptera in the families Dytiscidae (Platynectes), Hydrophilidae (Enochrus), and Gyrinidae (Merodineutes, Macrogyrus), and Heteroptera in the families Naucoridae (Nesocricos, Tanycricos), Gelastocoridae (Nerthra), and Veliidae (Rhagovelia, Papuavelia, Tarsovelia), with crustaceans scarce, and fishes generally absent above 2,000 m (Allen 1991). Alpine streams, lying at elevations above 3,000 m, lack most of these biotic elements except for a very limited and specialized assemblage of Trichoptera in the families Hydrobiosidae (Apsilochorema), Hydropsychidae, and Hydroptilidae and Diptera in the family Chironomidae.

Midreach The midreach zone (Figures 5.5.2 and 5.5.3) is intermediate in environmental conditions between the headwater and terminal reaches (for discussion of terminal reaches, see below). Depending on the length and gradient of an individual stream catchment, this can either be a brief and highly foreshortened zone, or conversely may comprise the majority of a stream’s length. The substratum is usually mixed alluvium consisting of boulders, rocks, and gravel, with occasional sand or cobble bars developing on the inner margins of bends, and water temperatures typically range between 18 and 24C. The fauna of such streams in New Guinea is diverse, including plotosid catfishes, rainbowfishes (Melanotaenidae), grunters (Terapontidae), and gobioid fishes (Gobiidae and Eleotridae), Crustacea in the family Parastacidae (Charax), Ephemeroptera in the families Baetidae, Leptophlebiidae (Thraulus), and Prosopistomatidae (Prosopistoma), Trichoptera in the family Hydropsychidae (Cheumatopsyche, Hydropsyche, Macrostemum), Zygoptera in the families Calopterygidae (Neurobasis), Chlorocyphidae (Rhinocypha), Coenagrionidae (Palaiagria, Pseudagrion, Teinobasis), Platycnemididae (Idiocnemis), and Protoneuridae (Nososticta, Selysioneura), Anisoptera in the family Libellulidae (Huonia), Lepidoptera in the family Pyralidae, Diptera in the family Chironomidae, Coleoptera in the family Gyrinidae (Rhombodineutes), and Heteroptera in the families Gerridae (Ptilomera, Tenagogonus, Limnometra, Stygiobates, Metrobatoides), Veliidae (Rhagovelia, Strongylovelia, and numerous genera of Microveliinae), Mesoveliidae (certain Mesovelia), Ochteridae (Ochterus), Leptopodidae (Valleriola), Hebridae (Hebrus), Nepidae (Cercotmetus), and Naucoridae (Sagocoris, Aptinoocoris, Cavocoris, Idiocarus).

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Figure 5.5.2. The upper midreach zones of perennial stream networks on New Guinea are composed of clear, rocky creeks, such as this tributary to the Ziwa River in the central mountains of the island. Such streams are numerous in forested upland terrain, and support extremely diverse biotas of native aquatic insects but only a limited array of native fishes. Photo: D. A. Polhemus.

Terminal Reach The terminal reach (Figures 5.5.4 and 5.5.5) is the section of a watercourse below the first sharp gradient that bars upstream migration of itinerant marine fishes, such as flagtails (Kuhlia). The elevation is generally less than 50 m (although though this may not be reached until far inland on large systems such as the Digul, Wapoga, and Mamberamo), and gradient is less than 5%. The substratum in larger streams and rivers is primarily fine sediment, intermixed to varying degrees with sand, gravel, and rocks. The water temperature is greater than 24C (generally 25–27); conductivity exceeds 80 ␮mhos (mainly 100–150 ␮mhos); dissolved solids are 60–100 mg/l; and pH is neutral to slightly basic, ranging from 6.5–7.8. Large lowland rivers such as the Digul and Mamberamo fall within this division, as do the lower courses of many rivers draining from the southern face of the central mountain ranges to the Arafura Sea. Characteristic fauna includes Barramundi (Lates calcarifer), ariid catfishes, numerous gobioid fishes (Gobiidae and

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Figure 5.5.3. The midreach of the Minajerwi River as it exits the southern foothills of the Papuan Central Ranges north of Timika is indicative of the high velocity and base flow exhibited by many rivers in the mountains of New Guinea. The coarse alluvial character of the bed is in marked contrast to the fine sediments of the terminal reach channel occupied by this same river just a few kilometers downstream. Photo: D. A. Polhemus.

Eleotridae), Ephemeroptera in the family Palingeniidae (Plethogenesia), Odonata in the families Coenagrionidae (Agriocnemis, Argiocnemis, Pseudagrion, Xiphiagrion) and Libellulidae (Agrionoptera, Huonia, Neurothemis, Orthetrum, Rhyothemis, Tetrathemis), Coleoptera in the family Gyrinidae (Spinosodineutes), Heteroptera in the families Gerridae (Rhagdotarsus, Limnometra, Limnogonus, Ciliometra), Veliidae (Microvelia), Mesoveliidae (certain Mesovelia), Notonectidae (Nychia) and Corixidae (Micronecta). The terminal reaches of the larger lowland rivers in Papua experience periodic mass hatches of large-sized palingeniid mayflies, with body lengths exceeding 3 cm, which create an impressive sight on the water and represent an important food source for native fishes. The above three stream divisions may each be further segregated into two zones, erosional and depositional. Erosional zones include waterfalls, rapids, riffles, and other areas where there is a net loss of substrate or organic material due to the action of flowing water. Depositional zones include pools, the inner margins of stream bends, and other areas where such material is deposited. Since these two types of zones alternate and intergrade along the length of any given reach, they are not considered as discrete ecosystems per se, but they are often important habitat determinants for individual taxa. For example, the goby genus Stenogobius is generally restricted to depositional zones.

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Figure 5.5.4. Terminal reach streams in lowland areas, such as this rainforest creek in the Wapoga River basin, typically occupy low gradient beds of fine alluvium, and are among the faunally richest aquatic ecosystems in New Guinea in terms of both fishes and insects. Photo: D. A. Polhemus.

intermittent streams Intermittent streams (Figure 5.5.6) comprise seasonally flowing waters in discrete channels, with flow decreasing in volume to slow-exchanging pools prior to desiccation. Examples of such ecosystems include small creek beds in lowland alluvial forests, such as those seen near Kuala Kencana in the Timika area, and periodically flooded overflow channels adjoining larger rivers (Figure 5.5.6), which carry water briefly during spates, or for more prolonged periods during the rainy season. Pools in such systems generally persist for at least a few weeks to a few months, usually as discontinuous surface manifestations of diminishing hyporheic (subsurface) flow. Ecosystems of this type most frequently develop in porous, sandy channels where flow can readily retreat subsurface, and although water quality is variable along such reaches, in some cases becoming stagnant, it is often high due to some slight degree of flow coupled with natural sand filtration. Because intermittent streams differ environmentally from perennial streams in terms of flow regime and water continuity, they are biologically distinct in lacking many diadromous species (those that migrate between fresh and salt water), but by contrast often contain abundant insects and other small invertebrates that may be rare elsewhere. Such ecosystems generally support an impoverished fish biota except in the largest and most persistent pools, where fishes such as blue-eyes

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Figure 5.5.5. The Tirawiwa River upstream from its confluence with the Wapoga River, in north-central New Guinea, is typical of a low gradient, terminal reach river flowing across alluvial lowland terrane. Photo: D. A. Polhemus.

Figure 5.5.6. Overflow channels, such as this one along a tributary to the Doorman River near Dabra, in north-central Papua, function as intermittent streams, carrying surface flow during the wet season and then receding to scattered pools fed by hyporheic flow during the drier months of the year. Photo: D. A. Polhemus.

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(Pseudomugil), hardyheads (Craterocephalus), rainbowfishes (Melanotaenia), and glassfishes (Ambassis) may be present. The fish-free pools of these systems serve as important breeding refugia for a variety of aquatic insect species, however, including the Zygoptera in the families Protoneuridae (Nososticta), Platycnemididae (Idiocnemis), and Coenagrionidae (Teinobasis), Heteroptera in the families Gerridae (Tenagogonus, Limnometra), Hydrometridae (Hydrometra), Veliidae (Microvelia), Notonectidae (Enithares, Anisops), Corixidae (Micronecta), and various aquatic Coleoptera in the families Dytiscidae, Hydrophilidae, and Gyrinidae (Macrogyrus).

rheocrenes Rheocrenes, literally ‘‘flowing springs,’’ are perennial seeps and springs flowing short distances over rock surfaces (Figure 5.5.7) or in indistinct channels. These numerous, ubiquitous small seepages (which represent ‘‘leaks’’ from elevated aquifers) are typically found as natural occurrences on bedrock faces or banks of

Figure 5.5.7. Rheocrenes, such as this seeping bedrock face and outflow pool near Etna Bay, are common in the mountains of New Guinea and support a highly specialized insect biota rich in endemic Coleoptera, Heteroptera, and Odonata. Photo: D. A. Polhemus.

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deeply incised streams (particularly adjacent to waterfalls), as well as artificially along road cuts. Water quality of such ecosystems is variable, with their waters sometimes iron-rich as evidenced by bacterial precipitation of orange ferric hydroxide. Two divisions can be recognized: thermal, with average water temperature noticeably (at least 10C) above the mean annual temperature of the air at the same locality, and non-thermal, with water temperature near or below the mean annual air temperature. Non-thermal rheocrenes, by far the most common type, have a distinctive and often highly endemic biota consisting of a flora of algae, mosses, ferns, Diptera in the family Dolichopodidae, Odonata in the families Megapodagrionidae (Argiolestes) and Corduliidae (Hemicordulia), Coleoptera in the families Dytiscidae and Hydrophilidae, and Heteroptera in the families Gelastocoridae (Nerthra), Ochteridae (Ochterus), Hebridae (Hebrus), Saldidae (Saldula), and Microveliinae (Aegilipsicola, Rheovelia, Brechyvelia). Although they may support distinct algal communities, thermal rheocrenes in general tend to have a highly impoverished fauna, as is typical of thermal waters in general throughout Papua.

artificial ditches and flumes Ditches and flumes are artificial streams constructed by humans to convey water to areas where it would otherwise not naturally flow. Such conduits often pass over or through ridges, and thus transgress natural drainage divides. Although some ditches were built by prehistoric Papuans for crop irrigation, particularly in the Baliem Valley, most were constructed during the past century for municipal water supplies or to provide drainage in urban and industrial areas. At higher elevations, such systems are generally associated with mining developments, such as the Grasberg mine above Timika, while at middle elevations they include a wide array of local water supplies ranging from rudimentary split bamboo flumes to joined plastic pipes many kilometers long. The environmental character and biota of ditches and flumes differs with their location and degree of use, but in upland Papua they often contain high-quality water comparable to midreach stream water. In general, a lack of shelter and slack water results in low faunal diversity, with the most prominent biotic elements being certain aquatic mollusks, Diptera, Heteroptera, Odonata, and occasionally atyid shrimps, with fishes generally scarce. In lowland areas, particularly in or near larger towns, effluent ditches are commonly constructed to remove water from small reservoirs, agricultural sites, and use facilities. Water quality in such effluent ditches is moderate to low, and the macrofauna, when present, consists mainly of introduced fishes and hardy invertebrates.

Lentic Ecosystems Lentic ecosystems are defined as standing or still waters, generally in definite basins. They may be divided into two types, lacustrine and palustrine, depending on the type of basin they occupy.

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lacustrine systems (lakes and ponds) Lacustrine ecosystems are standing waters occupying definite basins with discrete shorelines, and have predominantly open water with depth generally exceeding 2 m. Such ecosystems are numerous in Papua, and vary greatly in size, from small montane ponds to large lowland lakes, and even flooded World War II bomb craters.

Natural Lakes and Ponds This ecosystem class comprises natural freshwater lacustrine ecosystems (Figures 5.5.8–5.5.12) with salinities less than 0.5 o/oo, including limnocrenes (pond-like springs with subterranean limnetic water sources). Although relatively uncommon in the tropical Pacific as a whole, such ecosystems are well represented in Papua due to its young, rugged topography and extensive karst exposures. These factors have combined to create many areas with blind or poorly integrated drainages,

Figure 5.5.8. High alpine lakes, such as these formed at the base of melting glaciers near the summit of Mt Jaya, are numerous along the crest of the highest mountains in western New Guinea, but relatively poor in aquatic biota. Photo: D. A. Polhemus.

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Figure 5.5.9. Lake Andersen, near the Grasberg mine, is typical of alpine lakes formed in strike valleys amid the tilted limestone layers of the central mountains of Papua. Note the series of lakes that occupy basins of progressively lower elevation along the westward strike of the valley. Such ecosystems contain specialized suites of aquatic insects such as Coleoptera and Trichoptera, and also serve as important habitat for waterfowl such as Salvadori’s Teal. They are devoid of native fishes. Photo: D. A. Polhemus.

particularly in the Lengguru Fold Belt of the Vogelkop region (Figure 5.5.10), and to a smaller extent in the strike valleys along the crest of the central mountains (Figure 5.5.9). The lake systems of Papua are extensive, and each individual lake is distinctive in terms of its location, environmental features, and native biota (for a list of major lakes in Papua and their endemic biota (see Table 5.5.1). The natural biotas of such systems in the lowlands include many fishes, especially melanotaeniids and gudgeons, as well as Zygoptera in the families Lestidae (Lestes) and Coenagrionidae (Agriocnemis, Ischnura) and Heteroptera in the families Gerridae (Limnogonus), Mesoveliidae (Mesovelia), and Veliidae (Microvelia), in addition to a diverse waterfowl assemblage (Erftemeijer and Allen, 1989). These systems are frequently degraded by introduction of exotic food fishes, particularly carp and tilapia (see further discussion later in this chapter). By contrast, alpine lakes, such as Lake Andersen near the Grasberg mine (Figure 5.5.9), support a limited and specialized insect biota of Diptera in the family Chironomidae, and Coleoptera in the family Dytiscidae (tribe Bidessini), and also represent important habitats for native waterfowl such as Salvadori’s Teal.

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Figure 5.5.10. Lake Laamora in the Bird’s Neck region of Papua, shown here in the dry season, is trapped amid a parallel series of limestone anticlines upthrust in the Miocene, and has a seasonally fluctuating water level. It supports an endemic rainbowfish species. Photo: G. Allen.

Artificial Reservoirs Reservoirs consist of lacustrine waters occupying artificial basins or human-made impoundments. In contrast to other parts of the world, Papua lacks large-scale reservoirs created by high dams, although many smaller artificial basins and impoundments built for various purposes are scattered throughout the province, intermixed in a few areas along the north coast with small artificial basins created by warfare, in the form of water-filled bomb craters. In addition, the construction of logging roads often inadvertently creates unplanned reservoirs of varying size and duration due to the obstruction of streams by culverts and other temporary crossings. The environmental quality of such reservoirs and their resulting biotic assemblages vary with reservoir type. Primary storage reservoirs, built primarily for agricultural and domestic water supplies, frequently lie on or near source streams in remote upland sites, and have relatively stable surface levels and good water quality. As such, they often support submerged and floating water flora, which in turn provides excellent habitat for aquatic insects and mollusks. Distributional reservoirs, by contrast, lie mainly on agricultural lands or in populated areas, and are used mostly for temporary water storage and redistribution. As a result, they have fluctuating surface levels and moderate to poor water quality, with their waters often turbid. Accidental reservoirs created by warfare or road construction vary widely in environmental quality and may exhibit any of the above characteristics.

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Figure 5.5.11. Lake Sentani, near Jayapura, is formed in a basin lying between the accreted terrane of the Cyclops Mountains to the north (right side of picture), and the foothills of the Foja Mountains to the south (left side of picture). It contains at least three species of endemic fishes. Photo: D. A. Polhemus.

Although they are artificial ecosystems in terms of basin origin and structure, such ecosystems are often colonized by dispersive native lowland aquatic insect species, particularly wide-ranging Anisoptera in the families Libellulidae (Pantala, Diplacodes) and Aeschnidae (Anax), and Heteroptera in the families Belostomatidae (Appasus), Nepidae (Ranatra), Notonectidae (Anisops), Veliidae (Microvelia), and Mesoveliidae (Mesovelia). They may harbor rainbowfishes and a few other native fish species, but are also frequently stocked with livebearers and rivulines (for mosquito control) as well as food fishes such as tilapia.

Saline Lakes Although present as shoreline features or closed lagoons on other islands in the region with seasonally drier climates, particularly Timor and the Lesser Sunda Islands, saline lakes have not been documented in Papua. The shores of such lakes, which have waters with salinities exceeding 0.5 o/oo (and in some cases exceeding 100 o/oo), typically support a limited but extremely distinctive halophytic (saltadapted) biota, particularly among certain insect groups such as Diptera in the family Ephydridae and Heteroptera in the family Saldidae (Pentacora, Micracanthia, Saldula).

palustrine systems Palustrine ecosystems, more commonly referred to as ‘‘wetlands,’’ comprise various types of swamps and marshes with lentic waters less than 2 m deep (usually

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Figure 5.5.12. The vast wetlands bordering the Mamberamo River in the Meervlakte basin form a complex mosaic of lotic and lentic ecosystems, the latter including both lacustrine and palustrine components. Photo: D. A. Polhemus.

⬍1 m), occupying irregular or poorly-defined basins (Figures 5.5.12–5.5.15). This category encompasses a broad array of individual ecosystems that tend to form a continuum, and creating an unambiguous classification to accommodate the full range of variation involved has proven problematic, but as a general rule forested wetlands are considered to be swamps, while open, non-forested wetlands are classified as marshes. Palustrine ecosystems include wetlands at both high and low elevations, each with several types. Elevated wetlands, located in remote areas, are primarily natural systems in which native biota dominates, good examples being the marshes that are frequently encountered in the upland valleys of the central mountain ranges. Low elevation wetlands have in most parts of the Asia-Pacific region been severely modified by humans (IUCN, 1991), but in Papua they retain a largely natural character (Figure 5.5.13), due to a general absence of rice cultivation and consequent channelization. This situation is changing, however, in the Merauke and Jayapura regions, where physical alteration of wetland structure and introduction of invasive fishes is rapidly transforming such ecosystems.

Upland Bogs Upland bogs comprise small bodies of acidic open water on flat, elevated topography at elevations generally above 2,000 m in areas of high persistent rainfall (⬎300 cm year). The soil substratum beneath such bogs tends to be primarily organic, producing clear, cool (⬍16C) waters that are very low in dissolved minerals (con-

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Table 5.5.1. Endemic biota of major lakes in Papua Anggi Lakes 01 22⬘ S, 133 55⬘ E Insects Ischnura rhodosoma Lake Holmes 02 29⬘ S, 138 00⬘ E Fishes Chilatherina bleheri Melanotaenia maylandi (occurs in nearby creeks) Lake Sentani 02 42⬘ S, 140 30⬘ E Fishes Chilatherina sentaniensis Glossolepis incisus Glossogobius sp. Ayamaru Lakes 01 17⬘ S, 132 06⬘ E Fishes Melanotaenia ajamaruensis Melanotaenia boesmani Glossogobius hoesei Pseudomugil reticulatus Lake Kurumoi 02 10⬘ S, 134 5⬘ E Fishes Melanotaenia parva Lake Yamur 03 39⬘ S, 134 58⬘ E Fishes Variichthys jamoerensis Lake Laamora 03 41⬘ S, 134 17⬘ E Fishes Melanotaenia lakamora Mogurnda magna Lake Aiwaso 03 39⬘ S, 134 16⬘ E Fishes Melanotaenia lakamora Mogurnda aiwasoensis Mogurnda magna Lake Kamaka 03 43⬘ S, 134 11⬘ E Fishes Melanotaenia kamaka Melanotaenia pierucciae (occurs in nearby creeks) Craterocephalus fistularis Mogurnda pardalis (continued)

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Table 5.5.1. (Continued) Lake Mbutu 03 58⬘ S, 134 57⬘ E Fishes Pelangia mbutuensis Mogurnda mbuta Lake Kaifayama 03 53⬘ S, 134 46⬘ E Fishes Mogurnda kaifayama Paniai Lakes 03 55⬘ S, 136 20⬘ E Insects Ischnura ariel Archiboldargia mirifica Paniai Lake Crayfishes Charax boschmai Charax buitenijkae Charax communis Charax murido Charax pallidus Charax paniaicus Tage Lake 03 57⬘ S, 136 15⬘ E Fishes Oxelyotris wisselensis Tigi Lake 04 02⬘ S, 136 13⬘ E Fishes Oxelyotris wisselensis Crayfishes Charax communis Charax longipes Charax solus Note: The lakes listed contain distinctive assemblages of endemic fishes and invertebrates. Each lake represents a unique ecosystem, and all are priority conservation areas.

ductivity ⬍30 ␮mhos), stained yellow to brownish with humic solutes, and acidic (pH ⬍ 5.5). Such bogs are widely distributed in the Papuan uplands, and are distinguished by their strongly acidic water chemistry and impoverished invertebrate faunas, which include aquatic Diptera in the families Ephydridae and Dolichopodidae. Due to their elevation, these bogs uniformly lack fishes.

Upland Swamps and Marshes Upland marshes are perennial to seasonally intermittent, non-forested wetlands in upland areas (100–1,200 m) of moderate to high rainfall, but with better drainage than boglands. Their waters are clear and sometimes yellowish, with low to moderate dissolved mineral content (conductivity 30–80 ␮mhos), and circumneutral

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Figure 5.5.13. The diversity of juxtaposed lotic and lentic aquatic ecosystems in lowland New Guinea is well illustrated in this satellite image from the Kikori River basin of Papua New Guinea. Aquatic ecosystems visible in this image include lowland rivers and their tributary creeks, mixohaline mangrove swamps, freshwater swamps, and freshwater marshes. Photo: D. A. Polhemus.

(pH 5.5–7.5). Emergent aquatic plants (sedges and grasses) are often abundant, including Drosera, Gentiana, Utricularia, Brachyposium, Carex, and Scirpus. Their fauna is similar to that of bogs described above, but more abundant and diverse. Upland swamps are perennial to seasonally intermittent forested wetlands in upland areas (100–1,200 m) of moderate to high rainfall. Their waters are nonacidic, with characteristics similar to those of upland marshes. By comparison to marshes, however, their fauna is more diverse and often endemic, including a few fishes (often gudgeons of the genus Mogurnda), specialized Odonata in the family Coenagrionidae (Ischnura), and Heteroptera in the family Veliidae (Microvelia, Neusterensifer). Overall, upland marshes and swamps appear to be more productive than bogs, and support a greater faunal diversity. Such ecosystems tend to be localized and of relatively limited extent, forming either in natural depressions, or in areas where streams are impounded by either natural barriers or by the construction of road crossings.

Freshwater Lowland Swamps and Marshes These ecosystems comprise naturally occurring, shallow, standing, perennial limnetic waters in lowland areas at elevations ⬍ 100 m, which may occupy either definite or indistinct basins not immediately adjacent to the coastline, with emer-

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Figure 5.5.14. Lowland freshwater swamps, as seen here on Gam Island in the Raja Ampat group, intergrade along their seaward margins into mixohaline and saline mangrove swamps. Such ecosystems formerly covered extensive areas in Papua, but have been heavily degraded by logging. They are the favored habitat of certain fishes such as blue-eyes and gudgeons. Photo: D. A. Polhemus.

gent flora predominant. They are maintained by either stream inflow, or by exposure of the natural water table. Water quality in such systems may be variable, but salinity is always ⬍ 0.5 o/oo (conductivity 100–300 ␮mhos), with nearly neutral pH values of 6.0–7.5. Such freshwater lowland swamps and marshes represent a complex series of ecosystem types ranging from flooded taro and rice fields to natural marsh basins and riverine swamp forests. Included in this category are the vast lowland swamp forests of the Meervlakte, the huge palustrine basin along the central course of the Mamberamo River and its tributaries (Figure 5.5.12). Anthropogenically-altered ecosystems of this type are often favorable for the establishment and spread of invasive fishes, including snakeheads and tilapia (see discussion later in this chapter). Lowland freshwater marshes are perennial lowland wetlands lacking trees but with abundant emergent vegetation of other types. Natural systems of this type in Papua are exemplified by the extensive wetlands of the southern lowlands in the Digul and Trans-Fly regions; artificial systems are increasingly numerous and often agricultural, dominated by monocultural taro or rice. The characteristic fauna of both natural and artificial systems includes numerous fishes (especially blue-eyes, rainbows, glassfishes, and gudgeons), a diverse array of Odonata, partic-

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Figure 5.5.15. Mangrove swamps, such as this one on Batanta Island, are extensive along the coasts of New Guinea, and intergrade into freshwater swamps along their inland margins. They support a rich fish biota, and a distinct and diverse assemblage of aquatic insects, particularly surface-dwelling waterstriders. Photo: D. A. Polhemus photo.

ularly in the families Coenagrionidae (Agriocnemis, Pseudagrion) and Libellulidae (Agrionoptera, Neurothemis, Orthetrum, Rhyothemis, Tetrathemis), and various Heteroptera in the families Belostomatidae (Lethocerus), Nepidae (Ranatra), Gerridae (Limnogonus), Mesoveliidae (Mesovelia), and Veliidae (Microvelia). Lowland freshwater swamps, in contrast to marshes, are forested perennial lowland wetlands (Figure 5.5.14), with their water depth often fluctuating on a seasonal basis due to influxes of limnetic water from perennial streams. This ecosystem category includes a range of botanically diverse coastal plain and riparian forested wetlands, including sago swamps, pandanus swamps, and peat swamp forests, all of which are extensively represented in Papua, particularly along the southern coast bordering the Arafura Sea. Peat swamp forests could potentially be segregated as a separate ecosystem on the basis of acidic water chemistry, similar to the case with upland bogs. Distinguishing flora includes Metadina, Barringtonia, sago (Metroxylon sagu), various Pandanus species (for list see Stone, 1982), and Campnosperma brevipetiolata in very wet areas. Also assignable here are the ‘‘freshwater mangrove’’ forests, florally dominated by Myristica, Callophyllum, Syzygium, Campnosperma, Palaquium, Intsia and Diospyros, and similar riparian forests of Sonneratia caseolaris (for additional discussion see Johns 1982 and Chapter 5.7). This is prime habitat for certain fishes such as blue-eyes (Pseudomugil), and gudgeons (Oxyeleotris and Mogurnda). Typical insect fauna includes Odonata in the

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families Protoneuridae (Nososticta) and Libellulidae (Agrionoptera, Orthetrum), and Heteroptera in the families Gerridae (Limnometra, Rhagdotarsus), Veliidae (Microvelia, Strongylovelia), Hydrometridae (Hydrometra), Veliidae (Microvelia), Belostomatidae (Appasus), and Nepidae (Ranatra).

Saline Lowland Wetlands Lowland saline marshes, more commonly known as salt marshes, are non-forested lowland saline wetlands dominated by emergent vegetation, most characteristically Pickleweed (Batis maritima). Lowland saline swamps, by contrast, are forested lowland or riparian saline wetlands dominated by a diverse array of mangrove species (Figure 5.5.15), and often intergrade into true euhaline mangrove estuaries. Such mixohaline swamps support mixed floral assemblages of Avicennia, Nypa, Rhizophora, Bruguiera, and Sonneratia (for further discussion see Johnstone and Frodin, 1982), and a characteristic surface insect fauna (J. Polhemus and D. Polhemus 1996; Andersen 1992; Andersen and Weir 1999) of trepobatine Gerridae (Stenobates, Rheumatometroides) and haloveliine Veliidae (Xenobates). Salt marshes and mangrove swamps of these types are common along the coasts of Papua, with the latter extensively developed bordering the Arafura Sea, along the margins of Bintuni Bay, and at certain river mouths along the north coast. They are discussed more extensively in Chapter 5.4. Also falling within this ecosystem class are various intermittent lowland wetlands, consisting of lentic waters occurring seasonally in shallow basins. Their waters are generally warm (20–30C), mixohaline or poikilohaline (although evaporation may cause such waters to become hyperhaline as drying progresses), and basic, with pH values of 6.5–8.0. Characteristic biota includes certain littoral halophilic insects, including Diptera in the family Ephydridae (Ochthera) and Heteroptera in the family Saldidae (Pentacora, Saldula). Fishes may also be present, especially blue-eyes and various gudgeons. Examples of such ecosystems in Papua include certain seasonally dry lake basins formed between limestone anticlines east of Kaimana in the Vogelkop region, and salt pans that border the back margins of mangrove estuaries along the southern New Guinea coast.

Anchialine Pools An additional type of saline lowland wetland ecosystem found to a limited degree in Papua consists of anchialine pools. The name (from Greek anchialos, ‘‘near the sea’’) was suggested by Holthuis (1973) to define ‘‘pools with no surface connection to the sea, containing salt or brackish water, which fluctuates with the tides.’’ Such pools contain a distinctive biota consisting most typically of invertebrates of marine origin that have invaded through subterranean interstices, and often support unusual taxa not found elsewhere, particularly red shrimps, with fishes being rare or absent. These pools are generally small, with the majority being less than 100 m2 in area. Their surfaces are usually inland extensions of the oceanic water table, although mixohalinity, usually less than 10 percent, often results from dilution by seaward percolating groundwater. Ecosystems of this type, sometimes re-

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ferred to as ‘‘marine lakes’’ are known to occur on the Raja Ampat Islands of Mansuar and Misool. The pool on Mansuar is surrounded by mangrove and rainforest, has a soft silty bottom, a narrow band of algae and sponge along the perimeter. Five fish species are present in this ecosystem, including Kalyptatherina helodes, the only member of the family Telmatherinidae known outside of Sulawesi. This same fish species is also found in clear waters of sheltered mangrovecoral reef inlets throughout the Raja Ampat Islands. On the islands adjacent to the Papuan region (e.g., Timor), anchialine pools occur primarily in elevated fossil reef rock, and are variable in depth depending on tidal stage, with certain very shallow pools appearing only at high tide. Their surface waters are generally mixohaline, ranging from 1–10 o/oo, but occasionally approach euhaline levels, and are usually clear and circumneutral, with temperatures ranging from 22–30C. Waters within individual pools are usually homiohaline, but with sharp, stable, vertical salinity stratification evident in deeper pools. Although there are no direct surface connections with ocean, tidal fluctuations are also usually still evident, because the water surface level is merely an inland extension of the marine water table, with the mixohalinity resulting from dilution of intruding ocean water with seaward-percolating groundwater. Such pools may occur singly, but are more typically found in groups with subsurface interconnections, and thus represent surface manifestations of otherwise subterranean ecosystems. The biota of anchialine pools is unique and distinctive, with some faunal species, particularly red shrimps, not found elsewhere. Introduction of alien fishes quickly degrades or eliminates such crustacean communities, a process well documented elsewhere in the Pacific, particularly Hawai’i.

Subterranean Aquatic Ecosystems The anchialine pools discussed in the preceding section are localized surface exposures of larger and more extensive subterranean aquatic ecosystems that occur in all areas of tropical karst terrain. Such ecosystems may be lotic, such as the ‘‘Baliem Swallet,’’ where the Baliem River disappears underground for a considerable distance near Mt Trikora, or lentic, in the form of standing pools within cave systems. Despite the known existence of such ecosystems in Papua, their faunal exploration has been extremely limited. Cursory collections have been made from pools in a few accessible cave systems in the Baliem Valley, revealing that certain insect species occurring above the surface in surrounding areas, particularly the genus Microvelia in the Heteroptera, will colonize pools in the twilight zones of such caves. To date, no blind cave fishes, crustaceans, or insects have been recorded from Papua, although such taxa are known from adjacent Papua New Guinea (Allen 1996; Holthuis 1980; Chapter 5.13).

seaward interfaces The inland water ecosystems described above intergrade into marine systems at several points, the most important being saline marshes and swamps, and the

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estuaries that form at the seaward ends of the terminal reaches of perennial streams. Recent classifications of marine environments in the insular tropical Pacific address such estuarine ecosystems in a manner similar to that used above for freshwater systems. The estuarine transition zone between limnetic and euhaline waters is primarily one in which mixohaline waters in delineable basins exhibit continuous or periodic surface connection to the ocean, allowing the entry of a diverse euryhaline marine fauna, including certain snappers, glassfishes, cardinalfishes, damselfishes, gobies, and gudgeons (this definition excludes waters inhabited by stenohaline marine inshore fauna such as corals, urchins, etc.). The level of the water surface exhibits tidal fluctuations, which may also produce strong inflows and outflows, and there is generally a pronounced stratification of halinity (concentration of sodium chloride), temperature, and (usually) oxygen concentration. Two distinct subtypes of natural estuaries may be recognized based on freshwater inflows and diadromous fauna.

True Estuaries These are drowned river and stream mouths fed by limnetic water from perennial stream runoff. Their inland extent is determined by measurable tidal fluctuation and topography, such that estuaries of this type tend to be much more extensive on the south coast of Papua than on the north. In Papua such estuaries also tend to be horizontally stratified, with relatively large freshwater inflows relative to channel or basin volumes, leading to pronounced differences in salinity from head to mouth as one progresses along the length of the estuary. Additionally, such estuaries usually have a degree of vertical stratification, resulting from the fact that freshwater has a lower density than salt water, and therefore tends to ‘‘float’’ on top of a salt water wedge as the stream discharges into the ocean. The predominance of horizontal or vertical stratification is dependent on a variety of factors, including the volume and velocity of freshwater inflow, the depth and size of the estuary basin, and the degree of mixing created by winds and currents. In general, estuaries at the mouths large rivers tend to be homiohaline along any particular reach, while those of smaller stream systems tend to exhibit poikilohalinity resulting from wide seasonal fluctuations in freshwater discharge. Along many coasts in Papua true estuaries are often dominated by mangroves, and serve as important migratory pathways for larval and juvenile diadromous fishes and other animals (Erftemeijer et al 1989). They also support unique assemblages of marine water striders in the family Gerridae (Halobates, Rheumatometroides, Stenobates) which partition such estuaries on the basis of horizontal salinity gradients (Herring, 1961; J. Polhemus and D. Polhemus 1996; Andersen and Cheng 2004).

Estuarine Limnocrenes Estuaries of this type consist of nearshore basins with subterranean limnetic water sources (generally basal springs) and open connections to the sea. In contrast to true estuaries, their waters tend to be uniformly homiohaline, due to annually

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stable limnetic discharges. The biota is generally similar to that of true estuaries, but may also include submerged vascular plants, and lacks transient diadromous stream fauna.

Threats to Inland Water Ecosystems in Papua Although the overall condition of freshwater ecosystems in the New Guinea region is excellent, there are still obvious threats to the biota, which tend to manifest themselves on local rather than regional scales. These threats may be grouped into three general categories: physical alteration of habitat, utilization of biotic resources, and invasive species. Each of these threat categories is discussed separately below.

physical alteration of habitat Logging Large-scale industrial logging, particularly by international timber companies, is a clear threat to watershed integrity throughout the New Guinea region. The obvious and disastrous effects of clearcutting aside, even selective logging by such companies results in an extensive network of poorly-planned and constructed secondary roads that create widespread siltation and stream impoundment problems. Although treefalls are a natural element of the New Guinea rain forest and the small impoundments resulting from them are encountered on nearly every forest stream in the region, particularly in the lowlands, logging tends to greatly increase the number of such channel obstructions, increasing pool habitat and decreasing riffles. Logging roads also tend to employ rudimentary bridges that subsequently collapse, creating further impoundments. Opening the forest canopy also increases insolation (exposure to sunlight) and thereby increases water temperature. The overall effect, then, is to create a stream that is warmer, more slowly flowing, and traps more sediment. Much of the large-scale logging in Papua is undertaken by foreign companies with poor environmental records, or their local Indonesian subsidiaries. In addition to large-scale operations by companies such as PT Inhutani II and PT Astra, local military garrisons often set up illegal logging operations to subsidize their pay, usually with no consideration of environmental effects. By contrast, the advent of small-scale logging, utilizing ‘‘walkabout sawmills’’ appears to result in rather light and transient damage to streams and watersheds. Such operations leave a lighter environmental footprint because they usually target only particular tree species, such as rosewood, which are widely scattered in the forest; they do not operate in one area for a long period of time; and they do not require the creation of an extensive road network.

Shifting Cultivation The impacts of shifting cultivation are similar to those of clearcut logging, but on a far more localized scale. In traditional village settings, the effects of shifting

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cultivation were mitigated over time by the fact that such garden patches were relatively small in size and widely dispersed. In many cases, if all available garden areas had been used at least once, entire villages simply relocated to alternative sites, allowing the old gardens to go back to forest. As population has increased in many highland areas, however, the number of gardens has proliferated while the number of years they are allowed to lay fallow has decreased, and local governments have discouraged villages from changing location. In general, shifting cultivation tends to have disproportionate impacts on first order streams (the smallest streams in a given drainage network), which are characteristic of the ridge slopes on which gardens are usually established. Creeks passing through newly cleared garden areas are usually exposed to intense sunlight and high air temperatures, and obstructed by massive tangles of vines and tree branches that in many cases make them nearly impossible to traverse. These ecosystem impacts produce significant faunal changes, with deep forest species that require cooling shade, particularly certain genera of Odonata (Selysioneura, Tanymecosticta), being absent in such areas. Provided that a patchwork of forest and garden plots remains intact, however, such forest biota will eventually recolonize streams in former garden areas once a canopy of native trees is re-established.

Oil Palm In common with clearcut logging, oil palm plantations result in wholesale ecosystem conversion that has broad impacts across entire stream catchments. The creation of a plantation requires initial land clearing equivalent to clearcut logging (which may in fact be the first step if the proposed plantation area is covered with primary forest), after which a new canopy structure of oil palms eventually becomes established. Nutrient inputs from such plantations into adjacent streams appear to be high, probably due to fertilizer and other agrochemical runoff, leading to a proliferation of algae and consequent impacts on the benthic biota. Because oil palm development is generally undertaken on relatively flat lowland sites, it disproportionately impacts the terminal reaches of streams via clearance and drainage channelization of alluvial and swamp forests, with consequent impacts on diadromous biota similar to those described subsequently for mining.

Mining Large-scale mining operations have had obvious local impacts to certain river systems in Papua, most notably the Ajkwa, which lies downstream of the Freeport Grasberg gold and copper mine. Although large scale-mining produces dramatic local impacts that are highly visible, reasonable attempts to mitigate these impacts, which include siltation, chemical contamination, and catchment dewatering for pipeline slurries, have been undertaken at Grasberg. A more pernicious set of impacts often arises from small-scale gold mining efforts that are common throughout the New Guinea region. As noted by Susupu and Crispin (2001): ‘‘Environmental issues do not seem to be a strong concern for members of the small-

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scale artisanal mining community. Issues such as damage to river beds, solids in water and destruction of riverbanks are not addressed.’’ The most persistent impact to freshwater ecosystems from such small-scale mining arises not from physical disturbance to streambeds, however, but from the mercury used in the mining process. By way of example, some 4 tons of mercury per year is presently sold to alluvial miners in Papua New Guinea, based on wholesalers’ records (Susupu and Crispin 2001). This mercury is used to extract gold from black sand either in between the sluice-box compartments or via simple panning. In the Wau/Bulolo area, where dredge mining occurred from the late 1920s through the 1960s, bulldozers still occasionally uncover large puddles of mercury, and similar situations are reported from long-term mining sites on Bougainville (Susupu and Crispin 2001). In Papua, local mercury pollution is now also occurring in the Timika area due to illicit gold refining operations being conducted by military units using barrels of concentrate stolen from Grasberg mine. Being non-soluble, mercury remains in river sediments indefinitely, and may be difficult to detect, since it is possible for river water to flow clear of mercury even when high levels of mercury are present in the river bed. Such mercury contamination, however, frequently enters the riverine food chain, where it is amplified through successive trophic levels, eventually posing severe risks to local human populations who consume fish and crustaceans. In contrast to logging or oil palm plantations, which degrade entire catchments via wholesale landscape conversion, mining effluents generally impact only the main stem of any given catchment, leaving most tributaries undisturbed and available as potential reservoirs of biotic recolonization. The degradation of main stem rivers, however, particularly in the terminal reaches, can have serious impacts on certain diadromous faunal elements such as fish and prawns, preventing completion of the longitudinal migrations essential to their life cycles and thereby potentially extirpating them from certain river systems.

Petroleum Petroleum development has relatively limited impact on inland waters, because the environmental disturbances associated with it tend to be small, scattered, and highly localized. Outside of the possibility of spills and pipeline leaks, which can obviously have serious short-term local impacts, the major threat from petroleum development results from forest degradation or clearance adjacent to the network of service roads, which provide conduits into previously undisturbed tracts of forest. In general, due to the scattered nature of the operations and shifting well sites, the overall ecosystem impacts of petroleum development are in some aspects similar to those of selective logging or shifting cultivation. In addition, because petroleum operations are restricted to a only a few particular areas in Papua such as the Vogelkop Peninsula, they do not appear to pose a broad-scale threat to freshwater ecosystems in Indonesian New Guinea on the same order as logging or even mining.

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Dams Dams and hydropower developments are sparse in the New Guinea region, and their impacts on freshwater systems are currently minimal. Larger scale projects, such as the proposed large dam on the main stem of the Mamberamo River in northern Papua, would clearly have significant basin-wide impacts were they to be constructed, but such plans are currently shelved due to economic constraints. By contrast, small mini-hydros, which are commonly used in the mountains of New Guinea to provide electricity for local mission stations, have minimal biotic impact.

Ungulates The impacts of ungulates on New Guinea aquatic systems are underappreciated, but can be significant and extensive, in both upland and lowland areas. In the highlands, cattle grazing has been observed to create widespread slope terracing and converts valley bottoms into muddy marshes, increasing river siltation and water turbidity. Introduced Rusa Deer have similarly impacted savanna lowland habitats in southeastern Papua. Feral pigs, although widespread in New Guinea, have not had the same disastrous impacts to native forests as observed on smaller islands in Polynesia. Feral pigs are intensively hunted throughout the region, which probably serves to keep their numbers in check to some extent. It is unknown if they act as vectors of the water-borne disease leptospirosis, as they do in the Hawai’ian Islands, but this seems likely.

utilization of biotic resources Live Aquarium Fish Trade With a single exception, there appears to be little impact on the native fauna due to the live aquarium fish trade. As far as can be determined there is very little commercial harvesting of wild fishes for the aquarium trade with the exception of the illegal trade for Saratoga or Bony Tongue (Scleropages jardinii: family Osteoglossidae), which occurs in the southeastern border area of Papua. Saratoga is popular in the aquarium trade, probably because of its similar appearance to the Asian Arowana (S. formosus), which is a much sought-after ‘‘good-luck’’ fish in eastern Asia, particularly China and Japan, where they are known as Dragonfish. The huge popularity of the Dragonfish has apparently resulted in a demand for other species of bony tongues. Saratoga is a popular aquarium and sports-fish native to southern New Guinea and northern Australia. It breeds annually just prior to the wet season (September to November). After external fertilization the female orally incubates a brood of about 30–130 eggs until they hatch 1–2 weeks later (Allen et al. 2002). The female then guards the newly hatched young, which remain close to her mouth for the next 4–5 weeks. The young fingerlings are particularly vulnerable at this stage of the life cycle and are easily harvested. The species is protected by law in Indonesia, and subject to various regulations in Australia.

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Beginning in the 1990s villagers in the Torassi or Bensbach River area, in the Western Province of PNG immediately adjacent to the border with Papua, have been collecting and selling wild Saratoga fingerlings to merchants from across the border in nearby Merauke (Hitchcock, in press). These fish, as well as illegally captured fingerlings from Papua, are then exported to Asia, where they commanded considerable prices for several years. Australian fish breeders report that saturation of the market by Torassi Saratoga led to a collapse in prices and dramatic decline in demand for the species, which has negatively impacted upon the Australian export trade in wild-caught and captive-bred fingerlings. There is also evidence from local villagers living along the Bensbach River that seasonal harvesting of Saratoga over the past decade has resulted in a dramatic decline in population numbers. Therefore a critical need exists for more detailed study of this problem as well as a sound management plan that will insure the sustainability of the fishery. In addition, effective policing of the illegal trade is needed on the Papuan side of the border. There is scant information on the harvest of other ornamental species. Rainbowfishes of the family Melanotaeniidae are the only New Guinea group that is regularly seen in the international aquarium trade. Most of the species were introduced to the trade by various foreign collectors, often operating illegally. Rainbowfishes spawn readily in captivity and there is now a large captive breeding pool that apparently satisfies most of the commercial demand, thus negating the need for wild-caught fish. However, there is probably limited capture of wild fish by Indonesian merchants in places such as Sorong and Jayapura, although reliable data are lacking. At least one merchant was operating in Sorong as recently as six years ago. His trade revolved mainly around rainbowfishes, especially the brightly colored Boeseman’s Rainbow (Melanotaenia boesmani), which is endemic to the Ayamaru Lakes region of the central Vogelkop Peninsula. The species was introduced to the aquarium industry in 1983 by a German collector, and it has steadily increased in popularity. By 1989 Ayamaru villagers were catching so many live fish for the aquarium trade the species was on the brink of becoming endangered (Allen 1995). An estimated 60,000 male rainbows were captured each month for shipment to Jakarta exporters. Fortunately, the Indonesian government eventually placed controls on the industry.

Impact of Food Fish Harvesting on Native Fishes There are virtually no data on the harvest of native fishes for human consumption or the possible impact of this activity on native fishes in general. Compared to the considerable harvest of marine fishes, the take of freshwater fishes seems relatively insignificant. Nevertheless, people living along the major river systems depend on freshwater fishes for a significant portion of their diet. Most of the larger villages have regular fish markets, which appear to be dominated by forktail catfishes, large gudgeons (Eleotris and Oxyeleotris), and various introduced fish, especially carp and tilapia. Forktail catfishes (family Ariidae) are represented in New Guinea fresh

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waters by 21 species and are probably the most important food fish in this habitat. Although they are heavily targeted by gill netting and traditional fishing methods their numbers do not appear to be declining, at least in major Papuan river systems such as the Digul and Mamberamo. A variety of fishing methods are employed including hook and line from canoes, home-made traps, and various nets ranging from simple one-person hoop nets to large seines and gill nets. Streams, some of considerable size, are sometimes diverted and the former channel containing isolated pools with dense fish concentrations are then netted or speared. Some villages also employ Derris (Fabaceae) root to poison ponds, stagnant pools or slow flowing sections of creeks. In addition, local fishers are usually adept at catching by hand gudgeons and other fishes that hide in crevices. Traditional fishing methods appear to have insignificant impact on the native fish fauna. After all, they have been used for centuries and continue to be sustainable. The problem lies with more modern techniques, especially when outboard motors have been introduced in combination with gill nets. It is our opinion that gill nets should be banned from areas of special biological significance in Papua, such as Lake Sentani and Lake Yamur. Gill netting has certainly played a major role in the demise of the Freshwater Shark (Carcharinus leucas) in Lake Yamur and the Giant Sawfish (Pristis microdon) in Lake Sentani.

invasive species In relation to its overall size, the New Guinea region exhibits a remarkably low incidence of invasive freshwater species. This fortuitous situation appears to result from the fact that the region is lightly inhabited, has not experienced extensive colonization and settlement by foreign peoples (although this situation is changing in Indonesian New Guinea with a continuing influx of western Indonesian settlers that were initiated through now-defunct government-sponsored transmigration programs), and is still not well integrated into the global economy. The result is that freshwater ecosystems in many parts of the island and its proximal archipelagoes remain among the most pristine on earth. New Guinea’s general ecological integrity notwithstanding, the presence of exotic freshwater fishes is an increasing problem throughout the island. Allen (1991) reported the presence of 22 species representing 19 genera, 11 families, and all six continents. Since then at least six more introductions have been noted, and more can be expected, especially on the Indonesian side of the island. In the present chapter we provide details of the more recent introductions as well as a general overview of the invasive problem. Most of the introductions have had a negative impact, either by competing for space and limited food resources, or by feeding on native species, including their eggs and fry. Tilapia (Oreochromis mossambica) has adversely affected the environment, creating turbid conditions in formerly clean lakes, and badly overcrowding the indigenous fauna due to its prolific breeding. Several species including tilapia,

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walking catfish, carp, and climbing perch appear to be undergoing rapid population increases and therefore pose a serious threat to native fishes. The current distributional pattern of introduced fishes is closely tied to transmigration areas of Papua Province, particularly the larger population centers such as Jayapura and Timika. The transmigration program of the Indonesian government is no doubt responsible for many of the introductions. Newly arrived settlers often bring their pets and fish-pond stock from other parts of the archipelago. Thus, there is a major potential for further introductions. Of primary concern is the relatively recent appearance of four invasive species (tilapia, snakehead, climbing perch, and walking catfish) in the Bensbach River system of southwestern Papua New Guinea (Hitchcock 2002). At least some of these possibly entered the river via drainage ditches associated with the building of the Trans-Irian Highway, which in 1982 crossed the international border in two locations on the Upper Bensbach. Tilapia and walking catfish are more recent introductions, having been first noticed in the area in about 1995. Of equal concern is the appearance of two South American fishes, Prochilodus argenteus (Prochilodontidae) and Colossoma bidens (Characidae), and Barbonymus goniotus (Cyprindiae) from western Indonesia in the Ramu system of Papua New Guinea. The origin of these introductions remains a mystery, but they may have been species that were experimentally raised for potential introduction during an ill-conceived fish stock enhancement program sponsored by the Food and Agriculture Organization of the United Nations (FAO) in the 1980s. Allen et al. (2002) noted that the Mamberamo River in Papua Province had the highest percentage (17.1) of introduced fishes of any major river system in New Guinea. The appearance of species such as tilapia, walking catfish, snakehead, and three species of cyprinids is particularly alarming, given the relative isolation of this system and lack of major population centers. Another problem area is the Timika region of southern Papua Province. Prior to the opening of the Freeport gold and copper mine, there were no invasives in the region. But a huge influx of transmigrants has seen the introduction of tilapia, climbing perch, walking catfish, and snakehead (Allen et al 2000). In addition, the Blue Panchax (Aplocheilidae) from southeast Asia was introduced in the 1990s, apparently for mosquito control. Across New Guinea as a whole, invasive species appear more concentrated in lakes and wetlands, although certain lowland streams and river systems, particularly in the Mamberamo and Sepik-Ramu basins, are badly contaminated. The amazingly intact character of New Guinea’s wetland systems in a physical sense may in fact be limiting the spread of invasives, due to a lack of canals and periodically flooded agricultural field systems, coupled with natural seasonal drying. By contrast, the introductions of invasive fish into lotic (i.e., flowing water) environments is of great concern, since this enables highly vagile invasives such as tilapia, mosquitofish, or snakeheads to penetrate repeatedly both riverine and ephemeral riparian wetland habitats after seasonal flooding. Particularly problematic in this regard has been the introduction and continuing spread of snakeheads (Channa

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spp.) because of their ability to survive buried in the mud of ephemeral wetlands for months utilizing their accessory breathing organ (Courtenay and Williams 2004). This predaceous invasive has the potential to spread throughout the entire coastal wetland zone of southern New Guinea, from Etna Bay eastward to at least the Lakekamu River. Although the invasive fish species already present in New Guinea appear to be undergoing population expansions, thereby posing a grave threat to native species (Allen 1991), the specific impacts of such invasives on aquatic organisms endemic to New Guinea have for the most part not been determined. Similarly, little work has been undertaken regarding the identity or spread of other invasive freshwater animal species, particularly invertebrates. The following section provides additional detail on many of the most significant invasive fishes documented from the New Guinea region, and their varying degrees of ecological impact as known to date.

Carp Carp (Cyprinus carpio) are common in a few areas such as the upper Baliem River in Papua, Lake Kopiago, and the Lower and Middle Sepik and Ramu river systems of PNG (Allen 1991). Like many invasive fish species, carp modify their environment to conditions for which they are better suited to survive in than native fish species. World-wide, carp are regarded as a pest fish because of their tendency to uproot and destroy aquatic vegetation which results in increased turbidity and deterioration of habitats (Fuller et al. 1999). Carp have also been found to not only impact native fish species directly through egg predation, but also negatively impact waterfowl by increasing turbidity causing a reduction in food availability needed by both birds and native fish (Fuller et al. 1999).

Tilapia Tilapia (Oreochromis or Sarotherodon spp.) are perhaps one of the most adaptable and widespread species of fish in existence, and have been stocked throughout the world. These highly invasive fish are now abundant in the Timika region of Papua (Allen et al. 2000), and in the lower Ramu and middle and lower Sepik rivers of Papua New Guinea, and have become the most important food fish in the Sepik area (Allen 1991). Tilapia have ecological impacts similar to carp in that they uproot aquatic plants, are known to feed on wetland taro, and reduce food supplies for native bird species (Englund and Eldredge 2001). In contrast to carp, tilapia are even more invasive in tropical areas because their ability to withstand saline and brackish-water environments (Englund and Eldredge 2001) allows them to spread along a coastline.

Snakeheads Native to areas of Indonesia west of Weber’s Line, snakeheads (Channa striata) currently are found on Waigeo Island off western Papua (Allen et al. 2000); in streams near Bintuni on the Vogelkop Peninsula (Allen 1991); in the Timika re-

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gion (Allen et al. 2000); and in the vicinity of Merauke (Hitchcock 2002). Because migrants prefer eating this fish, it is commonly first found near their settlements (Allen et al. 2000) and would be expected to be spread throughout Papua by migrant communities. Species in the genus Channa are voracious and highly effective predators, and further establishment in New Guinea would have highly detrimental impacts to all freshwater biota. Snakeheads have been implicated in the extinction of at least four species of fish in Madagascar, displacing the formerly common native Cichlid genus Paratilapia from the central plateau and Lake Aloatra (Courtney et al. 2004; Courtenay and Williams 2004). In New Guinea, snakeheads appear to have caused a general reduction of native fish numbers and diversity in areas where they are present (Allen et al. 2000).

Trout Although Brown Trout (Salmo trutta) were introduced in 1949 to the highland regions of Papua New Guinea, and by 1952 had become established in this area (Werry 1998), there is no record of similar introduction into the upland streams of Indonesian New Guinea. In Papua New Guinea, the effect of trout on the native ichthyofauna appears to be minimal because they have survived in high elevation (⬎ 2,000 m) areas lacking native fishes (Allen 1991), although they have been documented to prey upon two species of endemic waterbugs, Nesocricos mion and Tanycricos acumentum, the latter of which also occurs in the central ranges of Papua (Polhemus and Polhemus 1985, 1986). The impacts of this predation are unknown, however, and these insect species still remain common enough after the introduction of trout to be used as bait by highland tribesman (Polhemus and Polhemus 1985).

Livebearers (Poeciliids) At least three poeciliid species have been recorded from New Guinea: Mosquitofish (Gambusia affinis), Guppies (Poecilia reticulata), and Green Swordtails (Xiphophorus helleri). Although Mosquitofish and the other two species of poeciliids were introduced into New Guinea to control mosquitoes, their impact in this regard has been minimal, and they instead crowd out effective native predators of mosquitoes such as rainbowfishes (Allen 1991). Given their continuing popularity as supposed biocontrol agents, however, such fishes will likely continue to be introduced to Papua, particularly in streams near larger towns. The situation in Papua New Guinea provides a cautionary example: guppies are now common in the Goldie River and in streams around Port Moresby, and are often the only fish present. In 2004, surveys funded by Conservation International also found guppies in the lower Gumini River system of Milne Bay Province; these guppies were the only invasive fish species found in this otherwise faunally pristine stream. Green Swordtails, although unknown from the main body of Papua province, are present in at least one upland stream on Biak Island, and are highly invasive, having the ability to penetrate streams far inland (Englund and Eldredge 2001). Invasive poeciliid fishes have been documented to cause the local extirpation of

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native aquatic invertebrates, such as species in the spectacular endemic Hawai’ian damselfly genus Megalagrion (Englund 1999). Additionally, international agency personnel need to be educated that native species such as rainbowfishes are often as good or better at controlling mosquito populations than introduced taxa. Any introduction of poeciliids for mosquito control is highly misguided because of potential non-target impacts to both native vertebrates and invertebrates, and because mosquito larvae, particularly those of the anophelenes that vector malaria, are generally found in puddle-type habitats and seasonally flooded pans on the floors of lowland forests, habitats not generally frequented by poeciliid fishes.

Walking Catfish Walking Catfish (Clarias batrachus) have increased their numbers around the Timika region since 1997, and are now common in some areas there (Allen et al. 2000). This species was first introduced into New Guinea in the Lake Sentani region and is now also found in the Vogelkop Peninsula (Allen 1991) as well as southeastern savanna areas near Merauke (Hitchcock, 2002). Similar to snakeheads, Walking Catfish have an accessory air-breathing organ allowing them to survive for long periods of time out of the water or in low-oxygen water. The impacts of this species on native biota are largely unknown (Fuller et al. 1999), but it does have excellent dispersal abilities once established because of its facultative air-breathing capabilities.

Blue Panchax The Blue Panchax (Aplocheilus panchax) is an ornamental aquarium species apparently being introduced for mosquito control, and is now rapidly spreading in peat swamps and disturbed areas in the Timika region (Allen 2000). Similar to Mosquitofish, this species can tolerate brackish water and may exert similar predation pressures on native invertebrates. However, impacts to native biota are currently unknown.

Climbing Perch or Climbing Gouramies Apparently first introduced into the western half of New Guinea, climbing perch (also called climbing gouramies) are now one of the most widely distributed invasive fish species on the island, being found from the Timika region of Indonesian New Guinea to the Morehead River area of Papua New Guinea (Allen 2000). This species is one of the hardiest of fishes (Fuller et al. 1999) because its accessory airbreathing organ allows it to survive for up to six days out of water (Allen et al. 2000). The use of its strong fin spines allows it to traverse considerable distances on land, and it is found in fresh to brackish water (Fuller et al. 1999). The impacts of this fish on native biota in New Guinea are unknown.

Gouramies Introduced primarily as a food fish, two families and three species of gourami are found in New Guinea. Two species in the family Belontiidae, the Snakeskin Gou-

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rami (Trichogaster pectoralis) and Threespot Gourami (Trichogaster trichopterus), have restricted ranges and do not appear to be highly invasive (Allen 1991). Werry (1998) states T. pectoralis is established in the southern lowland floodplain systems and certain areas around Port Moresby. The Giant Gourami (Family Osphronemidae), Osphronemus goramy, was introduced in 1957 but most introductions appear to have been unsuccessful and not in sufficient quantities to provide a fishery (Werry 1998). Impacts to native biota from gouramies are likely minimal because these species have not spread or even become successfully established except in a few instances.

Problems Caused by Invasive Species Many of the above invasive fish species have been documented to cause the extinction or severe range reduction of native fishes and invertebrates in other areas of the world, but their impacts in New Guinea are conjectural. Similarly, the introduced Cane Toad (Bufo marinus) is present in Papua, but its effects on native aquatic biota have not yet been determined. To date, no introduced species of freshwater insects or mollusks have been encountered in Papua, indicating that most if not all of the current invasives are fishes. Given that invasive species problems are still at an early stage in New Guinea, it is imperative to assess the current freshwater aquatic biodiversity of the island before such invasives spread further, in order to have a proper frame of reference from which to judge impacts. Of concern are continuing fish introductions promoted by the United Nations and its various agencies such as Food and Agriculture Organization to provide alternative protein sources and cash earning opportunities (Werry 1998). Werry (1998) documented that six species of fish were introduced to New Guinea between 1991 and 1997 through FAO programs. These introductions, and introductions of food fish by migrants, represent a continuing threat to the unique vertebrate and invertebrate biodiversity found in the varied inland water ecosystems of Papua, and New Guinea as a whole.

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894 / d a n a . polhemus & g erald r . allen Allen, G.R., K.G. Hortle, and S.J. Renyaan. 2000. Freshwater Fishes of the Timika Region, New Guinea. P.T. Freeport Indonesia, Timika. Allen, G.R., H. Ohee, P. Boli, R. Bawole, and M. Warpur. 2002. Fishes of the Yongsu and Dabra areas, Papua, Indonesia. In Richards, S.J., and S. Suryadi. (eds.) A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, D.C. Allen, G.R., and S.J. Renyaan. 2000. Fishes of the Wapoga River System, northwestern Irian Jaya, Indonesia. Pp. 50–58, 148–155 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 9, Conservation International, Washington, D.C. Andersen, N.M. 1975. The Limnogonus and Neogerris of the Old World. Entomologica Scandinavica Supplement 7: 1–96. Andersen, N.M. 1992. A new genus of marine water striders (Hemiptera, Veliidae) with five new species from Malesia. Entomologica Scandinavica 22: 389–404. Andersen, N.M., and L. Cheng. 2004. The marine insect Halobates (Heteroptera: Gerridae) biology, adaptations, distribution and phylogeny. Oceanography and Marine Biology: An Annual Review 42: 119–180. Andersen, N.M., and T.A. Weir. 1999. The marine Haloveliinae (Hemiptera: Veliidae) of Australia, New Caledonia and southern New Guinea. Invertebrate Taxonomy 13: 309–350. Baehr, M. 1990. Revision of the genus Ochterus Latreille in the Australian region (Heteroptera: Ochteridae). Entomologica Scandinavica 20: 449–477. Balke, M. 1995. The Hydroporini (Coleoptera: Dytiscidae: Hydroporinae) of New Guinea: systematics, distribution and origin of the fauna. Invertebrate Taxonomy 9: 1009–1019. Balke, M. 1999. Two new species of the genus Copelatus Erichson, 1832, subgenus Papuadytes Balke, 1998, from Papua New Guinea (Insecta: Coleoptera: Dytiscidae). Annalen des Naturhistorischen Museum Wien 101 B: 273–276. Balke, M. 2001. Biogeography and classification of New Guinea Colymbetini (Coleoptera: Dytiscidae). Invertebrate Taxonomy 15 (2): 259–275. Balke, M., and L. Hendrich. 1992a. Ein neuer Rhantus Dejean aus West-Neuguinea (Coleoptera: Dytiscidae). Entomologische Zeitschrift 102 (3): 37–39. Balke, M., and L. Hendrich. 1992b. Ein neuer Schwimmka¨fer der Gattung Hydaticus Leach (Coleoptera: Dytiscidae) aus dem Hochland von West Papua. Mitteilungen der schweizerischen entomologische Gesellschaft 65: 297–302. Balke, M., L. Hendrich, and G. Wewalka. 1992. Carabdytes upin n.gen., n.sp. aus Neuguinea (Coleoptera: Dytiscidae). Entomologische Zeitschrift 102 (6): 93–100. Balke, M., D.J. Larson, and L. Hendrich. 1997. A review of the New Guinea species of Laccophilus with notes on regional melanism (Coleoptera: Dytiscidae). Tropical Zoology 10 (2): 295–320. Balke, M., D. Larson, L. Hendrich, and E. Konyorah. 2000. A revision of the New Guinea water beetle genus Philaccolilus Guignot, stat.n. (Coleoptera:Dytiscidae). Deutsche Entomologische Zeitschrift 47 (1): 29–50. Bistro¨m, O., M. Balke, and L. Hendrich. 1993. A new species of Hyphydrus Illiger 1802 (Coleoptera: Dytiscidae) from West New Guinea, and notes on other species of the genus. Tropical Zoology 6: 287–298. Bott, R. 1974. Die su¨sswasserdrabben von Neu Guinea. Zool. Verh. 136: 1–36. Brinck, P. 1976. The Gyrinidae of the Bismarck Archipelago and the Solomon Islands (Coleoptera: Gyrinidae). Entomologica Scandivanica 7: 81–90.

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Inland Water Ecosystems: Classification, Biota, and Threats / 895 Brinck, P. 1981. Spinosodineutes (Coleoptera: Gyrinidae) in New Guinea and adjacent islands. Entomologica Scandinavica Supplement 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. Brooks, G.T. 1951. A revision of the genus Anisops (Notonectidae, Hemiptera). University of Kansas Science Bulletin 34, pt. 1 (8): 301–519. Courtenay, W.R, Jr., and J.D. Williams. 2004. Snakeheads (Pisces, Channidae)—a biological synopsis and risk assessment. US Geological Survey Circular 1251: 1–143. Courtenay, W.R., Jr., J.D. Williams, R. Britz, M.N. Yamamoto, P.V. Loiselle. 2004. Identity of introduced snakeheads (Pisces, Channidae) in Hawai‘i and Madagascar, with comments on ecological concerns. Bishop Museum Occasional Papers 77: 1–13. Demoulin, G. 1954. Contribution a` l’e´tude des Palingeniidae (Insecta, Ephemeroptera). Nova Guinea, Zoology 33: 305–344. 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. Royal Entomological Society of London, London. Englund, R.A. 1999. The impacts of introduced poeciliid fish and Odonata on endemic Megalagrion (Odonata) damselflies on Oahu Island. Hawaii. Journal of Insect Conservation 3: 225–243. Englund, R.A., and L.G. Eldredge. 2001. Fishes. Pp. 32–40 in Staples, G.W., and R.H. Cowie (eds.) Hawaii’s Invasive Species, A Guide to Invasive Plants and Animals in the Hawaiian Islands. Mutual Publishing and Bishop Museum Press, Honolulu. Erftemeijer, P.L.A., and G.R. Allen. 1989. Bird observations at Danau Kurumoi, Irian Jaya. Kukila 4 (3/4): 153–155. Erftemeijer, P., G. Allen, and Zuwendra. 1989. Preliminary resource inventory of Bintuni Bay and recommendations for conservation and management. PHPA/Asian Wetlands Bureau-Bogor Indonesia: 1–151. Fuller, P.L., L.G. Nico, and J.D. Williams. 1999. Nonindigenous fishes introduced into inland waters of the United States. American Fisheries Society, Bethesda, Maryland. Gentili, E. 1980. The genus Laccobius in Melanesia (Coleoptera: Hydrophilidae). Pacific Insects 22 (3–4): 385–400. Gentili, E. 1981. I Laccobius di Nuova Guinea conservati al Museo ‘‘G. Doria’’ di Genova. Ann. Museo Civico di Storia Naturale Genova 83: 271–274. Gentili, E. 1988. Verso una revisione del genere Laccobius (Coleoptera, Hydrophilidae). Annuario Oss. Fis. Terr. Mus. Stoppani Semin. Arc. Milano 9 (n.s.) (1986): 31–47. Gentili, E. 1989. Alcune novita´ sul genere Laccobius (Coleoptera, Hydrophilidae). Annuario Oss. Fis. Terr. Mus. Stoppani Semin. Arc. Milano 10 (n.s.) (1987): 32–39. Grant, P.M. 1985. Systematic revision of the Thraulus group genera (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) from the Eastern Hemisphere. PhD diss., Florida State University. Haines, A. 2001. Freshwater Snails of the Tropical Pacfic Islands. Institute of Applied Sciences, Suva. Herring, J.L. 1961. The genus Halobates (Hemiptera: Gerridae). Pacific Insects 3 (2–3): 223–305. Hitchcock, G. 2002. Fish fauna of the Bensbach River, southwest Papua New Guinea. Memoirs of the Queensland Museum 48 (1): 119–122.

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896 / d a n a . polhemus & g erald r . allen Hitchcock, G. In press. Cross-border trade in Saratoga fingerlings from the Torassi or Bensbach River, southwest Papua New Guinea. Traffic Bulletin of the IUCN. 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. 1973. Caridian shrimps found in land-locked salt water pools at four IndoWest Pacific localities (Sinai Peninsula, Funafuti Atoll, Maui and Hawaii islands), with description of one new genus and four new species. Zoologische Verhandelingen 128: 1–48. Holthuis, L.B. 1980. A new cavernicolous freshwater crab from New Guinea (Crustacea: Decapoda). Zoologische Meddelelser 55 (27): 313–320. 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. Hungerford, H.H., and R. Matsuda. 1958. The Tenagogonus-Limnometra complex of the Gerridae. University of Kansas Science Bulletin 39: 371–457. IUCN. 1991. A Directory of Asian Wetlands. IUCN Conservation Monitoring Centre, Cambridge. Johns, R.J. 1982. Plant zonation. Pp. 309–330 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr. W. Junk, The Hague. Johnstone, I.M. and D.G. Frodin. 1982. Mangroves of the Papuan subregion. Pp. 513–528 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr. W. Junk, The Hague. Kormilev, N.A. 1971. Ochteridae from the Oriental and Australian regions (HemipteraHeteroptera). Pacific Insects 13: 429–444. Lansbury, I. 1962. Notes on the genus Anisops in the Bishop Museum (Hem.: Notonectidae). Pacific Insects 4 (1): 141–151. Lansbury, I. 1963. Notes on water-bugs of Solomon Islands and New Hebrides. Pacific Insects 5 (1): 5–10. Lansbury, I. 1965. A new tribe and genus of Notonectidae (Heteroptera: Notonectidae) from Borneo. Pacific Insects 7 (2): 327–332. Lansbury, I. 1966. Notes on the genus Aphelonecta (Hemiptera-Heteroptera: Notonectidae). Pacific Insects 8 (3): 629–632. Lansbury, I. 1968a. The Enithares (Hemiptera-Heteroptera: Notonectidae) of the Oriental Region. Pacific Insects 10 (2): 353–442. Lansbury, I. 1968b. A revision of the genus Paranisops (Heteroptera: Notonectidae). Proceedings of the Royal Entomological Society of London (B) 33 (11–12): 181–188. Lansbury, I. 1968c. 50. Notonectidae (Hemiptera-Heteroptera) of Rennell Island. The Natural History of Rennell Island, British Solomon Islands 5: 95–98. Lansbury, I. 1969. The genus Anisops in Australia (Hemiptera-Heteroptera, Notonectidae). Journal of Natural History 3: 433–458. 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.

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Inland Water Ecosystems: Classification, Biota, and Threats / 897 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. 1974. Notes on the genus Enithares Spinola (Hem., Notonectidae). Entomologist’s Monthly Magazine 109: 226–231. Lansbury, I. 1975. Notes on additions, changes and the distribution of the Australian water-bug fauna (Hemiptera-Heteroptera). Memoirs of the Natural History Museum of Victoria 36: 17–24. 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.

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898 / d a n a . polhemus & g erald r . allen 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. 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. Neboiss, A. 1986a. Atlas of Trichoptera of the SW Pacific–Australian Region. Dr. W. Junk Publishers, The Hague. Neboiss, A. 1986b. Taxonomic changes in caddis-fly species from the South-West Pacific–Australian region with descriptions of new species (Insecta: Trichoptera). Memoirs of the Museum of Victoria 47: 213–223. Neboiss, A. 1986c. Identity of two caddis-fly species described by Brauer and Nava´s (Trichoptera). Aquatic Insects 8: 99–104. Neboiss, A. 1987. Preliminary comparison of New Guinea Trichoptera with the faunas of Sulawesi and Cape York Peninsula. Pp. 103–108 in Bournaud, M., and H. Tachet (eds.) Proceedings of the 5th International Symposium on Trichoptera. Dr. W. Junk Publishers, Dordrecht. Neboiss, A. 1989. The Ocecetis reticulata species-group from the South-West Pacific (Trichoptera: Leptoceridae). Bijdragen tot de Dierkunde 59 (4): 191–202. Neboiss, A. 1994. A review of the genus Paranyctiophylax Tsuda from Sulawesi, Papua New Guinea and northern Australia (Trichoptera: Polycentropodidae). Memoirs of the Museum of Victoria 54: 191–205. ¨ ber papuanische Gyriniden. Senckenbergiana 7 (5): 172–177. Ochs, G. 1925. U Ochs, G. 1955. Die Gyriniden-fauna von Neuguinea nach dem derzeitigen stand unserer kenntnisse (Coleoptera: Gyrinidae). Nova Guinea, New Series 6 (1): 87–154. Ochs, G. 1960. Nachtrag zur Gyriniden-fauna von Neuguinea. Opuscula Zoologica 53: 1–4. Petocz, R. G. 1989. Conservation and Development in Irian Jaya. A Strategy for Rational Resource Utilization. E. J. Brill, Leiden. 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. 1994. Conservation of aquatic insects: worldwide crisis or localized threats? American Zoologist 33: 588–598. Polhemus, D.A. 1998. Aquatic insects of the Wapoga River area, Irian Jaya, Indonesia. Pp. 39–42 and 90–94 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. Conservation International, Washington, D.C. Polhemus, D.A. 2002. Aquatic insects of the Dabra area, Mamberamo River basin, Papua, Indonesia. Pp. 57–60 and 129–137 in Richards, S.L., and S. Suryadi (eds.) A Biodiversity Assessment of the Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, D.C.

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Inland Water Ecosystems: Classification, Biota, and Threats / 899 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. 1989a. 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. 1989b. The Aphelocheirinae of tropical Asia (Heteroptera: Naucoridae). Raffles Bulletin of Zoology 36: 167–310. 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. 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, J.T., and I. Lansbury. 1997. Revision of the genus Hydrometra Latrielle in Australia, Melanesia, and the South-west Pacific (Heteroptera: Hydrometridae). Bishop Museum Occasional Papers 47: 1–67. 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. 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

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900 / d a n a . polhemus & g erald r . allen 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. Tijdschrift 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 & 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. Stone, B.C. 1982. New Guinea Pandanaceae: first approach to ecology and phytogeography. Pp. 401–436 in Gressitt, J.L. (ed.). Biogeography and Ecology of New Guinea. Dr. W. Junk, The Hague. Susapu, B., and G. Crispin. 2001. Report on small-scale mining in Papua New Guinea. Mining, Minerals and Sustainable Development Project, International Institute for Environment and Development, (IIED), Rep. 81. 29 pp. 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. Wells, A. 1990. The hydroptilid tribe Stactobiini (Trichoptera: Hydroptilidae) in New Guinea. Invertebrate Taxonomy 3: 817–849. Wells, A. 1991. The hydroptilid tribes Hydroptilini and Orthotrichiini in New Guinea (Trichoptera: Hydroptilidae: Hydroptilini). Invertebrate Taxonomy 3: 487–526. Werry, Lloyd P. 1998. A review of freshwater fish introductions in Papua New Guinea. Science in New Guinea 24: 33–36.

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5.6. Coastal Vegetation of Papua robert j. johns, garry a. shea, and pratito puradyatmika h e ve g e t at i o n of the beach ridges and flats ranges from pioneer herbaceous communities on the first beach ridge to tall mixed forest inland. Papua has the most extensive area in Indonesia with a total of 3,819 km2 of coastal and beach vegetation. This represents 65.2% of the total for Indonesia. The coastal vegetation of Papua is dominated by herbaceous communities, shrubberies, and littoral forest. The transition between these is usually abrupt, but elements of all formations are sometimes intermingled. Communities in the tidal swamps have a close physiographic relationship to other strand communities. In addition to the land component, there is a littoral component composed of a number of seagrass communities. Coastal beach ridges and swales (linear swampy depressions) are located along exposed seashores and on old beach ridges within the estuaries. The stripped pattern of the ridges and flats is hardly apparent on the ground, but shows up clearly from the air because of differences in height, composition, and color of the forest. The terrain is usually traversed by more or less permanently swampy depressions or swales that are aligned parallel to the ridges. The vegetation in the swales ranges from floating and submerged plants in their deepest parts, to swamp grasses, reeds, sago palm, pandans, and swamp woodland in progressively shallower water. Mangroves are present in brackish swales within the tidal zone.

T

Littoral Communities The seagrass flora of New Guinea is rich in species, with 10–13 species (Johnstone 1978a,b, 1982; Brouns and Heijs 1985; Heijs and Brouns 1986; Chapter 5.3). The seagrass beds are usually of a mixed nature, consisting of six to seven species. In these multi-species meadows Thalassia hemprichii is the dominant seagrass, followed by Enhalus acoroides and Cymodocea rotundata. Cymodocea serrulata, Halophila ovalis, Halodule uninervis, and Syringodium isoetifolium may be locally abundant. The seagrass communities have been described for New Guinea in Johnstone (1982) and Chapter 5.3. Seagrasses provide grazing for dugongs, sea turtles, certain species of fish, and some species of sea urchins that possess cellulose-digesting bacteria. Productivity in seagrass beds is high (Whitten et al. 1987, cited in MacKinnon et al. 1996) but only 5% of seagrass production is consumed directly; the rest enters the offshore food chain as decomposing material. Net primary productivity in Thalassodendron ciliatum meadows off Sulawesi was measured at 16.4 tons/ha, higher than the rate Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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for many lowland forests. Many adult and juvenile stages of fish and invertebrates spend parts of their life cycles in seagrass beds, feeding or taking shelter. Important commercial and subsistence species include rabbitfish, goatfish, and mullets (Polunin 1983; Burbridge and Maragos 1985), and edible invertebrates such as crabs, shrimps, clams, and sea cucumbers. Other common invertebrates include sea stars, gastropods, and sometimes corals. Some of the common seaweeds in the beds are also edible. Because seagrass beds are in shallow, nearshore, and often accessible areas, they are popular fishing grounds in Indonesia (MacKinnon et al. 1996). Seagrass beds serve several other useful functions, including stabilization of offshore sand reservoirs, and regular transport of carbonate sand to beaches under dynamic systems nearer to the shore. The beds are a source of commercially valuable algae and may be particularly favorable for seaweed mariculture (Burbridge and Maragos 1985). As coastal ecosystems in shallow, well-lit waters, seagrass beds are susceptible to damage from increased sedimentation levels, dredging, and thermal and chemical pollution (Zieman 1975) as well as overexploitation.

Herbaceous Beach Vegetation The littoral pioneers that dominate the herbaceous beach vegetation are deeprooting and tolerant of salt, wind, and high soil temperatures. Most species have floating seeds, and are consequently widely distributed throughout the Malesian region and northern Australia (Rand and Brass 1940). They are usually dominated by the creeper Ipomea pes-capraea, particularly on the seaward slopes. This is often mixed with other creepers, Canavallia maritime and Vigna marina. Coastal grasses and sedges are more common on the crests of the sand dunes and back dunes, including the species Ischaemum muticum, Fimbristylis, and Remirea maritima (Cyperus pedunculatus). Van Balgooy (1996) describes the situation in the Aru Islands where the I. pes-capraea formation merges into coastal forest. Sesuvium portulacestrum is a tasteless pink-flowered vegetable growing along the coast. In the Merauke area a low shrubby vegetation dominated by Wedelia and Stachytarpheta grows on the dunes. These littoral pioneer species are tolerant of salt, wind, and high soil temperatures. The parasite Cassytha filiformis (Lauraceae) grows on many coastal plants, and is also a common herb in eucalypt savanna. Grasses and sedges such as Ischaemum muticum, Fimbristylus spp., and Cyperus pedunculatus are more prominent on the crest. Commonly the lauraceous parasite Cassytha filiformis overgrows a variety of host plants. Seedlings of common beach trees and shrubs often grow among these grasses and sedges. The mixed species community may be named after the dominant species (e.g., Ischaemum muticumFimbristylus sericea-Cyperus). Single species often dominate the ridge crests, such as Ischaemum muticum, Fimbristylus sericea, Spinifex littoreus, and Cyperus spp.

Scrub Communities Behind the herbaceous beach vegetation is a transition zone where shrubs and low trees are scattered among the herbaceous vegetation. The dominant species in the

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scrub community are Pemphis acidula, Messerschmidia (Tournefortia), Scaevola, and Allophylus cobbe, associated with various herbaceous and woody creepers and climbers. In some places, this may give way to a dense scrub, composed of Hibiscus tiliaceus, Desmodium umbellatum, and woody climbers such as Flagellaria indica. Hibiscus tiliaceus can dominate this community.

Coastal Shrub and Tree Communities Sometimes a shrubby beach community occurs which is very similar to the Barringtonia formation, but lacking the larger trees. Common species include Clerodendron inerme, Morinda citrifolia, and Wedelia biflora. Pemphis acidula is a dwarf shrub on coral rocks or can be a larger shrub on non-coralline rocky shores. Cassytha filiformis is a common parasite on many coastal plants. Cycas rumphii stems, often with branched trunks, are restricted to coral headlands and igneous rocks. The beach forests of New Guinea are similar to surrounding areas in Malesia and Northern Australia. The structure and species composition of beach forest varies from area to area. A number of communities can be identified according which species are actually present and dominant on a particular site, such as Barringtonia asiatica, Calophyllum inophyllum, Terminalia catappa, Barringtonia asiatica-Calophyllum inophyllum. In the Aru Islands (van Balgooy 1996) the Barringtonia formation occurs on both sandy and rocky coasts. Patches of Barringtonia asiatica are not common in the Merauke area. Mixed beach or littoral forests often develop on inland beach ridges and flats. Pterocarpus indicus, Terminalia spp., and Syzygium spp. are often dominant species, or Barringtonia asiatica-Calophyllum inophyllum can dominate. Many of the species listed above for the beach woodland communities are also present. Mixed beach forest is characterized by abundant palms in the shrub and lower tree layers, and by many trees with poor stem form. Climbers are usually plentiful and in many places tree trunks are covered by the climbing fern Stenochlaena. Substrate and the influence of salt spray prevents these communities from developing into a climatic climax. Thus, the communities are referred to here as edaphic climax communities.

Mixed Woodland or Open Forest Communities Characteristic species of these communities are: Ardisia elliptica, Caesalpina bonduc, Calophyllum inophyllum, Casuarina equisetifolia, Cocos nucifera, Colubrina asiatica, Crinum asiaticum, Cycas rumphii, Desmodium umbellatum, Dodonaea viscosa, Erythrina variegata, Guettarda speciosa, Heritiera littoralis, Hernandia nymphaeifolia, Hisbiscus tiliaceus, Mammea odoratus, Maranthes corymbosa, Messerschmidia argentea, Morinda citrifolia, Pandanus bibur, Pandanus tectorius, Pluchea indica, Pongamia pinnata, Premna corymbosa, Scaevola taccada, Sophora tomentosa, Tacca leontopetaloides, Terminalia catappa, Thespesia populnea, and Wedelia biflora.

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Calophyllum inophyllum, Terminalia catappa, Thespesia populnea, and Hernandia peltata can form a narrow strip of coastal vegetation. Other common genera are Cerbera and Aegiceras. Small subcanopy shrubs and trees include Vitex trifolia, Hibiscus tiliaceous, Scaevola, and Pandanus odoratisimus. Other common species are Myristica hollrungii, Diospyros spp., Pandanus tectorius, and P. pedunculatis. Finschia chloroxantha can also be locally common. In some areas the understory is dominated by Wedelia biflora, Crinum asiaticum, and Wedelia biflora. On uplifted coral Pandanus dubius is a common species. On Waigeo Island, coastal forest is dominated by Intsia bijuga, Pongamia pinnata, Homalium sp., Ficus prasinicarpa, Glochidion, Syzgium, and Timonius. On flatter coastal areas Hibiscus tiliaceous and Scaevola sericea were common. On many smaller limestone islands were almost pure stands of Vandopsis lissochiloides, Paphiopedilum praestans, Ficus prasinicarpa, Glochidion, and Pongamia pinnata. Small lateritic islands were covered with dense stands of Dicranopteris linearis.

Casuarina Forest Casuarina equisetifolia, a common pioneer on beach ridges and flats, often forms dense stands above the high-water mark on sandy coasts, river mouths, and offshore bars, usually behind a narrow zone of herbaceous beach vegetation where the beach is not very steep. These forests are characterized by their grayish-green foliage. The southwest coast of Papua, around Kokenau, is characterized by long lines of tall casuarinas—the coast is referred to as the ‘‘Casuarina Coast.’’ The coastal strip of the Mamberamo is bounded by a long row of Casuarina equisetifolia. Casuarina forest also occurs to the west and east of the mouth of the Mamberamo River. On both sandy and rocky coasts on the Aru Islands (van Balgooy 1996), the vegetation is sometimes dominated by Casuarina equisetifolia. Casuarina is unable to regenerate on the litter carpet of its own dead, fallen photosynthetic twigs. Thus, Casuarina equisetifolia is gradually replaced by various broad-leaved trees, which form a mixed forest or woodland. Some very large, emergent Casuarina trees may persist as remnants of the former forest. The main canopy and understory are composed of the species that are common in the mixed woodland beach communities. A good example occurs about 1 km west from the mouth of the Unurupa River on the seaward facing coast. This was selected for quantitative study. The dominant species are Casuarina equisetifolia, Intsia spp., Sterculia spp., and Heritiera littoralis. The Casuarina occurs as an overmature emergent. Hibiscus tiliaceus, Barringtonia asiatica, and Calophyllum inophyllum are common along the beach margin. Heritiera littoralis is typically an element of the adjacent mangrove community that occurs behind the beach ridge. One pioneer Casuarina community occurs on a sand island or spit near the mouth of the estuary north west of Puriri Island. Casuarina equisetifolia is the dominant species, and occurs mainly as a single species community with some Hibiscus tiliaceus along the beach fringe. The estimated total basal area of trees

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over 10 cm dbh is 10.1 m2/ha. The estimated total stem volume is 21.8 m3/ha. The vertical and horizontal structure typical of this community is illustrated in the profile diagram in Figure 5.6.1. A survey of Casuarina-Mixed Forest was carried out by Shea et al. (1996) for the vegetation on a beach ridge about 1 km west of the mouth of the Unurupa River on the seaward facing coast. The most important genera and species based on a combined ‘‘importance value’’ (Curtis 1959; see Appendix 5.6.1) are Lumnitzera littorea, Heritiera littoralis, Hibiscus tiliaceus, Sterculia sp., Sonneratia alba, Heritiera littoralis, Terminalia catappa, and Intsia sp. The Casuarina occurs as an overmature emergent. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 27 m2/ha. The estimated total stem volume is 181.5 m3/ha. The vertical and horizontal structure typical of this community is illustrated in the profile diagram in Figure 5.6.2. Species in this plot were Sterculia sp., Intsia acuminata, Manilkara sp., Intsia sp., Sonneratia alba, Terminalia sp., Lumnitzera sp., Hibiscus sp., Casuarina equisetifolia, Diospyros sp., Heritiera sp., and Osbornia sp. The mangrove species Lumnitzera, Sonneratia, and Heritiera occupy the inner margins of beach ridges, but may also be found as large relic trees on the seaward side of beaches (e.g., along the beach south of Pasir Hitam).

Cocos nucifera Vegetation At the end of the 19th and at the beginning of the 20th century coconuts were planted almost everywhere, including in plantations, for the production of copra. A quantitative analysis was carried out for the vegetation on a beach ridge adjacent

Figure 5.6.1. Pioneer Casuarina equisetifolia-Hibiscus tilaceus forest on sand island in Minajerwi estuary, southern Papua. A. Casuarina equisetifolia; B. Hibiscus tilaceus. Source: Shea et al. (1996).

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Figure 5.6.2. Profile diagram of mixed beach forest community. Casuarina equisetifolia over Lumnitzera littorea-Sonneratia alba-Adenanthera-Sterculia. A. Sterculia sp.; B. Intsia acuminata; C. Manilkara sp.; D. Intsia sp.; E. Sonneratia alba; F. Terminalia sp.; G. Lumnitzera sp.; H. Hibiscus sp.; I. Casuarina equisetifolia; J. Diospyros sp.; K. Heritiera sp.; L. Osbornia sp. Source: Shea et al. (1996).

to the abandoned village at Pasir Hitam in the Minajerwi Estuary. The dominant species are Cocos nucifera, Casuarina equisetifolia, Terminalia catappa, and Hibiscus tiliaceus. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 28.3 m2/ha. The estimated total clear bole volume is 155.7 m3/ha (Shea et al. 1996). The vertical and horizontal structure typical of this community is illustrated in the profile diagram in Figure 5.6.3. Trees recorded at this site were Cocos nucifera, Hibiscus tiliaceus, Casuarina equisetifolia, Barringtonia, and Terminalia catappa. Around Merauke are extensive areas dominated by Cocos nucifera. This could be explained either by the gradual seaward movement of the coastline or, alternatively, due to planting. Under the extensive stands of coconuts in the Merauke

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Coastal Vegetation of Papua / 907

Figure 5.6.3. Beach dominated by Cocos nucifera-Casuarina equisetifolia-Terminalia catappa-Hibiscus tiliaceus at Pasir Hitam, southern Papua. A. Cocos nucifera; B. Hibiscus tiliaceus; C. Casuarina equisetifolia; D. Barringtonia sp.; E. Terminalia catappa. Source: Shea et al. (1996).

area, there is often a rich ground flora which includes Crinum, Cenchrus, Fimbristylis, Setaria and Euphorbia.

Acknowledgments We wish to express our sincere thanks to Dr W. Vink for his constructive comments and to the editors for their advice and assistance.

Literature Cited Brouns, J.J.W.M., and F.M.L. Heijs. 1985. Tropical seagrass ecosystems in Papua New Guinea. A general account of the environment, marine flora and fauna. Proc. K. Ned. Akad. Wetensch. C88: 145–182. Burbridge, F., and J.E. Maragos. 1985. Coastal Resource Management and Environmental Assessment Needs for Aquatic Resources Development in Indonesia. Washington Institute for Environment and Development, Washington, D.C. Heijs, F.M.L., and J.J.W.M. Brouns. 1986. A survey of seagrass communities around the Bismarck Sea, Papua New Guinea. Proc. K. Ned. Akad. Wetensch. C89: 11–44.

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908 / robert j . j o h n s, garry a . shea, & pratito puradyatmika Johnstone, I.M. 1978a. The ecology and distribution of Papua New Guinea seagrasses. I. Additions to the seagrass flora of Papua New Guinea. Aquatic Bot. 5: 229–233. Johnstone, I.M. 1978b. The ecology and distribution of Papua New Guinea seagrasses. I. The Fly Islands and Raboin Island. Aquatic Bot. 5: 235–243. Johnstone, I.M. 1982. Ecology and distribution of seagrasses. Pp. 497–512 in Gressit, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr W. Junk Publishers, The Hague. MacKinnon, K., G. Hatta, H. Halim, and A. Mangalik. 1996. The Ecology of Kalimantan. The Ecology of Indonesia Series, Volume III. Periplus Editions, Singapore. Polunin, N.V.C. 1983. The marine resources of Indonesia. Oceanogr. Mar. Biol. Ann. Rev. 21: 455–531. Rand, A.L., and L.J. Brass. 1940. Summary of the 1936–37 New Guinea Expedition. Bull. Am. Mus. Nat. Hist. 77: 341–380. Shea, G.A., P. Puradyatmika, A. Mandessy, and D. Martindale. 1996. Biodiversity Survey in the P.T. Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. P.T. Hatfindo Prima, Bogor, Indonesia. van Balgooy, M.M.J. 1996. Vegetation sketch of the Aru Islands. Ned. Comm. Internat. Natuur Besch Meded. 30: 1–13. Zieman, J.C. 1975. Seasonal variation of Turtle, Thalassia testudinum Ko¨nig, with reference to temperature and salinity effects. Aquatic Bot. 3: 149–175.

Appendix 5.6.1 Data Collection Methods The data collection methods for rainforest plots are described by Shea et al. (1996). Forest vegetation types were sampled using a sample plot method. A vegetation proforma was used to record stand structure, life forms, species composition, tree measurements, and tree co-ordinates. Stem diameter, bole height, crown height, total height, crown diameter, and special features were recorded for trees measuring 10 cm or more in diameter. This information was used for quantitative analysis and the preparation of profile diagrams for each of the sampled stands. In addition, collections were made for species that occurred within the stand, including both vascular and non-vascular plants. The important measurable quantities used for community sampling were the number of individuals or density (abundance). Density is the number of individuals per unit area, and is expressed in terms of individuals per ha (10,000 m2). The frequency of species is the number of times a species is recorded in a given number of small quadrats or at a given number of sample points. Species cover was assessed as either crown and shoot area (for herbaceous communities) or basal area (for forest communities). Basal area is the area outline of a plant near the ground surface. For trees, basal area involves measuring the tree diameter, usually at breast height (dbh), that is, 1.3 m above the ground, and calculating area by the formula p  r2, where p equals 3.14 and r is the half the diameter, or radius. For trees with buttresses, diameter is measured above buttress. In addition, tree height, stem diameter, and stem volume were also measured. Definitions and methods followed Mueller-Dombois and Ellenberg (1974). The ‘‘importance value’’ Curtis (1959) for component species is defined as the sum of

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Coastal Vegetation of Papua / 909

the relative density, relative frequency, and relative dominance, and is calculated as follows. Relative density is calculated by multiplying the number of species by 100 and then dividing by the total number of individuals in the plot or quadrat. Relative dominance is calculated by multiplying the dominance of a species by 100, and then dividing by the dominance of all species. Relative frequency is calculated by multiplying the frequency of each species by 100, and then dividing by the sum of the frequencies of all species. The three figures are summed to yield the importance value of a species, which reaches a maximum of 300 in stands consisting of only one tree species.

Literature Cited Curtis, J.J. 1959. The Vegetation of Wisconsin: An Ordination of Plant Communities. University of Wisconsin Press, Madison. Mueller-Dumbois, D., and H. Ellenberg. 1974. Aims and Methods of Vegetation Ecology. Wiley, New York. Shea, G.A., P. Puradyatmika, A. Mandessy, and D. Martindale. 1996. Biodiversity Survey in the P.T. Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. P.T. Hatfindo Prima, Bogor.

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PAGE 909

5.7. Lowland Swamp and Peat Vegetation of Papua robert j. johns, garry a. shea, and pratito puradyatmika w a mp v e g et a t i on , including peat swamp, occurs at all altitudes in Papua. These communities have been poorly studied and in earlier accounts of the vegetation of New Guinea (Paijmans 1976; Johns 1982) little reference is made to them. Swamp forest is most extensive in the lowlands along the southern coasts of Papua where Campnosperma, a tall forest tree reaching a height of up to 30–35 m, is the main forest dominant in swamps over large areas. The other common types of swamp vegetation are the sago swamp (Metroxylon sagu) and Pandanus swamp. As the areas of low-lying swamp forests are infilled with alluvial materials, the soil level builds up in the forest and a gradual transition occurs from swamp forest to mixed lowland tropical rainforest. The floristically most diverse and best-developed of the swamp forests are mixed in species composition, often including many species also found in the better drained rainforest. Tidal swamp vegetation (mangroves) is treated in Chapter 5.5. Brief mention is made here of freshwater swamp communities such as the small stands of Sonneratia which occur inland in poorly drained depressions, not in tidal swamps. Extensive areas of peat and swamp vegetation occur in the lowlands of New Guinea at altitudes of 3–35 (–50) m asl. Due to the large scale of the maps, the areas shown would also include areas of mangrove (tidal swamp vegetation). Detailed mapping of swamp and peat vegetation, with ground truthing, would be required for better maps. The vegetation ranges from open freshwater aquatic vegetation to medium or tall swamp forest, often with a sago substratum. Soils are often peaty mixed with recent fine alluvium (riverine). Deep peat deposits, up to 10 m thick, have been recorded in the southern regions of Papua (Shea et al. 1998). Swamp forest is the dominant plant community in the mainly wooded backswamps of the larger rivers such as the tributaries of the Mamberamo River, while scattered patches of swamp forest occur within grass swamps on levees and meander scrolls of old river courses. In transitions between swamp forest and swamp woodland, the forest tends to occupy the better-aerated and better-drained sites. Swamp forest is tallest in strongly fluctuating, but permanent, swamps just behind river levees, while in quiet swamps further away from the river it is lower, thinner stemmed, and usually more open. In Papua New Guinea extensive areas in Bougainville have a very distinct swamp forest dominated by Terminalia brassii. This species does not grow in Papua but has potential for the reforestation of lowland swamps.

S

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

910

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Lowland Swamp and Peat Vegetation of Papua / 911

Lowland Swamp Forest in Papua Species composition of swamp forests varies depending largely on site history and seed availability. Swamp forest communities are widespread and patchy in their distribution. The canopy in swamp forest is even and rather open, or occasionally dense, and individual trees can reach some 30 m in height. Common canopy trees are Campnosperma, Terminalia canaliculata, Nauclea coadunata, Syzygium, Alstonia scholaris, Bischofia javanica, and Palaquium. Many other trees can be scattered through the swamp forest. There is a sparse subcanopy but several species are common: Alstonia spatulata, Barringtonia, Diospyros, Garcinia, and Gymnotroches axillaris. Where flooding is prolonged in the swamp forest there are few shrubs and herbs. Often thin lianas, fleshy climbers, and climbing ferns are common. The most extensive areas of lowland swamp forest are in south Papua and the Mamberamo Basin.

mixed lowland swamp forest Where the inundation is short open swamp vegetation is formed with grasses up to 1 m. The swamp forest (van Royen 1963) has scattered specimens of Acacia, Banksia dentata, Borassus, Calophyllum, Eucalyptus, Garcinia, Livistona, Licuala, Litsea, Timonius, Tristania suaveolens, and an occasional Dillenia alata. The undergrowth has numerous grass species, including Aristida, Eragrostis, Eriachne, Imperata cylindrica, Ischaemum, Pseudopogonatherum, and Saciolepis. Other associated genera include Desmodium, Melastoma, Merremia, and Nepenthes (van Royen 1963). Several types of swamp forest have been surveyed by Shea et al. (1998) from south Papua, particularly within the Freeport Contract of Work Area. These are described here. The first example is a swamp forest in which the canopy was dominated by Alstonia scholaris, Hopea novoguineensis, Garcinia dulcis, and Terminalia copelandii. Other common trees included Dillenia alata, Pandanus tectorius, Gmelina, and Rhus taitensis (Table 5.7.1). The estimated total basal area for trees with a diameter greater than 5 cm dbh is 15.6 m2/ha and the total clear bole volume is 62.7 m3/ha. Mixed swamp forests also occur between the east levee road and the Minajerwi River in south Papua. In one stand sampled by Shea et al. (1998) at the east end of the cross levee road at Biodiversity Research Site No. 3, the dominant canopy trees were Vatica papuana, Stemonurus, Terminalia copelandii, and Campnosperma brevipetiolata. Other genera present included Linociera, Intsia bijuga, Palaquium, Cerbera, Dillenia alata, Myristica, palm, Mallotus, Syzygium, and Hopea novoguineenis. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 21.0 m2/ha and the total volume was 169.7 m3/ha (Table 5.7.2). In a second stand Hopea novoguineensis, Terminalia copelandii, Pandanus tectorius, and Alstonia scholaris were recorded as the dominant canopy species. Other trees in the community were Canarium decumanum, Dillenia alata, Poikilospermum papuana, and Myristica (Table 5.7.3). The estimated total basal area for trees with a

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PAGE 911

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7.15 3.86

Garcinia dulcis

Gmelina sp.

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Source: Shea et al. (1998).

62.74

7.67

Terminalia copelandii

Total

1.33

Syzygium argentea

3.11

Polyosma sp. 1.77

4.49

Pandanus tectorius

Rhus taitensis

0.58

Myristica sp.

12.02

5.57

Dillenia alata

Hopea novoguineensis

1.48

Canarium decumanum

Alstonia scholaris 2.16

11.56

Species

Calophyllum papuana

Total clear bole volume (m3/ha)

325

35

5

15

15

20

5

80

10

35

45

15

10

35

Absolute density

100.00

10.77

1.54

4.62

4.62

6.15

1.54

24.62

3.08

10.77

13.85

4.62

3.08

10.77

Relative density (%)

15.57

1.62

0.20

0.76

0.66

0.99

0.13

2.98

0.96

1.85

1.50

0.52

0.38

3.01

Absolute dominance (m2/ha)

100.00

10.41

1.27

4.91

4.22

6.38

0.82

19.14

6.20

11.89

9.64

3.36

2.43

19.34

Relative dominance (%)

5.38

0.50

0.13

0.38

0.38

0.38

0.13

0.75

0.25

0.63

0.75

0.38

0.25

0.50

Absolute frequency

100.00

9.30

2.33

6.98

6.98

6.98

2.33

13.95

4.65

11.63

13.95

6.98

4.65

9.30

Relative frequency (%)

300.00

30.48

5.13

16.50

15.81

19.51

4.68

57.71

13.92

34.28

37.44

14.95

10.16

39.41

Importance value

Table 5.7.1. Swamp forest dominated by Alstonia scholaris, Hopea novoguineensis, Garcinia dulcis, and Terminalia copelandii

912 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 912

1.70 0.39 1.49 1.21 9.49 0.23 0.44 5.22 1.16 2.29 4.27 1.49 1.26 1.97

Aglaia sp.

Buchanania macrophylla

Calophyllum sp.

Campnosperma sp.

Campnosperma brevipetiolata

Canarium indicum

Canarium sp.

Cerbera sp.

Cryptocarya sp.

Dacryoides sp.

Dillenia alata

Diospyros sp.

Eugenia sp.

Eugenia speciosa

Species

Total clear bole volume (m3/ha)

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10

15

5

25

15

10

20

5

5

10

5

5

5

5

Absolute density

1.18

1.78

0.59

2.96

1.78

1.18

2.37

0.59

0.59

1.18

0.59

0.59

0.59

0.59

Relative density (%)

0.21

0.16

0.13

0.63

0.28

0.18

0.64

0.06

0.04

0.93

0.13

0.13

0.04

0.13

Absolute dominance (m2/ha)

1.02

0.77

0.62

2.99

1.33

0.86

3.07

0.30

0.20

4.43

0.64

0.64

0.21

0.61

Relative dominance (%)

0.01

0.01

0.00

0.01

0.01

0.01

0.01

0.00

0.00

0.01

0.00

0.00

0.00

0.00

Absolute frequency

1.18

1.78

0.59

2.96

1.78

1.18

2.37

0.59

0.59

1.18

0.59

0.59

0.59

0.59

Relative frequency (%)

(continued)

3.38

4.32

1.81

8.91

4.88

3.23

7.80

1.48

1.39

6.79

1.82

1.82

1.39

1.80

Importance value

Table 5.7.2. Mixed swamp forest dominated by Vatica russak, Stemonurus, Terminalia copelandii, and Campnosperma brevipetiolata

Lowland Swamp and Peat Vegetation of Papua / 913

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9.55 5.86 0.59 0.22 2.98 3.89

Macaranga mappa

Mallotus ricinoides

Mallotus sp.

Myristica sp.

1.19

Hopea sp.

Linociera sp.

2.88

Hopea novoguineensis

Intsia bijuga

0.36

0.83

Gmelina sp. 0.25

0.33

Glochidion sp.

Haplolobus floribundus

0.71

Garcinia sp.

Gnetum gnemon

1.98

Total clear bole volume (m3/ha)

Ficus adenosperma

Species

Table 5.7.2. (Continued)

03-15-07 07:34:23

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35

20

5

10

40

20

10

20

5

5

5

5

10

15

Absolute density

4.14

2.37

0.59

1.18

4.73

2.37

1.18

2.37

0.59

0.59

0.59

0.59

1.18

1.78

Relative density (%)

0.49

0.42

0.04

0.09

1.25

1.06

0.16

0.35

0.08

0.05

0.09

0.05

0.08

0.31

Absolute dominance (m2/ha)

2.33

2.01

0.2

0.45

5.93

5.04

0.77

1.68

0.36

0.22

0.43

0.23

0.38

1.46

Relative dominance (%)

0.02

0.01

0.00

0.01

0.02

0.01

0.01

0.01

0.00

0.00

0.00

0.00

0.01

0.01

Absolute frequency

4.14

2.37

0.59

1.18

4.73

2.37

1.18

2.37

0.59

0.59

0.59

0.59

1.18

1.78

Relative frequency (%)

10.61

6.75

1.38

2.81

15.4

9.78

3.14

6.42

1.54

1.40

1.62

1.41

2.75

5.01

Importance value

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Source: Shea et al. (1998).

Total

169.66

40.96

Vatica russak

4.40 10.54

Terminalia copelandii

Terminalia sp.

10.70

2.71

Syzygium sp.

Terminalia complanata

0.53

15.42

Stemonurus sp. 0.99

0.39

Sloanea pullei

Sterculia sp.

1.22

Rhus taitensis

Sterculia parcinsonii

0.61

Palm sp.

Pterygota sp.

3.06 7.85

Palaquium sp. 0.38

4.68

Palaquium lobbianum

Paraserianthes falcataria

0.99

Palaquium amboinensis

845

180

20

20

35

20

5

5

130

5

5

5

5

45

10

5

5

100.00

21.30

2.37

2.37

4.14

2.37

0.59

0.59

15.38

0.59

0.59

0.59

0.59

5.33

1.18

0.59

0.59

21.01

4.90

1.00

0.50

1.27

0.43

0.09

0.10

2.49

0.04

0.13

0.09

0.05

0.79

0.33

0.47

0.09

100.00

23.34

4.77

2.36

6.06

2.05

0.42

0.45

11.83

0.19

0.61

0.43

0.26

3.77

1.58

2.25

0.44

0.42

0.09

0.01

0.01

0.02

0.01

0.00

0.00

0.07

0.00

0.00

0.00

0.00

0.02

0.01

0.00

0.00

100.00

21.3

2.37

2.37

4.14

2.37

0.59

0.59

15.38

0.59

0.59

0.59

0.59

5.33

1.18

0.59

0.59

300.00

65.95

9.51

7.09

14.35

6.79

1.60

1.64

42.60

1.38

1.80

1.61

1.44

14.42

3.95

3.43

1.63

Lowland Swamp and Peat Vegetation of Papua / 915

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65.86

Total

Source: Shea et al. (1998).

18.84

1.42

Poikilospermum papuana

Terminalia copelandii

15.51

Pandanus tectorius

Dillenia alata 0.47

0.85

Canarium decumanum

Myristica sp.

1.69

Alstonia scholaris

15.87

11.21

Species

Hopea novoguineensis

Total clear bole volume (m3/ha)

175

40

10

25

5

70

10

5

10

Absolute density

100.00

22.86

5.71

14.29

2.86

40.00

5.71

2.86

5.71

Relative density (%)

11.78

3.18

0.28

2.41

0.19

3.55

0.24

0.33

1.59

Absolute dominance (m2/ha)

100.00

27.02

2.4

20.44

1.63

30.11

2.05

2.82

13.52

Relative dominance (%)

2.50

0.50

0.13

0.38

0.13

0.75

0.25

0.13

0.25

Absolute frequency

100

20

5

15

5

30

10

5

10

Relative frequency (%)

300.00

69.87

13.12

49.73

9.49

100.11

17.77

10.68

29.24

Importance value

Table 5.7.3. Mixed swamp forest dominated by Hopea novoguineensis, Terminalia copelandii, Pandanus tectorius, and Alstonia scholaris

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Lowland Swamp and Peat Vegetation of Papua / 917

diameter greater than 5 cm dbh was 11.8 m2/ha and the total clear bole volume was 65.9 m3/ha. This forest was impoverished, with only eight species recorded, possibly due to limited seed sources. Another swamp forest had the canopy dominated by Hopea novoguineensis, Terminalia copelandii, Alstonia scholaris, and Polyosma. Other trees here included Garcinia dulcis, Pandanus tectorius, and Rhus taitensis (Table 5.7.4). The estimated total basal area for trees with a diameter greater than 5 cm dbh was 22.8 m2/ha and the total clear bole volume was 106.3 m3/ha.

campnosperma swamp forest The dominant species in swamp forest, Campnosperma brevipetiolata, is widespread particularly on alluvial and peat sites along the southern regions of Papua. It commonly dominates permanently flooded sites with soils that are often peaty with a thick layer of organic matter on the surface. C. brevipetiolata is a tall forest tree 30–35 m (rarely to 50 m) in height, with a long thick straight bole, dense flattopped crown, and large leaves to up 17–18 cm long. It possibly grows in association with Campnosperma coriacea, but we have seen only one herbarium collection from Papua although this species is widespread in swamp forests in Papua New Guinea. Pure stands can be easily recognized in aerial photographs by their graytoned, smooth, even canopy. From the air, the stands are sometimes mistaken for stands of Calophyllum suberosum, which habitually grows in swamps or periodically inundated ground and river banks that are sometimes subject to tidal influence (Stevens 1980). Several collections of C. suberosum are known from the Asmat and the Digul River in southern Papua. Stevens (1980) noted that the (floating) seeds may be dispersed by rivers. Buttresses are common on many trees; others have stilt roots (C. suberosum has stilt roots to 2 m tall) and knee roots. The forest trees are densely covered with epiphytes and climbers. Sago can form a common, even a dense, understory in these forests. Lundquist (1942) made terrestrial reconnaissance of the areas northwest, north, east, and south of the McCluer Gulf/Bintuni Bay from Teminabuan to the east end of the bay, and also mapped the south side, Bomberai (west of Babo). On the south coast the mapping covered areas from Lakahia Bay to the Uta River. The Campnosperma forests were deeply inundated in the wet season. Lundquist observed about 45,000 ha, of which 29,000 ha were dense forest. Sago usually occurred in the understory; the more open the forest canopy was. the greater the density of sago. The dbh of Campnosperma was usually below 30 (– 40) cm. Vink (pers. comm.) observed extensive areas of Campnosperma forest along the north coast of Papua.

melaleuca swamp forest The swamp forest of the monsoon areas in southeast Papua are dominated by Melaleuca leucadendra in southern New Guinea. These forests are flooded for most of the year (van Royen 1963). The Melaleuca, or paper-bark, forest has an even,

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PAGE 917

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5.93

Garcinia dulcis

03-15-07 07:34:25

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2.86

Rhus taitensis

Source: Shea et al. (1998).

Total

106.25

28.98

7.60

Polyosma sp.

Terminalia copelandii

4.57

Pandanus tectorius

31.38

1.27

Hopea novoguineensis

2.18

Ficus sp.

0.7

Canarium indicum

Dillenia sp.

0.67

Canarium decumanum 1.87

0.51

Calophyllum sp.

Canarium sp.

2.70

15.04

Calophyllum papuana

Alstonia scholaris

Species

Total clear bole volume (m3/ha)

440

70

20

50

15

190

35

5

5

5

5

5

5

10

20

Absolute density

100.00

15.91

4.55

11.36

3.41

43.18

7.95

1.14

1.14

1.14

1.14

1.14

1.14

2.27

4.55

Relative density (%)

22.82

5.45

0.68

1.88

1.18

7.80

1.40

0.30

0.34

0.29

0.14

0.14

0.11

0.44

2.66

Absolute dominance (m2/ha)

100.00

23.90

3.00

8.25

5.16

34.17

6.12

1.33

1.49

1.27

0.62

0.62

0.47

1.93

11.68

Relative dominance (%)

5.13

0.63

0.50

0.63

0.25

1.00

0.75

0.13

0.13

0.13

0.13

0.13

0.13

0.25

0.38

Absolute frequency

100.00

12.20

9.76

12.20

4.88

19.51

14.63

2.44

2.44

2.44

2.44

2.44

2.44

4.88

7.32

Relative frequency (%)

Table 5.7.4. Swamp forest dominated by Hopea novoguineensis, Terminalia copelandii, Alstonia scholaris, and Polyosma

300.00

52.00

17.30

31.81

13.45

96.86

28.71

4.91

5.06

4.85

4.19

4.19

4.04

9.09

23.54

Importance value

918 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 918

Lowland Swamp and Peat Vegetation of Papua / 919

almost pure, one-layered canopy (Paijmans 1976) up to 30 m tall. The stands are usually open, with slender boles to 30 m tall. They form narrow bands along seasonally dry swamps in southeast Papua (van Royen 1956, 1963). The trunks of many trees are charred due to frequent fires during the dry season. In some areas in south Papua New Guinea Melaleuca trees can have a double root system. In the dry season there is an exposed root system some 2–3 m above ground level. This exposed system functions during the wet season (R. Johns, pers. obs.). Melaleuca leucadendra has characteristic white stems. The forest is flooded for most of the year, the ground often remaining wet throughout the dry season. Wetter areas often support a floating vegetation dominated by Azolla and Ceratophyllum.

erythrina swamp Erythrina fusca is dominant in these forests, forming a closed canopy and lacking a distinct sub-canopy layer. Scattered trees in these forests include Buchanania, Carota, Corynocarpus australasica, Garcinia sp., Glochidion sp., Hibiscus tiliaceous, Sarcocephalus sp., and Vitex cofassus (van Royen 1963). Common climbers are Microsorium and Stenochlaena. Numerous epiphytes occur in the forest, including Myrmecodia, Hydnophytum, and also Dendrobium.

barringtonia, leptospermum swamp This swamp forest is poorly developed, composed of Barringtonia tetraptera and Leptospermum abnorme, with scattered Alyxia, Rhodamnia, and Syzygium. Van Royen (1963) listed the undergrowth species as Fimbristylis griffithii, with scattered Eriocaulon, Helminthostachys zeylanica, Ilysanthes, and Xyris complanata. Epiphytes are abundant in this type of swamp forest. These include Amylotheca, Cyclophorus, Dischidia, Dendrobium, Eria, Hoya, Pyrrosia, and Psilotum nudum.

pandanus swamp Swamp areas dominated by Pandanus are widespread in Papua and occupy a habitat similar to that of the sago palm. The pandanus forms an open community to 8–10 m in height on stagnant to frequently flooded sites in freshwater to brackish situations. Stone (1982) listed P. tectorius from saline and brackish swamps near the coast in New Guinea. Freshwater swamps and swamp woodlands can have several species of Pandanus: P. hollrungii, P. hysterix, P. kaernbachii (Stone 1982), P. lauterbachii, P. leiophyllus, P. scabribracteatus, and the widespread savanna and coastal species P. tectorius. Sometimes trees are scattered through the canopy. These communities are very poorly known. P. scabribracteatus, which is very closely related to P. kaernbachii, occurs in the southern border areas of Papua near the Papua New Guinea border. It is gregarious in shallow swamps, forming a canopy at 10–14 m height. As noted by Stone (1982), freshwater crabs may be found in the leaf axils. Pandanus swamp communities are extensive in the valleys to the north of the Central Ranges in New Guinea. Shea et al. (1998) distinguished several swamp communities from south Papua with Pandanus tectorius often dominant. They distinguished the forests by the

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PAGE 919

920 / robert j . johns , garry a. shea, & p ratito puradyatmika

species composition of the associated trees: Terminalia copelandii, Hopea novoguineenis, Garcinia dulcis, Polyosma, Rhus taitensis, Gmelina, and Pandanus tectorius (Table 5.7.5); and Hopea novoguineensis, Terminalia copelandii, Alstonia scholaris, Polyosma, Garcinia dulcis, Pandanus tectorius, and Rhus taitensis (Table 5.7.6). Swamp forest can be dominated by Terminalia copelandii-Hopea novoguineenisGarcinia dulcis-Polyosma. Other trees include Rhus taitensis, Gmelina, and Pandanus tectorius (Table 5.7.5). The estimated total basal area for trees with a diameter greater than 5 cm dbh was 18.2 m2/ha and the total clear bole volume was 89.4 m3/ha. Some stands along the PT Freeport Indonesia road to Cargo Dock at Mile 23 in south Papua have a substratum of pandans rather than sago palms. One stand was sampled that was located off the PTFI road to Cargo Dock near Mile 18. The dominant genera were Pandanus, Nauclea, Cryptocarya, Palaquium, Campnosperma, and Syzygium (Table 5.7.6). The plant community was represented by the name of the four most dominant genera: Pandanus, Nauclea, Cryptocarya, and Palaquium. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 15 m2/ha and the total volume was 63 m3/ha.

metroxylon sagu (sago) swamp Pure sago swamp and sago in mixed forest, covers very large areas in the McCluer Gulf/Bintuni Bay from Teminabuan to the east end of the bay, and also on the south side, Bomberai west of Babo (Lundquist 1942). Exploitation of sago by the local population had only a minimal effect on the vegetation. Extensive stands of sago occur in the southeast of Papua. The swamp remains flooded for long periods and the canopy is closed. Undergrowth is limited to Fimbristylis, with patches of Lemna floating in areas of open water. Similar swamp forests also occur in the Mamberamo basin. Brass (1938) includes a detailed list of the plants occurring in the sago swamp. Sago palm (Metroxylon sagu) grows best in shallow, permanently swampy woodland where there is a regular inflow of fresh water. Under favorable conditions, sago may reach 15 m in height, with starch-producing stems reaching 10 m in height. Sago generally multiplies by forming suckers around the base of old stems, usually after flowering. However, sago may also reproduce by seed and shows considerable variation and cross-breeding. All gradations occur from stands of pure sago palm, virtually without trees, to forest with an open lower layer of sago. Under dense sago there is no undergrowth; the peaty soil is layered with fallen dead fronds, and the palm’s numerous tiny pneumatophores form the only live ground cover (Paijmans 1976). Large areas of sago probably owe their origins to the activities of humans, because the sago regenerates aggressively where villagers have cleared the mixed lowland swamp forest for firewood and construction timber for building village houses. Once the rainforest trees have been felled the swamps are quickly dominated by sago which, due to its density, inhibits the regeneration of rainforest trees.

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PAGE 920

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

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03-15-07 07:34:26

8.38 2.50 16.61 0.86 6.05 6.85 48.15 89.40

Species

Garcinia dulcis

Gmelina sp.

Hopea novoguineensis

Pandanus tectorius

Polyosma sp.

Rhus taitensis

Terminalia copelandii

Total

Source: Shea et al. (1998).

Total clear bole volume (m3/ha)

285

120

20

20

5

90

5

25

Absolute density

100.00

42.11

7.02

7.02

1.75

31.58

1.75

8.77

Relative density (%)

18.18

9.30

1.33

1.35

0.39

3.65

0.64

1.51

Absolute dominance (m2/ha)

100.00

51.15

7.31

7.44

2.16

20.09

3.54

8.31

Relative dominance (%)

3.38

1.00

0.38

0.25

0.13

0.88

0.13

0.63

Absolute frequency

100.00

29.63

11.11

7.41

3.70

25.93

3.70

18.52

Relative frequency (%)

Table 5.7.5. Swamp forest dominated by Terminalia copelandii, Hopea novoguineenis, Garcinia dulcis, and Polyosma

300.00

122.89

25.44

21.87

7.62

77.59

8.99

35.60

Importance value

Lowland Swamp and Peat Vegetation of Papua / 921

PS

PAGE 921

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PS

Source: Shea et al. (1998).

99.81

Total

3.38

Litsea sp. 59.91

7.03

Intsia sp.

Metroxylon sp.

0.85

7.15

Diospyros sp. 5.38

7.59

Campnosperma sp.

Garcinia sp.

0.36

Alstonia sp.

Endospermum sp.

8.16

Aglaia sp.

Species

Total clear bole volume (m3/ha)

235

155

5

5

10

15

10

20

5

10

Absolute density

100.00

67.3

2.1

2.1

4.2

6.3

4.2

8.5

2.1

4.2

Relative density (%)

24.20

19.64

0.40

0.63

0.61

0.14

1.02

0.98

0.06

0.73

Absolute dominance (m2/ha)

100.00

81.1

1.6

2.6

2.5

0.6

4.2

4.0

0.3

3.1

Relative dominance (%)

Table 5.7.6. Forest dominated by Campnosperma and Intsia with a Pandanus substratum

2.4

1.0

0.1

0.1

0.2

0.3

0.1

0.4

0.1

0.1

Absolute frequency

100.0

43.6

4.2

4.2

9.3

10.7

4.2

16.6

4.2

4.2

Relative frequency (%)

300.0

192.0

7.8

8.8

16.3

17.6

12.5

29.1

6.5

11.4

Importance value

922 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 922

Lowland Swamp and Peat Vegetation of Papua / 923

Other swamp forest communities are dominated by Hopea novoguineensis, Terminalia copelandii, Alstonia scholaris, and Garcinia dulcis. Other trees in these include Ardisia, Calophyllum papuana, Glochidion, Canarium indicum, Rhus taitensis, Garcinia dulcis, Myristica, Polyosma, and Metroxylon sagu (Table 5.7.7). The estimated total basal area for trees with a diameter greater than 5 cm dbh was 24.2 m2/ha and the total clear bole volume was 114.2 m3/ha. Forests dominated by Metroxylon sagu, can occur with the canopy dominated by Terminalia copelandii, Alstonia scholaris, and Hopea novoguineensis. Other trees included Myristica fatua, Garcinia dulcis, Rhus taitensis, and Dillenia alata (Table 5.7.8). The estimated total basal area for trees with a diameter greater than 5 cm dbh (or basal diameter in the case of short stemmed sago palms) was 35.3 m2/ha and the total clear bole volume was 53.6 m3/ha. Sago swamp forest can also have the canopy layer with numerous Terminalia copelandii, Syzygium argentea, and Alstonia scholaris. Other trees include Rhus taitensis, Polyosma, Canarium, Garcinia dulcis, and Pandanus tectorius (Table 5.7.9). The estimated total basal area for trees with a diameter greater than 5 cm dbh was 41.7 m2/ha and the total clear bole volume was 88.9 m3/ha. Terminalia copelandii can also be mixed with Hopea novoguineensis and Garcinia dulcis. Associated genera included Polyosma papuana, Alstonia scholaris, and Dillenia alata (Table 5.7.10). The estimated total basal area for trees with a diameter greater than 5 cm dbh was 104 m2/ha and the total clear bole volume was 161 m3/ha. Peat swamps are often dominated by woodland and forest communities with a sago palm substratum. Some of these communities are open with a tree overstory, while other are disturbed where the mature overstory trees have died due to excessive inundation and sedimentation. Two stands were sampled in south Papua. A stand located off the PTFI road to Cargo Dock near Mile 21 had as its dominant genera Metroxylon, Campnosperma, Intsia, and Pandanus. Other important genera were Palaquium, Diospyros, Ficus, and Garcinia (Table 5.7.11). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 12 m2/ha and the total volume was 54 m3/ha. The dominant species in a second stand were Metroxylon sagu, Palaquium, Octomeles sumatrana, and Diospyros. This stand was located off the PTFI road to Cargo Dock at approximately Mile 18 in south Papua. Other dominant trees were Palaquium, Campnosperma, Syzygium, Aglaia, Terminalia canaliculata, Pandanus, Endospermum, Intsia, and Garcinia (Table 5.7.12). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 31.5 m2/ha and the total clear bole volume was 158.4 m3/ha. A plot was also placed in forest dominated by Intsia bijuga, Hopea novoguineensis, Artocarpus, and Palaquium lobbianum with a Metroxylon sagu substratum. Other trees included Stemonurus, Macaranga mappa, Terminalia complanata, Polyalthia glauca, Pometia pinnata, Vatica papuana, Homalanthus, Alstonia scholaris, and Ficus benjamina (Table 5.7.13). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 23.3 m2/ha and the total clear bole volume was 176.8 m3/ha. This stand is located off the PTFI road to Cargo Dock at approximately Km 20–21 in south Papua.

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PAGE 923

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2.38 2.00 4.78 3.17

Canarium indicum

Canarium sp.

Garcinia dulcis

Glochidion sp.

03-15-07 07:34:28

PS

0.81

Rhus taitensis

Source: Shea et al. (1998).

Total

114.24

37.09

0.72

Polyosma sp.

Terminalia copelandii

0.67

1.33

Myristica sp.

Poa sp.

0.00

39.36

Metroxylon sagu

Hopea novoguineensis

0.64

2.89

Calophyllum papuana

Gmelina sp.

2.84

15.57

Ardisia sp.

Alstonia scholaris

Species

Total clear bole volume (m3/ha)

425

135

10

5

5

10

5

165

5

5

20

5

15

5

10

25

Absolute densit

100.00

31.76

2.35

1.18

1.18

2.35

1.18

38.82

1.18

1.18

4.71

1.18

3.53

1.18

2.35

5.88

Relative density (%)

24.20

7.78

0.30

0.14

0.15

0.33

0.25

8.35

0.19

0.47

0.90

0.33

0.46

0.55

0.58

3.41

Absolute dominance (m2/ha)

100.00

32.16

1.25

0.56

0.62

1.38

1.04

34.51

0.77

1.96

3.73

1.34

1.91

2.26

2.41

14.09

Relative dominance (%)

16.88

2.00

1.88

1.75

1.63

1.50

1.38

1.25

1.13

1.00

0.88

0.75

0.63

0.50

0.38

0.25

Absolute frequency

100.00

11.85

11.11

10.37

9.63

8.89

8.15

7.41

6.67

5.93

5.19

4.44

3.70

2.9

2.22

1.48

Relative frequency (%)

300.00

75.78

14.71

12.11

11.43

12.62

10.36

80.74

8.62

9.06

13.62

6.96

9.15

66.4

6.98

21.46

Importance value

Table 5.7.7. Swamp forest dominated by Hopea novoguineensis, Terminalia copelandii, Alstonia scholaris, and Garcinia dulcis

924 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 924

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

1.34

Garcinia dulcis

CH50

03-15-07 07:34:29

PS

Source: Shea et al. (1998).

53.59

1.98

Rhus taitensis

Total

1.02

Polyosma sp. 24.83

5.84

Myristica fatua

Terminalia copelandii

0.00

Metroxylon sagu

0.66

0.34

Ficus sp.

Inocarpus fagifer

0.83

Dillenia alata

8.60

1.66

Ardisia sp.

Hopea novoguineensis

6.48

Alstonia scholaris

Species

Total clear bole volume (m3/ha)

175

60

10

5

10

10

5

40

10

5

5

5

10

Absolute density

100.00

34.29

5.71

2.86

5.71

5.71

2.86

22.86

5.71

2.86

2.86

2.86

5.71

Relative density (%)

35.26

4.71

0.47

0.20

0.87

23.75

0.23

2.00

0.39

0.11

0.26

0.23

2.05

Absolute dominance (m2/ha)

100.00

13.37

1.34

0.58

2.46

67.34

0.66

5.66

1.11

0.30

0.73

0.66

5.81

Relative dominance (%)

3.5

1.00

0.13

0.13

0.25

0.25

0.13

0.75

0.25

0.13

0.13

0.13

0.25

Absolute frequency

100.00

28.57

3.57

3.57

7.14

7.14

3.57

21.43

7.14

3.57

3.57

3.57

7.14

Relative frequency (%)

300

76.22

10.62

7.01

15.32

80.19

7.09

49.94

13.96

6.73

7.16

7.09

18.66

Importance value

Table 5.7.8. Sago swamp forest dominated by Metroxylon sagu, Terminalia copelandii, Alstonia scholaris, and Hopea novoguineensis Lowland Swamp and Peat Vegetation of Papua / 925

PAGE 925

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49.35 88.94

Terminalia copelandii

Total

Source: Shea et al. (1998).

6.68

5.14

Rhus taitensis

Syzygium argentea

6.15

Polyosma sp.

0.00

Metroxylon sagu 3.24

4.19

Garcinia dulcis 1.98

1.14

Dillenia alata

Pandanus tectorius

2.70

Calophyllum sp.

Myristica sp.

8.36

Alstonia scholaris

Species

Total clear bole volume (m3/ha)

270

145

10

20

30

15

5

10

15

5

5

10

Absolute density

100.00

53.70

3.70

7.41

11.11

5.56

1.85

3.70

5.56

1.85

1.85

3.70

Relative density (%)

41.70

9.40

1.68

1.40

1.36

0.79

0.47

23.75

0.81

0.24

0.37

1.43

Absolute dominance (m2/ha)

100.00

22.55

4.02

3.37

3.26

1.88

1.14

56.94

1.95

0.57

0.89

3.43

Relative dominance (%)

3.63

0.88

0.25

0.38

0.75

0.25

0.13

0.25

0.25

0.13

0.13

0.25

Absolute frequency

100.00

24.14

6.90

10.34

20.69

6.90

3.45

6.90

6.90

3.45

3.45

6.90

Relative frequency (%)

300.00

100.39

14.62

21.12

35.06

14.33

6.44

67.54

14.40

5.87

6.19

14.03

Importance value

Table 5.7.9. Sago swamp forest dominated by Metroxylon sagu, Terminalia copelandii, Syzygium argentea, and Alstonia scholaris

926 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 926

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CH50

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PS

Source: Shea et al. (1998).

Total

Polyosma sp.

Terminalia copelandii 161.12

0.57

51.26

0.72

Rhus taitensis

0.55

Glochidion sp.

3.67

6.22

Garcinia dulcis

54.36

3.17

Dillenia alata

Polyosma papuana

0.72

Canarium indicum

Metroxylon sagu

4.26

Canarium decumanum

30.44

0.81

Calophyllum papuana

Hopea novoguineensis

4.39

Alstonia scholaris

Species

Total clear bole volume (m3/ha)

405

5

125

5

15

35

135

5

30

10

5

20

5

10

Absolute density

100.00

1.23

30.86

1.23

3.70

8.64

33.33

1.23

7.41

2.47

1.23

4.94

1.23

2.47

Relative density (%)

104.02

0.17

9.18

0.19

0.80

83.11

6.51

0.18

1.12

0.74

0.14

0.93

0.20

0.75

Absolute dominance (m2/ha)

100.00

0.16

8.82

0.18

0.77

79.90

6.25

0.17

1.08

0.71

0.13

0.90

0.20

0.72

Relative dominance (%)

4.38

0.13

0.88

0.13

0.13

0.38

0.88

0.13

0.50

0.25

0.13

0.50

0.13

0.25

Absolute frequency

100.00

2.86

20.00

2.86

2.86

8.57

20.00

2.86

11.43

5.71

2.86

11.43

2.86

5.71

Relative frequency (%)

300.00

4.26

59.69

4.27

7.33

97.12

59.59

4.26

19.91

8.90

4.22

17.26

4.29

8.90

Importance value

Table 5.7.10. Mixed sago swamp forest dominated by Metroxylon sagu, Terminalia copelandii, Hopea novoguineensis, and Garcinia dulcis Lowland Swamp and Peat Vegetation of Papua / 927

PAGE 927

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

0.38

Garcinia sp.

CH50

03-15-07 07:34:30

PS

Source: Shea et al. (1998).

53.63

2.65

Pandanus sp.

Total

0.11

Metroxylon sagu 1.37

20.18

Macaranga sp.

Palaquium sp.

0.16

Intsia bijuga

Myristica sp.

0.25 10.55

Heritiera sp.

0.13

0.79

Harpullia sp.

0.55

Campnosperma brevipetiolata

Ficus sp.

16.51

Species

Diospyros sp.

Total clear bole volume (m3/ha)

360

55

20

5

55

5

20

10

5

5

5

15

160

Absolute density

100.00

15.28

5.56

1.39

15.28

1.39

5.56

2.78

1.39

1.39

1.39

4.17

44.44

Relative density (%)

11.93

0.84

0.23

0.04

5.79

0.04

1.40

0.08

0.05

0.08

0.11

0.14

3.14

Absolute dominance (m2/ha)

100.00

7.02

1.96

0.33

48.55

0.33

11.75

0.66

0.40

0.64

0.95

1.13

26.29

Relative dominance (%)

Table 5.7.11. Open forest dominated by Metroxylon, Campnosperma, Intsia, Pandanus, and sago

3.8

0.7

0.4

0.1

0.5

0.1

0.3

0.2

0.1

0.1

0.1

0.2

1.0

Absolute frequency

100.00

18.42

10.53

2.63

13.16

2.63

7.89

5.26

2.63

2.63

2.63

5.26

26.32

Relative frequency (%)

300.00

40.72

18.04

4.35

76.98

4.35

25.20

8.70

4.42

4.67

4.97

10.56

97.05

Importance value

928 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 928

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

7.59 7.15 0.85 5.38 7.03 3.38

Campnosperma brevipetiolata

Diospyros sp.

Endospermum sp.

Garcinia sp.

Intsia bijuga

Litsea sp.

CH50

03-15-07 07:34:30

PS

0.81 4.44 3.88 3.59

Pandanus sp.

Syzygium sp.

Terminalia canaliculata

Teijsmanniodendron sp.

Source: Shea et al. (1998).

158.39

28.18

Palaquium sp.

Total

16.57

Nauclea sp.

Octomeles sumatrana

0.95 0.16

Myristica sp.

59.91

0.36

Alstonia sp.

Metroxylon sagu

8.16

Aglaia sp.

Species

Total clear bole volume (m3/ha)

385

5

5

15

20

75

10

5

15

155

5

5

10

15

10

20

5

10

Absolute density

100.00

1.30

1.30

3.90

5.19

19.48

2.60

1.30

3.90

40.26

1.30

1.30

2.60

3.90

2.60

5.19

1.30

2.60

Relative density (%)

31.51

0.43

0.69

0.79

0.18

3.46

1.49

0.06

0.21

19.64

0.40

0.63

0.61

0.14

1.02

0.98

0.06

0.73

Absolute dominance (m2/ha)

100.00

1.36

2.20

2.52

0.58

10.97

4.73

0.18

0.65

62.33

1.28

1.99

1.92

0.45

3.24

3.10

0.18

2.32

Relative dominance (%)

4.5

0.1

0.1

0.2

0.4

0.8

0.2

0.1

0.2

1.0

0.1

0.1

0.2

0.3

0.1

0.4

0.1

0.1

Absolute frequency

Table 5.7.12. Forest dominated by Metroxylon sagu, Palaquium, Octomeles sumatrana, and Diospyros

100.00

2.22

2.22

4.44

8.89

17.78

4.44

2.22

4.44

22.22

2.22

2.22

4.44

6.67

2.22

8.89

2.22

2.22

Relative frequency (%)

300.00

4.88

5.72

10.86

14.66

48.23

11.77

3.70

8.99

124.81

4.80

5.51

8.97

11.02

8.06

17.19

3.70

7.14

Importance value Lowland Swamp and Peat Vegetation of Papua / 929

PAGE 929

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

CH50

03-15-07 07:34:32

0.2 0.1 0.6 2.2 0.1

Endospermum sp.

Evodia sp.

Ficus benjamina

Gardenia sp.

0.5

Dysoxylum molle 0.5

0.1

Decaspermum fruticosum

Endospermum

1.6

Canarium indicum

Elaeocarpus altisectus

0.6

Cananga odorata

0.5

Barringtonia racemosa 0.5

7.8

Artocarpus fretessii

Campnosperma petiolata

0.1

Anaxagorea sp.

1.8

2.8

Alstonia scholaris

Calophyllum inophyllum

3.7

Aglaia sp.

Species

Total clear bole volume (m3/ha)

PS

1

2

1

1

2

1

1

1

3

2

2

2

1

2

1

2

3

Absolute density

0.476

0.952

0.476

0.476

0.952

0.476

0.476

0.476

1.429

0.952

0.952

0.952

0.476

0.952

0.476

0.952

1.429

Relative density (%)

0.011

0.242

0.059

0.018

0.026

0.097

0.052

0.009

0.168

0.062

0.082

0.188

0.065

0.485

0.013

0.252

0.297

Absolute dominance (m2/ha)

0.045

1.040

0.252

0.075

0.112

0.416

0.225

0.039

0.721

0.266

0.354

0.808

0.279

2.083

0.057

1.083

1.277

Relative dominance (%)

0.003

0.005

0.003

0.003

0.005

0.003

0.003

0.003

0.008

0.005

0.005

0.005

0.003

0.005

0.003

0.005

0.008

Absolute frequency

0.476

0.952

0.476

0.476

0.952

0.476

0.476

0.476

1.429

0.952

0.952

0.952

0.476

0.952

0.476

0.952

1.429

Relative frequency (%)

0.998

2.945

1.205

1.028

2.017

1.368

1.178

0.992

3.578

2.171

2.259

2.713

1.231

3.988

1.009

2.988

4.134

Importance value

Table 5.7.13. Forest dominated by Intsia bijuga, Hopea novoguineensis, Artocarpus, and Palaquium lobbianum with a Metroxylon sagu substratum

930 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 930

2.2 6.6 0.2 8.3

Homalanthus sp.

Hopea novoguineensis

Horsfielda sp.

Intsia bijuga

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

90.9 1.7 0.7 0.2 0.3 7.9 0.1 0.1 0.8 1.2 1.9

Myristica fatua

Myristica sp.

Nauclea orientalis

Palaquium lobbianum

Pandanus tectorius

Pentaphalagium sp.

Pimelodendron pinnatum

Planchonella lepticola

Polyalthia glauca

0.2

Melochia sp.

Metroxylon sp.

1.4

Macaranga mappa

Metroxylon sagu

0.1

Linociera sp.

13.9

0.1

Glochidion averhoa

Intsia palembanica

0.1

Gironniera sp.

CH50

03-15-07 07:34:32

PS

5

1

2

1

1

2

1

1

2

2

102

1

6

1

6

4

2

4

5

1

1

2.381

0.476

0.952

0.476

0.476

0.952

0.476

0.476

0.952

0.952

48.571

0.476

2.857

0.476

2.857

1.905

0.952

1.905

2.381

0.476

0.476

0.241

0.098

0.073

0.013

0.017

0.480

0.026

0.026

0.118

0.291

15.392

0.023

0.205

0.022

1.098

0.717

0.040

0.538

0.265

0.013

0.013

1.036

0.420

0.315

0.056

0.072

2.060

0.112

0.112

0.508

1.248

66.127

0.099

0.882

0.094

4.718

3.082

0.172

2.310

1.138

0.058

0.057

0.013

0.003

0.005

0.003

0.003

0.005

0.003

0.003

0.005

0.005

0.255

0.003

0.015

0.003

0.015

0.010

0.005

0.010

0.013

0.003

0.003

2.381

0.476

0.952

0.476

0.476

0.952

0.476

0.476

0.952

0.952

48.571

0.476

2.857

0.476

2.857

1.905

0.952

1.905

2.381

0.476

0.476

(continued)

5.798

1.373

2.220

1.009

1.024

3.965

1.065

1.064

2.413

3.153

163.270

1.051

6.596

1.046

10.433

6.892

2.076

6.120

5.900

1.01

1.009

Lowland Swamp and Peat Vegetation of Papua / 931

PAGE 931

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

CH50

03-15-07 07:34:33

PS

Source: Shea et al. (1998).

176.8

0.2

Xanthophyllum sp.

Total

4.4

Vatica russak

0.7 0.6

Triumfetta sp.

Triocarpus sp.

2.9

0.2

Syzygium macrophylla

Terminalia complanata

0.2

Sloanea pullei 1.6

0.1

Pygeum parviflorum

0.9

0.1

Premna corymbosa

Stemonurus sp.

0.1

Pometia pinnata

Syzygium sp.

2.3

Total clear bole volume (m3/ha)

Pometia acuminate

Species

Table 5.7.13. (Continued)

210

2

2

1

2

5

7

2

2

1

1

1

1

4

Absolute density

100.000

0.952

0.952

0.476

0.952

2.381

3.333

0.952

0.952

0.476

0.476

0.476

0.476

1.905

Relative density (%)

23.276

0.043

0.317

0.049

0.090

0.340

0.169

0.117

0.044

0.026

0.008

0.016

0.010

0.211

Absolute dominance (m2/ha)

100.000

0.184

1.362

0.210

0.388

1.460

0.726

0.503

0.189

0.111

0.034

0.070

0.045

0.907

Relative dominance (%)

0.525

0.005

0.005

0.003

0.005

0.013

0.018

0.005

0.005

0.003

0.003

0.003

0.003

0.010

Absolute frequency

100.000

0.952

0.952

0.476

0.952

2.381

3.333

0.952

0.952

0.476

0.476

0.476

0.476

1.905

Relative frequency (%)

300.000

2.089

3.267

1.162

2.292

6.222

7.393

2.408

2.094

1.063

0.987

1.022

0.997

4.716

Importance value

932 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 932

Lowland Swamp and Peat Vegetation of Papua / 933

An upper layer of trees may be present in some peat swamp forests with a dense lower layer of sago. Common tree species include Alstonia scholaris, Campnosperma, Nauclea coadunata, Terminalia canaliculata, Mitragyna ciliata, Timonius, Syzygium, Mangifera, Garcinia, Palaquium, Bischofia javanica, Barringtonia, and Gynotroches axillaris. Sago may be mixed with Pandanus spp. to form a dense forest community. Various species of Pandanus may be involved, but the taxonomy of these is still unclear. On permanently wet sites the sago swamp forest community there may be an overstory of stunted trees, including Palaquium spp., Octomeles sumatrana, Diospyros spp., Campnosperma spp., Syzygium spp., Terminalia canaliculata, Aglaia sp., Alstonia scholaris, Intsia bijuga, Nauclea coadunata, Myristica sp., Litsea spp., Garcina spp., and Teijsmanniodendron sp.

Peat Swamp Vegetation In Malesia peat forests occur in four regions: Sumatra (54,990 km2), Kalimantan (49,300 km2), Sulawesi (1,320 km2), and Papua (22,570 km2) (Dick 1991). The extensive areas of swamp vegetation occur in the lowlands of New Guinea at an altitude of 3–35 (–50) m asl. The peat vegetation ranges from open water aquatic vegetation to medium or tall peat swamp forest with a sago substratum. Soils are peaty mixed with recent fine alluvium (riverine). Deep peat deposits, up to 10 m thick, have been recorded in the southern regions of Papua (Shea et al. 1998). The peat swamp forests were mapped from satellite imagery but with limited ground truthing. In his description of plant zonation Johns (1982) did not mention the peat swamp formation. Whitmore (1984a) recorded that peat swamp forests only occur as small patches mixed within more extensive freshwater swamps throughout New Guinea (Reynders 1962; Jacobs 1974). Whitmore (1984a,b) noted that there is no record that these New Guinea peat swamps have domed surface and the correlated complex of forest types found in other peat swamps in west Malesia. Since 1990 the extent of peat swamp forest in Papua has been mapped from satellite imagery, backed by limited ground survey. Peat formation begins at the inland edge of mangroves. Fine sediments carried down by the rivers are trapped behind the tangled mangrove roots and build up new land. As the coastline advances seawards due to sedimentation from erosion in the mountains, tidal inundation is less frequent, and the landward margin of the mangrove becomes less saline. Other plant communities develop in their wake. Because of the tidal influence and the backing up of freshwater in estuaries, the water table is high, and the soil is permanently waterlogged. The micro-organisms that would normally decompose falling plant debris are unable to survive in the anaerobic conditions, so that partly decomposed organic matter builds up to form a layer of peat over the mangrove clay soil. Rivers deposit alluvium along their banks, forming raised levees and creating backswamps. Cut off from the river water by the levees, the peat receives most of its water input from rainfall. Peat

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

CH50

03-15-07 07:34:33

PS

PAGE 933

934 / robert j . johns , garry a. shea, & p ratito puradyatmika

swamp forest species replace species which are characteristic of the mangrove formation. Since the peat swamps of Papua have been little studied, the following general description of the ecology of peat swamps is based on the peat swamps of Kalimantan, which have been extensively studied. MacKinnon et al. (1996) provides the following description of peat swamp forest and the soil. A peat soil is one with 65% or more organic matter content. The large peat deposits found behind coastal mangrove forest in Kalimantan and deltaic areas of Sarawak and Brunei are ombrogenous (rain-fed) peat swamps. The surface of the extensive lowland peat swamps is markedly domed and is not subject to flooding. The peat deposits are usually at least 50 cm thick, but they can be very deep, and depths of up to 20 m have been recorded. The surface of the peat is a solid, fibrous, and sometimes soft crust overlying a semi-liquid interior that contains large pieces of wood and other vegetable remains. Because most of the incoming water is from rain, it is extremely deficient in mineral nutrients. The peat and its drainage water are very acidic (with a pH of 4 or less) and poor in nutrients (oligotrophic), especially calcium. The many small rivulets that drain the peat swamp are tea-colored, and are described as ‘‘black-water’’ rivers. The vegetation occurring in peat swamps form a continuous sequence from open water to tall mixed swamp forest. Vegetation cover is dependent on the depth and quality of water, and drainage and flooding conditions. In relatively deep water, plant growth begins with communities of free-floating aquatics. As the water becomes less deep, rooting water plants are able to establish themselves. Stagnant water becomes dominated by herbaceous communities of mainly sedges, herbs, and ferns, while grasses predominate in swamps with moving water. Shrubs and trees appear in shallower swamps, resulting in various woodland communities differing in height and density. Swamp forest is the final stage in the sequence. Adjacent to herbaceous swamps, where the water table is permanently near the surface, open sago communities have a ground cover of shrub Pandanus, Hanguana malayana, sedges, or Phragmites karka. Some common dominants are Metroxylon sagu-Pandanus, Metroxylon sagu-Hanguana malayana, Metroxylon saguScirpus grossus, and Metroxylon sagu-Phragmites karka. Where the water table is well below the surface for at least part of the year, the ground layer is mainly composed of grasses, gingers, and ferns. Some of the more common communities are dominated by Metroxylon sagu-Stenochlaena and Metroxylon sagu and Nephrolepis. Most of the tree families of lowland evergreen forest are found in swamp forest. In the peripheral mixed swamp forest, where drainage is best, species composition is similar to that of lowland forest. There are very few plant species endemic to peat swamp forests; this may be because of the relatively recent origin of this habitat, which is probably less than 11,000 years old (Muller 1965).

mixed lowland peat swamp forest Mixed peat swamp forest communities are extensive in south Papua. The canopy is generally rather open, but may occasionally be rather dense. Under favorable

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

CH50

03-15-07 07:34:33

PS

PAGE 934

Lowland Swamp and Peat Vegetation of Papua / 935

conditions, the canopy may reach 30 m in height. The canopy is often of uniform height with few emergents. Some of the common trees found in the canopy are Alstonia scholaris, Terminalia canaliculata, Nauclea coadunata, Palaquium, Syzygium, Intsia, and Campnosperma. Generally, the lower tree stratum is open. Common lower canopy trees include Alstonia spatulata, Barringtonia, Diospyros, Garcinia, and Gynotroches. Pandanus and sago palms commonly form a substratum. Where the pandans and sago palm form a dense substratum, there is no undergrowth. Elsewhere, the density of the shrub and herb layer appears to vary depending on light conditions and flooding. Open stands may have a dense ground layer of Hanguana malayana and tall sedges. Thin lianas, fleshy climbers, and climbing ferns often thickly cover trunks. Where flooding is prolonged and deep, there are generally few shrubs and herbs. Generally, buttresses are not conspicuous, but stilt roots, sprawling surface roots, loop roots, and knee-shaped roots abound. Since many of these communities are permanently or seasonally inundated with water; they cannot develop into climatic climax communities, and the most advanced types are referred to here as edaphic climax communities. The dominant species vary among sites and include Alstonia, Campnosperma, Nauclea, and Syzygium; Campnosperma, Intsia, Nauclea, and Palaquium; or Syzgium, Alstonia, Palaquium, and Garcinia. Mixed peat swamp forests occur near the Minajerwi River in south Papua. One stand was sampled at the east end of the cross levee road at Biodiversity Research Site No. 3. The dominant canopy genera/species were Vatica russak, Stemonurus, Terminalia complanata, T. copelandii, Campnosperma brevipetiolata, Linociera, Intsia bijuga, Palaquium, Cerbera, Dillenia alata, and Myristica. Other common taxa were Mallotus, Syzygium, and Hopea novoguineensis. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 21.0 m2/ha and the total volume was 169.7 m3/ha. A second stand was located behind the levee, east of the Minajerwi River which flows through peat swamp land. Deep peat has not accumulated in these backwaters. Dominant genera were Pimelodendron, Vatica russak, Sloanea, and Myristica. Other trees included Pimelodendron, Vatica, Sloanea, Myristica, Elmerrillia, Aglaia, and Syzygium (Table 5.7.14). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 40 m2/ha. The estimated total volume was 438 m3/ha. East of the Minajerwi River south of junction with Kopi River in south Papua, Shea et al. (1998) established Biodiversity Research Site No. 4 on the higher and less frequently flooded river banks of a river flowing through swamps. Peat had not accumulated on the river banks. The dominant genera included Pometia, Celtis, Octomeles, and Syzygium. Other common trees (Table 5.7.15) included Artocarpus, Canarium, Vatica, Tetrameles, Aglaia, Myristica, Homonoia, and Cryptocarya. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 36 m2/ha. The estimated total volume was 346 m3/ha. Some stands of swamp forest have a substratum of Pandanus rather than sago palms. The dominant genera are Pandanus, Nauclea, Cryptocarya, Palaquium,

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

CH50

03-15-07 07:34:34

PS

PAGE 935

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

CH50

6.59 1.47 0.79

Campnosperma sp.

Canarium sp.

Cerbera sp.

03-15-07 07:34:34

2.80 1.38 0.28 1.37 5.28 0.95

Garcinia sp.

Glochidion sp.

Gnetum gnemon

Haplolobus sp.

Harpullia javanensis

Elmerrillia javanica

Flindersia sp.

14.42 51.70

Diospyros sp.

0.32

4.22

Campnosperma inertia

Cryptocarya sp.

0.18

Buchanania sp.

2.79

7.47

Alstonia sp.

Chisocheton sp.

40.70

Aglaia sp.

Species

Total clear bole volume (m3/ha)

PS

5

5

5

5

10

5

5

20

5

15

5

15

5

5

5

5

15

Absolute density

0.86

0.86

0.86

0.86

1.72

0.86

0.86

3.45

0.86

2.59

0.86

2.59

0.86

0.86

0.86

0.86

2.59

Relative density (%)

0.11

0.63

0.25

0.06

0.21

0.29

3.69

1.28

0.06

0.50

0.11

0.21

0.63

0.40

0.04

0.63

3.13

Absolute dominance (m2/ha)

0.27

1.57

0.62

0.15

0.52

0.72

9.18

3.18

0.15

1.24

0.27

0.52

1.57

1.00

0.10

1.57

7.79

Relative dominance (%)

Table 5.7.14. Forest dominated by Pimelodendron, Vatica, Sloanea, and Myristica

0.1

0.1

0.1

0.1

0.2

0.1

0.1

2.0

0.1

0.2

0.1

0.2

0.1

0.1

0.1

0.1

0.2

Absolute frequency

0.98

0.98

0.98

0.98

1.96

0.98

0.98

19.61

0.98

1.96

0.98

1.96

0.98

0.98

0.98

0.98

1.96

Relative frequency (%)

2.12

3.41

2.46

1.99

4.21

2.56

11.02

26.24

1.99

5.79

2.12

5.07

3.41

2.84

1.94

3.41

12.33

Importance value

936 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 936

0.43

4.20 0.72 45.56 15.11

Litsea sp.

Macaranga sp.

Myristica sp.

Palaquium sp.

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

CH50

03-15-07 07:34:34

PS

Source: Shea et al. (1998).

Total

Xanthophyllum sp. 437.89

7.64

46.39

0.69

Tabernaemontana sp.

Vatica sp.

20.66 13.66

Teijsmanniodendron sp.

0.13

Sterculia sp.

Syzygium sp.

57.29

Sloanea sp.

13.14

2.88

Planchonella sp.

Pometia sp.

45.98

Pimelodendron sp.

0.45

0.50

Ixora sp.

Pentaphalangium sp.

3.96

15.79

Intsia sp.

Intsia palembanica

Homonoia sp.

580

50

35

5

35

55

5

20

15

10

65

10

35

65

15

5

5

5

5

5

100.00

8.62

6.03

0.86

6.03

9.48

0.86

3.45

2.59

1.72

11.21

1.72

6.03

11.21

2.59

0.86

0.86

0.86

0.86

0.86

40.19

1.21

4.22

0.14

1.80

2.10

0.05

4.18

1.40

0.39

4.48

0.09

1.58

4.03

0.17

0.35

0.14

0.35

1.19

0.09

100.00

3.01

10.50

0.35

4.48

5.23

0.12

10.40

3.48

0.97

11.15

0.22

3.93

10.03

0.42

0.87

0.35

0.87

2.96

0.22

10.2

0.6

0.4

0.1

0.5

0.8

0.1

0.2

0.3

0.2

0.7

0.2

0.6

0.7

0.3

0.1

0.1

0.1

0.1

0.1

100.00

5.88

3.92

0.98

4.90

7.84

0.98

1.96

2.94

1.96

6.86

1.96

5.88

6.86

2.94

0.98

0.98

0.98

0.98

0.98

300.00

17.51

20.46

2.19

15.42

22.55

1.97

15.81

9.01

4.66

29.22

3.91

15.85

28.10

5.95

2.71

2.19

2.71

4.8

2.07

Lowland Swamp and Peat Vegetation of Papua / 937

PAGE 937

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2.01 1.16 7.85 0.98 1.32 6.70

Cryptocarya sp.

Ficus sp.

Gnetum gnemon

Homonoia sp.

43.54

Celtis sp.

Crudia sp.

20.61

Celtis latifolia

Chisocheton sp.

1.98

3.57

Calophyllum sp. 12.93

1.63

Buchanania sp.

Canarium sp.

18.00

Artocarpus sp.

Cananga sp.

18.99

Aglaia sp.

Species

Total clear bole volume (m3/ha)

03-15-07 07:34:35

PS

25

25

17

25

8

8

25

8

58

8

17

17

42

8

Absolute density

3.86

3.86

2.62

3.86

1.23

1.23

3.86

1.23

8.95

1.23

2.62

2.62

6.48

1.23

Relative density (%)

1.00

0.24

0.21

0.72

0.24

0.26

4.89

1.64

1.79

0.24

0.67

0.26

2.16

1.51

Absolute dominance (m2/ha)

2.75

0.66

0.58

1.98

0.66

0.72

13.45

4.51

4.92

0.66

1.84

0.72

5.94

4.15

Relative dominance (%)

0.50

0.33

0.33

0.33

0.17

0.17

0.33

0.17

0.50

0.17

0.33

0.33

0.50

0.17

Absolute frequency

Table 5.7.15. Swamp forest dominated by Pometia pinnata, Celtis, Octomeles sumatrana, and Syzygium

5.44

3.59

3.59

3.59

1.85

1.85

3.59

1.85

5.44

1.85

3.59

3.59

5.44

1.85

Relative frequency (%)

12.05

8.11

6.79

9.43

3.74

3.80

20.90

7.60

19.32

3.74

8.06

6.93

17.86

7.24

Importance value

938 / robert j . johns , garry a. shea, & p ratito puradyatmika

PAGE 938

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6.73 46.81

Planchonella sp.

Pometia pinnata

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15.83 0.37 5.42

Tetrameles nudiflora

Tetraprasandra sp.

Teijsmanniodendron sp.

Source: Shea et al. (1998).

Total

331.16

14.52

31.75

Syzygium sp.

Vatica russak

2.97

Pygeum sp.

4.04

2.11

Prainea sp.

1.34

Octomeles sumatrana

Pimelodendron sp.

10.68 46.36

Myristica sp.

Pangium sp.

1.24

Linociera sp.

0.70

5.61

Ixora sp.

Palaquium sp.

7.93

Intsia bijuga

648

25

17

8

8

42

17

17

117

8

8

17

8

8

58

8

8

8

100.00

3.86

2.62

1.23

1.23

6.48

2.62

2.62

18.02

1.23

1.23

2.62

1.23

1.23

8.95

1.23

1.23

1.23

36.35

1.78

0.73

0.07

1.51

2.93

0.52

0.68

5.44

0.80

0.38

0.29

0.17

3.68

1.43

0.15

0.80

0.94

100.00

4.90

2.01

0.19

4.15

8.06

1.43

1.87

14.97

2.20

1.05

0.80

0.47

10.12

3.93

0.41

2.20

2.59

9.19

0.50

0.33

0.17

0.17

0.50

0.17

0.33

1.00

0.17

0.17

0.17

0.17

0.17

0.83

0.17

0.17

0.17

100.00

5.44

3.59

1.85

1.85

5.44

1.85

3.59

10.88

1.85

1.85

1.85

1.85

1.85

9.03

1.85

1.85

1.85

300.00

14.20

8.22

3.28

7.24

19.98

5.90

8.09

43.90

5.29

4.13

5.27

3.55

13.21

21.92

3.50

5.29

5.67

Lowland Swamp and Peat Vegetation of Papua / 939

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Campnosperma, and Syzygium. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 15 m2/ha and the total volume was 63 m3/ha (Table 5.7.16). Where river banks are higher and less frequently flooded, the area behind levees may be dominated by seasonal swamp forest. These sites tend to lack the deep peat found away for the river. These forests have a canopy structure that is similar to climax dryland forest, but differs in species composition and ground cover. Different sites are dominated by different species, and the community is classified based on the dominant species present at each site. One community located behind the east bank of the Minajerwi River was dominated by Aglaia, Elmerillia tsiampaca, Syzygium, Myristica, Palaquium, Pimeleodendron, Pometia, Sloanea, Teysmanniodendron, Vatica, and Xanthophyllum. Where river banks are higher and less frequently flooded, young forests dominated by Artocarpus altilis and Octomeles sumatrana develop. These species may grow in pure stands, or mixed together. In this early successional stage, herbaceous and thin woody climbers are abundant, and often cover the trees. Where flooding is frequent, the ground tends to be relatively bare. Elsewhere, there may be a dense ground cover of ferns, gingers, and grasses.

Aquatic Communities Common submerged species, which include Blyxa spp., Hydrilla verticillata, Ceratophyllum spp., and the algae Chara spp., Lychnothamnus barbatusm, and Nitella spp., can form single-species communities or they may occur in mixed-species communities. Communities dominated by a single species include: Blyxa, Hydrilla, Ceratophyllum, Chara, Lychnothamnus, and Nitella. Submerged communities can also be mixed in species composition. Enhalus acoroides is frequent along river mouths on the north coast of Papua, but is absent where rivers deposit large amounts of silt, such as along the southern coast of Papua (van Royen 1963). Free-floating species are dominated by Lemna, Spirodela polyrhiza, Pistia stratiotes (introduced), Utricularia, Hydrocharis dubia, Ludwigia adscendens, Eichhornia crassipes, and Azolla pinnata. These may also grow in mixed communities. Rooted aquatic communities are found where water depths are less than 3 m. Many of these rooted water plants have submerged stems, but leaves that float on the surface of the water. These species include Nymphoides aurantiaca, Nymphoides exilifolra, Nymphoides germinata, Nymphoides indica, Nelumbo nucifera, Nymphaea nouchali, Nymphaea pubescens, and Nymphaea violacea. These may grow in mixed communities, or may grow in a mosaic in which some species form single-species communities. Ipomoea aquatica often grows in dense beds along the open water edge of swamps. The stems and floating leaves of Ipomoea are an important green vegetable consumed by indigenous people. Distinct herbaceous communities can grow up to 2.5 m above the level of the dark-colored water. Unlike other aquatic communities, they are rooted in a partly floating mat of waterlogged peat and organic debris. Common species occurring

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0.65 5.38 7.03 3.38

Endospermum sp.

Garcinia sp.

Intsia sp.

Litsea sp.

03-15-07 07:34:36

PS

Source: Shea et al. (1998).

59.91

7.15

Diospyros sp.

99.81

7.59

Campnosperma sp.

Total

0.36

Alstonia sp.

Metroxylon sp.

8.16

Aglaia sp.

Species

Total clear bole volume (m3/ha)

235

155

5

5

10

15

10

20

5

10

Absolute density

99.83

66.01

2.14

2.14

4.21

6.35

4.21

8.42

2.14

4.21

Relative density (%)

24.19

19.62

0.40

0.63

0.61

0.14

1.02

0.98

0.06

0.73

Absolute dominance (m2/ha)

99.94

81.10

1.65

2.60

2.52

0.55

4.21

4.05

0.25

3.01

Relative dominance (%)

2.4

1.0

0.1

0.1

0.2

0.3

0.1

0.4

0.1

0.1

Absolute frequency

99.78

41.68

4.17

4.17

8.20

12.37

4.17

16.68

4.17

4.17

Relative frequency (%)

Table 5.7.16. Swamp forest dominated by Nauclea, Cryptocarya, and Palaquium, with a Pandanus substratum

300.00

188.79

7.96

8.91

14.93

19.27

12.59

29.15

6.56

11.39

Importance value

Lowland Swamp and Peat Vegetation of Papua / 941

PAGE 941

942 / robert j . johns , garry a. shea, & p ratito puradyatmika

in these communities include the sedges Cyperus spp., Thoracostachyum sumatranum, Eleocharis dulcis, Scirpus spp., and Scleria sp.; the tall, fleshy, broad-leaved herb Hanguana malayana; and Typha orientalis. The ferns Stenochlaena spp., Nephrolepis spp., Ceratopteris thalictroides, Ampelopteris prolifera, and Cyclosorus interruptus can be locally common. These plants may also grow in mixed communities, or form a mosaic of single-species communities. On the clay plains in south Papua west of Merauke, Tecticornia cinerea forms distinct communities. Both Tecticornia cinerea and Batis argillicola grow in a narrow belt on clayey soils, periodically flooded in the rainy season but also influenced by freshwater from rivers (van Royen 1956).

grass and sedge swamps Archbold et al. (1942) discussed a variety of grass swamps, the grasses often associated with Adina, Barringtonia, Timonius, and Dillenia (Wormia). Van Royen (1956, 1963) also discussed swamp communities in southeast Papua, naming the main representatives as Phragmites karka and Scleria orzoides. Hanguana malayana was another conspicuous species forming an extensive vegetation, often also with Nephrolepis. Recent studies of these communities have been undertaken by Shea et al. (1998). Sedge communities are widely dispersed in southern Papua and can be dominated by several sedges, including Thoracostachyum sumatranum, Cyperus cephalotes, Cyperus imbricatus, Cyperus platystylis, Eleocharis dulcis, Scleria, Scirpus grossus, Scirpus litoralis, Scirpus mucronatus, and a community with mixed species. Herbland communities on peat include Hanguana malayana (probably not this species but specific limits are poorly known), Typha orientalis and a community of mixed species (sedges and forbs). Swamp communities can also be dominated by different species of ferns including the following: Stenochaena, Nephrolepis, Ceratopteris thalictroides (an edible species used as a green vegetable), Ampelopteris prolifera, and Cyclosorus interruptus, all of which also occur in communities of mixed-species composition. Medium-height swamp grassland communities include floating island grassland communities in wet peat swamps, swamp margin communities, and river bank and stream-side communities. Common medium-height grass species occur in floating island grassland communities in wet peat swamps and along swamp margins. Common species of medium-height grasses that are dominant in communities include Echinochloa praestans, Hymenachne acutigluma, Ischaenum polystachyum, Leersia hexandria, Brachiaria mutica, Panicum auritum, and Panicum paludosum. These may grow in mixed communities, or may grow in a mosaic in which some species form single-species communities. The species occurring in these mixed-species grassland communities varies at different sites. Tall grassland communities in peat swamps grow in brackish environments along the lower reaches of the Minajerwi near the boundary with tidal swamps. The following communities are common on frequently flooded river banks:

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Lowland Swamp and Peat Vegetation of Papua / 943

Phragmites karka dominated brackish-water riparian peat swamp, and Pandanusdominated brackish-water riparian peat swamp. Further upstream, the freshwater environment of frequently flooded low levee banks is dominated by Phragmites karka on lower sites and Saccharum robustum on higher sites. Tall swamp grasses, mainly Phragmites karka, grows in swamps that are shallower than those occupied by mid-height swamp grass. P. karka grows under a range of conditions, from permanent swamps to sites that are dry for several months. It grows in both stagnant and moving water, and in fresh- and brackish-water environments. On wet sites, Phragmites karka tends to form a single-species community. On drier sites, the Phragmites karka communities are more mixed, with herbs and ferns growing at the base of the tall grass. The associated ferns are often Pteridium auriculum and Asplenium spp. Drier sites tend to have an alluvial substrate, while wet sites often have a peat substrate. These drier sites tend to be invaded by pioneer tree species, and are converted into woodlands and eventually into forests, which are described in subsequent chapters. Saccharum robustum may form pure stands on relatively high river levees which are subject to frequent but only brief flooding. This vegetation type occurs on river bank alluvium. Tall swamp grasses, mainly Phragmites karka, grow in areas that are intermittently dry. Shrubs and low trees are widely scattered within this community. Some common shrubs and low trees are Glochidion, Nauclea, Mitragyna, Ficus, Macaranga, and Casuarina. This community generally occurs on alluvium rather than peat soils.

Acknowledgments We wish to express our sincere thanks to Dr. Wim Vink for his extensive comments and additions to this chapter. The careful comments of the editors were also appreciated.

Literature Cited Anderson, J.A.R. 1963. The flora of the peat swamp forests of Sarawak and Brunei including a catalogue of all recorded species of flowering plants, ferns and fern allies. Gdns. Bull. Singapore 20: 131–228. Archbold, R.A., A.L. Rand, and L.J. Brass. 1942. Results of the Archbold Expedition to New Guinea, No. 41. Summary of the 1938–1939 New Guinea Expedition. Bull. Am. Mus. Nat. Hist. 288, 35 plates. Brunig E.F. 1973. Species richness and stand diversity in relation to site and succession of forests in Sarawak and Brunei (Borneo). Amazoniana 4: 293–320. Dick, J. 1991. Forest Land Use, Forest Use, Forest Use Zonation and Deforestation in Indonesia: A Summary and Interpretation of Existing Information. Ministry of Population and Environment (KLH) and the Environmental Impact Management Agency, BAPEDAL, Jakarta. Jacobs, M. 1974. Panorama Botanique de l’Archipel Malais (Pltes. Vasc.). Ressources

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PAGE 943

944 / robert j . johns , garry a. shea, & p ratito puradyatmika Naturelles de l’Asie Trop. Humide, UNESCO. Recherches sur les Ress. Nat. XII, 285–320. Johns, R.J. (1982). Plant zonation. Pp. 309–330 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Kluwer Academic Publishers Group, Dordrecht, The Netherlands. Lundquist, E. 1942. Verslag van een bosexploratie in Nieuw Guinea in de Vogelkop en lans de Zuid-West kust tot de Bloemenrivier. Mei–Augustus, 1941, Batavia, Dienst Landbouw en Visserij, afdeling Boswezen; mimeo. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO in association with Australian National University, Canberra. RePPProT. 1986. Review of Phase I and II Results for Irian Jaya. Regional Physical Planning Programme for Transmigration (RePPProT), Ministry of Transmigration, Jakarta. Reynders, J.H. 1962. Shifting cultivation in the Star Mountains area. Nova Guinea, Anthropology 10 (3): 45–73. Shea, G.A., D. Martindale, P. Puradyatmika, and A. Mandessy. 1998. Biodiversity Surveys in the PT Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. PT Freeport Indonesia. van Royen, P. 1956. Notes on the vegetation of clay-plains in Southern New Guinea. Nova Guinea NS 7: 175–180. van Royen, P. 1963. Sertulum Papuanum 7. Notes on the vegetation of Southern New Guinea. Nova Guinea Contrib. 35: 195–241. van Royen, P. 1965. An outline of the flora and vegetation of the Cycloop Mountains. Nova Guinea Botany 21: 460–463. Whitmore, T.C. 1984a. Tropical Rainforests of the Far East. Oxford Science Publications. Clarendon Press, Oxford. Whitmore, T.C. 1984b. Vegetation map of Malesia, at scale 1:5 million. Journal of Biogeography 11: 461–471.

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PAGE 944

5.8. Lowland Vegetation of Papua robert j. johns, garry a. shea, and pratito puradyatmika u r kn o w l ed g e o f t h e e co l o g y of the major lowland plant communities is superficial because there are few detailed studies in the Papuan lowlands. Much of the data included on plant ecology is extrapolated from studies that have been made in Papua New Guinea. The status of the Forestry Surveys in Papua is not known to the authors but they probably contain detailed floristic data on the composition of the forests. Most early ecological studies in Papua were compiled by botanists making detailed collections of the plants (Archbold et al. 1942; Brass 1941; Lam 1934, 1945; Ridley 1916; van Steenis 1957). In Papua New Guinea the lowland vegetation has been documented in the publications in the CSIRO Land Research Series (Robbins 1968; Paijmans 1967, 1970, 1976) but few quantitative data have been published. Paijmans (1970) analyzed the results from a survey of four 0.8 ha plots in rainforest in northeast Papua New Guinea. No similar studies are known from Papua. Vink (1998, p. 91) has emphasized that much information on the lowland rainforest of Papua are ‘‘hidden in typewritten or mimeographed internal reports of forest surveys (in Dutch) with a very limited distribution.’’ Shea et al. (1998) published the results of detailed surveys of the vegetation of the Freeport Contract of Work area in south Papua, which is the basis of this chapter. A bibliography of papers on plant ecology in New Guinea was published by Johns (1992). Results are also available from ecological studies on the lowland dipterocarp forests in Papua New Guinea (Johns 1982, 1987). As a cautionary note, two major points must be accepted when reading this chapter: our knowledge of the ecology of the vegetation of Papua is fragmentary, and the flora of Papua is poorly known (Chapter 3.1). This complicates ecological studies because many of the plant species collected in Papua could be new to science. The descriptions of vegetation types are superficial, as they are often based on field notes made on expeditions where the primary objective was the collection of taxonomic specimens. ‘‘Critical checklists’’ of the plant species occurring in the flora, a fundamental requirement for detailed ecological studies in any region, are largely absent. Although several species lists have been prepared for lowland vegetation in Papua New Guinea, such lists are seldom available in Papua. Exceptions are the lists for the Vogelkop (database at Kew and Leiden), and the lists published for the Cyclops Mts (van Royen 1965), and Waigeo Island (van Royen 1960), which were based on collections van Royen made during expeditions to those regions. Ridsdale (1968) includes a list of the species collected during the Border Demarcation Expedition to south Papua in 1967.

O

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

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Ecosystem Instability in Papua A paper on the instability of the tropical ecosystem in Papuasia (Johns 1986) identifies the five major aspects of ecosystem instability that apply in Papua. First, is earthquake damage, including landslides, tidal waves, and physical damage to rainforest as a result of tree falls, breaking of branches, and loosening of larger epiphytes. Second, El Nin˜o related phenomena, which occur regularly (ca every 7 years), are periods of droughts of varying intensity that have had a major ecological impact on the vegetation of Papua. When the El Nin˜o event occurs after several years of low rainfall, the consequences of such droughts can be catastrophic. Much of the lowland rain forest in New Guinea burned during the droughts of 1877–1878, 1888, 1891, 1902, 1914–1915, 1940–1942, 1982–1983, and more recently in 1991 (Vink 1998). Forest fires associated with these droughts have often burned hundreds of square kilometers of rainforest (similar to the fires in Kalimantan). The damage to savanna, swamp, and monsoon communities can also be extensive. Large areas of rainforest have also been burned (Johns 1986; Vink 1998). A third source of instability is flood damage. Very heavy rains can cause extensive local damage by undercutting stream banks, causing landslides (particularly in areas with mudstones and siltstones), and by destroying riverine vegetation along the banks of major rivers by altering river courses, destroying extensive riverine forest communities. Oxbow lakes are created (and destroyed) during such floods. Fourth is lightning damage. Most areas in Irian Jaya (as in Papua New Guinea) are subject to many lightning strikes. Lightning plays a major role in mangrove ecology by destroying large patches of mangroves; these dead patches are easily visible from aerial photographs (Johns 1986). Similar patches occur in stands of Cocos nucifera and probably in montane forests; possibly the gaps occurring in Nothofagus forest are due to lightening strikes. Cyclones are another major cause of fires in natural communities, not only in savanna and rainforest, but also in dry mangrove communities. A fifth source of ecosystem instability is wind damage (local cyclones). No reports of cyclones have come from Papua. However, local cyclonic events appear common. The Agathis labillardieri forests on Yapen Island occupy narrow elongate strips which are very similar to events observed in lower montane Castanopsis acuminatissima forest and mixed montane forest in Papua New Guinea, which are thought to be the result of earlier local cyclones. Papua has no volcanoes and consequently, unlike Papua New Guinea, there has been no extensive damage caused by volcanic eruptions, such as occurred with the eruptions of Mt Lamington in 1953 and the extensive destruction that occurred during the last eruption of Long Island. This has several consequences for Papua because the soils, lacking volcanic ash, tend to be poor and are not be able to support large migrant human populations. The only source of volcanic ash has been the thin ash beds along the Papuan border, resulting from volcanic eruptions

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Lowland Vegetation of Papua / 947

of Long Island (in Papua New Guinea) some 300 years ago. Like Kalimantan, but unlike Java which is also volcanic, the soils throughout Papua are generally of low nutritional quality. Population density is lower in Papua than Papua New Guinea because the soils of Papua are poorer than those of PNG. Therefore the effects of humans on Papuan lowland vegetation are relatively limited. The intensity of human impacts on Papua lowland forests could change catastrophically and irreversibly in the near future if the population continues to increase and the forests continue to be harvested unsustainably.

Lowland Tropical Rainforest The plant associations that make up the tropical lowland rainforest in Papua are some of the most complex and least understood of the earth’s vegetation types. The lowland forests are very diverse in species composition, with in excess of 1,500–2,000 tree species in over 80 plant genera, and an unknown number of subcanopy species, parasitic and epiphytic plants, ground herbs, and so on. Many plant species are local endemics. The reasons for the local distribution patterns of particular species are not understood. In contrast to the flora of Borneo, the distribution and species composition of the subcanopy and ground flora seems to be very local in New Guinea. Collections from areas less than 1 km apart appear to support quite different herbaceous and shrub floras, but the canopy trees can be the same. The rainforests are often composed of trees with large root buttresses, supporting trees that reach some 45 m in height. The canopy surface is often very uneven and species of Ficus often occur as emergent species. Several species of Ficus start as epiphytic shrubs growing on the large branches in the canopy. Gradually their roots encase the trunks of their host trees, and the figs outlive their host. Many areas of lowland rainforest have been destroyed by instability caused by the sources mentioned above. The lowland forests dominated by Intsia (I. palembanica and I. bijuga) and Pometia pinnata are indicators of earlier fires. Lowland rainforest occurs in the everwet areas of Indonesia. Mapping of the forests was done by RePPProT (1986, 1990). Dick (1991), based on RePPProT (1990), estimated the areas of dryland lowland rainforest in Indonesia (Table 5.8.1). Papua has 176,750 km2 of lowland tropical rainforest, which is 26.5% of the total for Indonesia. Tropical lowland rainforest is the most complex and species-rich terrestrial vegetation type in the world (Whitmore 1984a,b). The forest canopy provides the framework in which other kinds of plants grow, including climbers, epiphytes, parasites, and saprophytes. The main tree families are the Anacardiaceae, Annonaceae, Apocynaceae, Burseraceae, Clusiaceae, Combretaceae, Dipterocarpaceae, Ebenaceae, Elaeocarpaceae, Euphorbiaceae, Leguminosae, Meliaceae, Moraceae, Myristicaceae, Myrtaceae, Rubiaceae, Sapotaceae, Sapindaceae, and Sterculiaceae. This lowland forest contrasts with the montane zone where the Araucariaceae, Fagaceae (including Nothofagaceae), Podocarpaceae, Cupressaceae, and Lauraceae are usually important. The mature forest growing in the alluvial valleys and on the

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Table 5.8.1. Area of lowland tropical rainforest in Indonesia by region Region

Area of lowland forest (km2)

Sumatra

120,734

Kalimantan

270,216

Sulawesi

57,362

Nusa Tenggara

6,969

Maluku

34,743

Papua

176,750

Total

666,744

Source: Shea et al. (1998).

alluvial fans are considered the ‘‘climatic climax’’ vegetation for Papua. Paijmans (1976) described these forests as follows: tall, floristically and structurally very rich, mixed alluvium forest is the most luxuriant of the lowland rainforests in New Guinea. The forest structure is irregular throughout all layers. The canopy is very variable in height, closure, and in crown sizes, features that are particularly striking from the air. It is rather open and has many gaps in which only lower trees are present. Generally the canopy is about 30–35 m high, but irregularly scattered trees emerging partly or fully above the canopy may reach 50 m or more; many figs are emergents. Most emergents and many canopy trees have wide crowns, tall straight boles, and high and wide buttresses, some trees attaining a girth well over 25 m. Tree species constantly present in the upper stories are Pometia pinnata, Ficus spp., including strangling figs, Alstonia scholaris, and Terminalia spp. The lower tree strata are usually rather open, as the number of trees below the canopy is relatively low. Some trees with stilt roots are invariably present. Typical lower story trees are Garcinia, Diospyros, Myristica, Maniltoa, and Microcos. The cover and density of the shrub and tall herb layer vary with the amount of light penetrating through the canopy, and with the types of shrubs present. A predominance of spreading, multi-branched shrubs results in a high cover, whereas a dense layer of slender saplings may have only a low cover. Palms are a feature of the shrub layer. They consist of small tree palms, and include the common Licuala with its conspicuous fan-shaped leaves. Tall gingers and Marantaceae locally form a dense layer. Pandans are rarely common, and tree ferns, in contrast to heath forest, and bamboo are scarce. The low herb layer forms an irregular and patchy ground cover. It may be almost absent, as where shrub palms are abundant, or quite dense, as where it is formed by Selaginella, Elatostema, Marantaceae, or Commelinaceae. Otherwise the herb layer consists mainly of ferns, tree and rattan seedlings, and some forest grasses and sedges. Thin and thick woody lianas, fleshy epiphytic climbers, and climbing ferns are usually common. Climbing rattan is invariably present, but is dense only below canopy openings. Epiphytic ferns and orchids are plentiful in the crowns of canopy trees, especially in old trees with open crowns,

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thick branches, and a rough bark. Unlike those in some high mountain forests, most orchids have inconspicuous flowers. The natural forest is structurally a complex mosaic of gap, building, and mature phase stands. Since cyclones are absent or infrequent in Papua, rainforest gaps are usually small and the result of the death of individual overmature trees. When these trees eventually fall, they produce a gap. Pre-existing seedlings tend to grow rapidly in these relatively small gaps. Species composition varies from site to site, and the community at any particular location can be defined by the dominant species that occur on the site. Since the flora contains hundreds, perhaps thousands, of tree species, there are an infinite number of possible combinations. The canopy is often dense in the lowland forest, allowing little light penetration into the lower layers of the forest, resulting in a surprisingly sparse undergrowth despite the apparent luxuriance of the forests. Despite the low light intensity, terrestrial floras in the forest are composed of many species including the ferns, particularly in the family Thelypteridaceae, many species in the rubiaceous genera, and a host of other plant genera and species. Many epiphytic species occur in the lowland rainforest including epiphytic ferns such as Lecanopteris and Platycerium wandae, rubiaceous ant plants in the genera Myrmecodia and Hydnophytum, and many epiphytic orchids such as the massive Grammatophyllum speciosum and species of Dendrobium including D. antennatum, D. spectabile, and D. bracteosum. Among the many climbers are the species of Mucuna, which are a major feature of the lowland tropical rainforest in New Guinea. Many climbers in the genus Gnetum occur in the forests. Lam (1945) described the lowland rainforest along the Mamberamo, including notes on the many species of ferns and orchids, which were subsequently described by J.J. Smith. Van Royen recognized several types of lowland rainforest on Waigeo Island. This included mixed forest with Pometia, Alstonia, and so on; Vatica-Dillenia forest on lateritic loamy soil with underlying limestone or ultrabasics; Agathis is dominant in some forests and Intsia-Pinanga forest grows on coastal limestone cliffs with thin clay soils. Decussocarpus wallichianus is very common on bare limestone soils. Riverine forests dominated by Myristica, Ficus, Syzygium, and Pometia occur along the major rivers. Octomeles sumatrana can form pure stands. In the Cyclops Range, lowland rainforest is found on both metamorphic and ultrabasic rocks. The forest on metamorphics is dominated by Pometia pinnata, with the usual lowland trees: Intsia bijuga, Anisoptera, Dillenia, Dracontomelum, Firmiana, Haplolobus, Myristica, Pandanus, and Pleiogynium, with a large species of Sloanea common on drier ridge crests. A similar forest occurs on the alluvial soils that form a belt to the south of the Cyclops Mountains, bordering Lake Sentani. Ridsdale (1968) includes brief descriptions of the vegetation along the border with Papua New Guinea, south of the Star Mountains. The main forest type that is widespread in south Papua was Vatica russak-Campnosperma montana forest. Ridsdale (1968) includes a list of the associated species based on their collections. Along ridges in south Papua the dominant forest trees were Eucalyptopsis papuana

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and Syzygium sp. Vatica russak and Hopea papuana form a distinct forest type in smaller valleys in the area. Vink (1998) has included information on the rainforest of the Vogelkop Peninsula. He includes details from various surveys by the Forestry Department. This includes survey results from the several areas. In the Tisa River and Maturi River survey the forests were dominated by Pometia pinnata and Intsia; the Sidei-Wariki alluvial forests were dominated by Pometia pinnata, Teysmanniodendron bogoriense, and Intsia. The forests on the floor of the Kebar Valley are similar to the forest of the Arfak Plain. Intsia bijuga and I. palembanica are common as are Pometia pinnata and Alstonia. Similar forest composition is reported by Vink (1998) from the Warsamson Valley. In the Momi-Ransiki area Vatica rassak is a very important component of the lowland forest, accounting for some 34.6% of the trees. In one strip a density of Vatica of 74.5% was recorded. The information contained in the Forestry Department reports from Papua needs detailed study.

riparian forest Riparian vegetation occurs along the river beds and on the banks of the major rivers to the north (Mamberamo River) and to the south of the main ranges. The rivers to the north are much longer than those to the south which drain the Central Ranges to the Arafura Sea. The forests on the river banks are often inundated during high floods. Riparian forests also include the alluvial forests which are sometimes flooded. Often the communities are destroyed by meandering rivers. There is seldom a sharp delimitation between these species and those of the swamp forests. River bed deposits, subject to flash flooding, are initially colonized by grasses which form scattered tussocks or grow in small stands, sometimes dominated by a single species: Cyperus, Saccharum spontaneum, and Pennisetum macrostachyum. Other grasses pioneer on well drained sites. These communities may initially be single species communities or mixed species communities. Wet sites on medium to coarse sediments in river beds are dominated by the grass Phragmites karka. As these wetland sites dry, they are initially invaded by ferns, including Pteridium sp. and Asplenium sp. As these sites dry further, trees and shrubs eventually invade. The pioneer tree species become established among the Phragmites and eventually form an open emergent layer over the tall Phragmites. Common pioneer trees include Glochidion, Ficus, Paraserianthes, and Casuarina. Many different woodland communities occur in the river beds depending on local site history and available seeds. Some recognized by Shea et al. (1998) include: Glochidion and Phragmites karka; Ficus and Phragmites karka; Paraserianthes and Phragmites karka; Casuarina and Phragmites karka; Glochidion, Ficus, and Phragmites karka; and more mixed communities dominated by Casuarina, Paraserianthes, and Phragmites karka. Islands and gravel bars in the river channels are colonized by herbs and pioneer species, particularly Casuarina. These pioneer communities create favorable conditions for other plant species to invade over time. The pioneer species are eventually replaced by successional species. Shea et al. (1998) sampled several stands

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located on islands (gravel and small boulders) in the river bed of the Aimoca River. Two stands were sampled by Shea et al. (1998) along the Aimoca River. These are dominated by Casuarina-Linociera-Campnosperma-Pisonia or Casuarina-Artocarpus-Pandanus. In the first stand the dominant plants were Casuarina equisetifolia, Linociera sp., Campnosperma montana, and Pisonia. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 15.0 m2/ha and the total volume was 129.1 m3/ha. The species Casuarina equisetifolia, Artocarpus sp., and Pandanus sp. were dominant in another stand located on small island in the river bed of the Aimoca River. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 11.7 m2/ha and the total volume was 101.8 m3/ ha. As the Casuarina trees mature, other trees species colonize the stand and the grasses and ferns are shaded out. The understory trees may include Alstonia spectabilis, Artocarpus sp., Campnosperma sp., Dillenia sp., Glochidion sp., Gnetum gnenom, Litsea sp., Linociera sp., Macaranga sp., Ficus sp., Myristica sp., Arenga sp., Pandanus sp., Pisonia sp., Prainea sp., Pterygota sp., Pygeum sp., and Saurauia sp. Casuarina dominated communities are common on medium and coarse textured materials in the meander belt of the Ajkwa River. One stand, located just south of the Ajkwa River bridge, was sampled. Dominant trees were Casuarina equisetifolia, Litsea, Pygeum, Pandanus, Alstonia spectabilis, Prainea, Linociera, Myristica, Glochidion, and Dillenia. The four dominant genera were Casuarina, Litsea, Pygeum, and Pandanus. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 19.0 m2/ha and the total clear bole volume was 156.7 m3/ha. Octomeles sumatrana is a common species forming pure stands along river banks and in poorly drained back swamps. Octomeles can grow up to 20 m in height in 4–5 years and the small seedlings become rapidly established in secondary conditions. The seeds of Octomeles rapidly loose their viability and after 2–3 days germination rates fall rapidly. In Papua New Guinea Octomeles are often established on volcanic ash surfaces following eruptions. Eucalyptus deglupta, a rare species in Papua, also functions as a river bank species on disturbed sites in Papua New Guinea. Riparian Pandanus setistylus is a common species on muddy banks in rivers at low altitude. These communities also grow in oxbow lakes. Saccharum robustum is common amongst the screwpalms.

River Bank Vegetation Frequently flooded river banks are dominated by Phragmites karka on lower sites and Saccharum robustum on higher sites. The wild breadfruit, Artocarpus altilis, is one of the first trees to appear on the tops of low river banks. They are mixed with grasses and other river bank herbs, climbers, and shrubs. Where river banks are higher and less frequently flooded, young forests dominated by Artocarpus altilis and Octomeles sumatrana develop. In this early successional stage, herbaceous and thin woody climbers are abundant, and often cover the trees. Where flooding is

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frequent, the ground tends to be relatively bare. Elsewhere, there may be a dense ground cover of ferns, gingers, and grasses. Over time, other early secondary forest trees such as Ficus, Laportea, Nauclea, Terminalia, and Kleinhovia become established in the understory, with the Octomeles sumatrana forming a tall emergent layer. The Octomeles sumatrana emergents may eventually reach 60 m in height, with large diameters and wide crowns. Octomeles sumatrana cannot regenerate under its own canopy, and is eventually replaced by other species until the mixed forest climax is reached. Alluvial valley forests are often dominated by Octomeles, Pometia, Elaeocarpus, and Anisoptera, or by Blumeodendron, Octomeles sumatrana, Dacryoides, and Garcinia.

Flood Plain Forest In south Papua three sites were assessed by Shea et al. (1998) on the Otomona River near Mile 38 Camp. On the flood plain adjacent to the river, Paraserianthes falcataria may grow to form a tall overstory over a medium main story composed of species such as Anthocephalus cadamba, Artocarpus, Octomeles sumatrana, Syzygium, Pometia, and various species of Ficus, including the strangling fig, F. benjamina. Other sites were dominated by Dysoxylum, Paraserianthes falcataria, Pometia, and Casuarina; Paraserianthes, Anthocephalus, Pometia, Psychotria forest, and a community dominated by Calophyllum, Macaranga, Anthocephalus, and Octomeles. A stand on the flood plain of the Aimoca River was dominated by Octomeles sumatrana, Pometia pinnata, Anisoptera thurifera, and Elaeocarpus sp. Other trees included Elaeocarpus sp., Alangium sp., Chisocheton sp., Garcinia sp., Syzygium sp., and Anthocephalus cadamba. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 36.4 m2/ha and the total volume was 368.5 m3/ha. On the flood plain adjacent to the Aimoca River the forest was dominated by Blumeodendron papuanum, Octomeles sumatrana, Dacryoides sp. Other trees include Garcinia sp., Dysoxylum sp., Garcinia sp., Syzygium sp., and Anisoptera thurifera. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 38.4 m2/ha and the total clear bole volume was 433.7 m3/ha. Paraserianthes falcataria may grow to form a tall overstory over a medium main story composed of species such as Anthocephalus cadamba, Artocarpus, Octomeles sumatrana, Syzygium, Pometia, and various species of Ficus, including the strangling fig, Ficus benjamina. Three such stands, located adjacent to the Otomona River near Mile 38 Camp, were sampled. The dominant genera/species in one of the stands are Dysoxylum, Paraserianthes falcataria, Pometia pinnata, Casuarina equisetifolia, and Anthocephalus cadamba. Alternatively, the dominant trees on river flood plains can be Paraserianthes falcataria, Anthocephalus cadamba, Pometia pinnata, Psychotria sp., Sloanea sp., Octomeles sumatrana, and Teijsmanniodendron bogoriense. The plant community is represented by the name of the four most dominant genera ParaserianthesAnthocephalus-Pometia-Psychotria. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 33.4 m2/ha and the total clear bole volume

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was 417.6 m3/ha. Another distinctive forest type is dominated by Calophyllum, Macaranga, Anthocephalus cadamba, Octomeles sumatrana, Xanthophyllum papuanum, and Platea latifolia. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 39.0 m2/ha and the total clear bole volume was 368.0 m3/ha.

valley floor and valley side forests The most frequent genera of trees are Pometia, Celtis, Pterocarpus, Ficus, Dracontomelum, Vitex, Sloanea, Cerbera, Alstonia, Maniltoa, Cryptocarya, Guioa, Kingiodendron, and Tristiropsis. The most common canopy species in many areas destroyed by El Nin˜o related forest fires is Pometia pinnata. Elaeocarpus and Intsia are also common dominants in disturbed areas. The subcanopy of these forests is very diverse and dominated by many genera including Diospyros, Gnetum, Protium, Pimeliodendron, Myristica, Dendrocnide, Kibara, and Horsfieldia. Common widespread species in these genera are Diospyros discolor, Gnetum gnemon, Protium macgregorii, and Myristica chrysophylla. Various species of feather and fan palms are common including Licuala, as are several species of Pandanus, particularly in areas with slightly wetter soils. Forests dominated by Vatica russak are very common in Papua. For example, at lower altitudes the forest of the Go Isthmus in Waigeo Island is dominated by this species.

ridge top forests There is a marked change in the floristic composition of forests on the welldrained ridge tops. The soils are more stable along ridges. The canopy of the ridge forests tend to more open, the subcanopy is dominated by species of Diospyros, Myristica, Gnetum, and Pimeliodendron. At higher altitude this forest shows a transition to the lower montane forests with the presence of scattered trees of Lithocarpus.

lowland rainforest in alluvial valleys The forest in alluvial valleys is tall and floristically very rich. This type of dryland alluvial forest is the most luxuriant of the rainforests of New Guinea (Paijmans 1976) and represents the climatic climax forest in Papua. The canopy is generally 30–40 m high, but scattered emergent trees can reach 50 m or more in height. Some trees reach girths of well over 2.5 m. Most emergent trees and many canopy trees have wide crowns, straight boles, and high buttresses. Woody climbers and bole climbers are common. Epiphytes are abundant, and include forbs, orchids, ferns, bryophytes, lichens, algae, and bacteria. The ground layer is composed mainly of seedlings and advanced growth of tree species, scattered shrubs, and herbs. The structure of the tall closed-forest is multi-layered (mainly 5-layered) with tall to very tall emergents over a closed main canopy, with understory, shrub layer, and ground cover. The height of the main canopy is 30–40 m; emergent trees are frequent, to 50 m tall; tree buttresses are common, including low, medium and

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high buttresses. Stilt roots are common; breathing roots (pneumatophores) occasional. Trees with flowers and fruit on trunk or main branches (cauliflory or ramiflory) are common. Dominant leaf-size class is mesophyll; trees with compound leaves are common to abundant; leaf drip-tips are abundant. Large and small woody climbers are both abundant. Vascular epiphytes are common to abundant; non-vascular epiphytes are present, sometimes common; bryophytes are scattered in the ground cover. Strangling figs are common. Shrub palms are common in shrub layer; bamboos are absent; shrubby pandans are occasional in the shrub layer; rattans are occasional. Tree ferns are rare or occasional. The dominant families include the Annonaceae, Apocynaceae, Burseraceae, Dipterocarpaceae, Ebenaceae, Fagaceae, Leguminosae, Meliaceae, Moraceae, Myrtaceae, and Sterculariaceae. Tree species consistently present in the understory are Pometia pinnata, Ficus spp., including strangling figs, Alstonia scholaris, and Terminalia. The lower tree strata are usually rather open, as the number of trees below the canopy is relatively low. Some trees with stilt roots are invariably present. Typical lower story trees are Garcinia, Diospyros, Myristica, Maniltoa, and Microcos. The dryland evergreen rainforest is similar to that which occurs on alluvial fan and plains. The dryland evergreen rainforest is the most floristically and structurally complex forest community. The dominant species vary from site to site. A community common in alluvial valleys, including stable river banks, is dominated by Ficus benjamina, Pometia, Intsia, and Syzygium. The forests which dominate the flood plain of lowland rivers such as the Aimoca River are dominated by late secondary forest tree species. Two stands located along the Aimoca were sampled. One stand was located on the flood plain of the Aimoca River. The dominant genera and species were Calophyllum sp., Macaranga sp., Anthocephalus cadamba, Octomeles sumatrana, Pometia pinnata, and Xanthophyllum papuanum, Platea sp., and Saurauia sp. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 39.8 m2/ha and the total volume was 417.9 m3/ha. A similar forest was studied on the flood plain opposite the Mile 38 Camp in south Papua. The major species in the plot were Canarium sp., Macaranga sp., Gonostylus macrophyllus, Dispyros maritime, and Ficus sp. Tall closed-forest communities on alluvial fans was assessed by Shea et al. (1998). The stand was located north of the road between bridges at approximately Mile 33. The forest was dominated by Myristica-Hopea-Teijsmanniodendron-Litsea. A similar forest stand located along the road at Kuala Kencana was different is species composition. The dominant genera at this site were Pometia-SyzygiumLitsea-Campnosperma. Near the Otomona Bridge similar forest was dominated by Pometia-Litsea-Aglaia-Syzygium. Forest communities near the Ajkwa Bridge at Mile 32/33 were Pometia-Syzygium-Intsia-Teijsmanniodendron; another at Kali Kopi at Mile 37 was dominated by Pometia-Alphitonia-Teijsmanniodendron and Alstonia. Another stand near a large strangling fig near Kali Kopi at Mile 37 was dominated by Ficus benjamina over Macaranga-Buchanania-Canarium-Timonius. The last two types of alluvial forest may have been selectively logged in the past, with one or two valuable trees removed per ha. The impact was minimal since the

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trees were cut into planks and boards in the forest and carried by hand to the roadside. The dominant trees in a stand located north of road between bridges at approximately Mile 33 along PTFI Road were Myristica, Hopea papuana, Teijsmanniodendron bogoriense, Litsea, Mastixiodendron, Pometia pinnata, Alstonia, Haplolobus, Homonoia javanensis, Hopea papuana, Chisocheton, and Syzygium. The four dominant genera were Myristica, Hopea, Teijsmanniodendron, and Pometia. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 34.3 m2/ha and the total clear bole volume was 340.2 m3/ha. This stand is located along road at the Light Industrial Park in Kuala Kencana. Dominant genera and species are Pometia pinnata, Syzygium, Litsea, Campnosperma, Aglaia, Chisocheton, Palaquium, Pimelodendron, Homonoia, Haplolobus, Nauclea, Macaranga, and Canarium. The most important genera and species based on combined Importance Value are Pometia pinnata, Syzygium, Litsea, Aglaia, Homonoia, Macaranga, Xanthophyllum, Teijsmanniodendron, Pimelodendron, and Myristica. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 29.1 m2/ha and the total clear bole volume was 304.4 m3/ha. A stand located 5 km south of the Otomona Bridge is dominated by Pometia, Litsea, Aglaia, Syzygium, Teijsmanniodendron, Pimelodendron, Chisocheton, Homonoia, and Polyalthia. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 15.0 m2/ha and the total clear bole volume was 304.4 m3/ha. A stand located adjacent to a forest stream south of the Ajkwa Bridge in south Papua was dominated by Pometia pinnata, Syzygium, Intsia, Teijsmanniodendron, Alstonia, Pimelodendron, Myristica, Hopea papuana, and Paraserianthes falcataria. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 36.6 m2/ha and the total clear bole volume was 334.4 m3/ha. A stand located near Kali Kopi, south Papua, had the dominant genera Pometia, Alphitonia, Teijsmanniodendron, and Alstonia. The important genera in the stand were Pometia pinnata, Archidendron clyperia, Teijsmanniodendron, Alstonia scholaris, Alphitonia, Homonoia javanensis, Intsia bijuga, Aglaia, and Campnosperma. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 20.7 m2/ha and the total clear bole volume was 179.4 m3/ha. Shea and colleagues established a plot in a stand at the site of a large strangling fig near Kali Kopi in south Papua. Dominant tree genera and species were Ficus benjamina, Macaranga, Buchanania, Canarium, Timonius, Syzygium, and Palaquium. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 34.4 m2/ha and the total clear bole volume was 545.2 m3/ha.

secondary forest During the natural growth cycle of trees in the natural forest, gaps appear. These gaps vary in size from small gaps (associated with individual tree falls) to very large gaps (often associated with cyclones and landslides). Pre-existing seedlings tend to grow up in small gaps, whereas large gaps are colonized by species which

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were not present when the gap was formed. The forest developing in big gaps, consisting of light-demanding, or pioneer species, is called secondary forest. The trees in natural secondary forest stands are all about the same age, and more or less the same size. Secondary forests are generally poorer in species. Most of the secondary forest species cannot regenerate under a canopy (even their own canopy). Thus, seedlings of shade-tolerant species establish beneath the pioneers and as the pioneers mature and die they grow up to replace them. Species composition is largely determined by the availability of seeds of shade-tolerant species. Where long-lived pioneers invade, they can occupy the site for a century or more. The complete succession back to primary forest generally takes centuries. Floristically, secondary forest is of very different composition from primary forest. Most pioneer species belong to relatively few families, and many to genera which are scarce in or absent from primary forest. Macaranga species are among the most common pioneer invaders (Whitmore 1967). Many woody climbers start life in gaps and are carried up by the trees to persist in high primary forests. Succession in big gaps in natural forest is similar to the succession that takes place on garden sites, and follows the sequence as outlined below. The process is more rapid where seeds, seedlings, or advance growth of primary forest species are present on the disturbed site. Large gaps caused by natural events are rare. Cyclones are absent or very infrequent, and there are few landslides due to the gently sloping topography. Gaps are usually small and the result of the death of individual overmature trees. When these trees eventually fall, they produce a gap. Preexisting seedlings tend to grow up in these relatively small gaps and the species are typical of the climatic community. The most extensive natural stands of secondary forest are located in areas that have experienced high water flooding in the past. On these sites, exceptional floods result in the replacement of tall primary forest trees by species more representative of late secondary forests. Such forests occur east of the road near Mile 38 and Mile 39 Camps, and between the Ajkwa and Otomona rivers north of the bridges. A stand opposite the Mile 39 camp is classified as seral Macaranga, Ficus, Diospyros, and Gonystylus dominated forest. The main causes of secondary forest in lowland areas are the practice of shifting cultivation and timber harvesting for construction activities. Shifting cultivation involves the clearing of the primary forest followed by burning and cultivation. Extensive areas of lowland forest have been converted to a mosaic of gardens and secondary vegetation in Papua. Although unmeasured, tropical foresters estimate that the full process for the rainforest to develop from secondary vegetation, or from clear-cutting, to a climatic climax forest takes several hundreds of years, depending on the availability of nearby seed sources (Gomez-Pompa et al. 1991).

river bed vegetation types on alluvial valleys River bed and river bank seres are similar to that which occurs in the alluvial valleys. In the absence of fire, the pioneer seral herbaceous community is invaded by fast-growing light-demanding woody plants, tall gingers, ferns, and wild bananas, and is transformed into a mixed species scrubland. Sometimes trees are left

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standing during cultivation, or are planted for their useful products, and are scattered through this community type. They include Canarium indicum, Terminalia kaernbachii, Dracontomelon puberulum, Pangium edule, Gnetum gnemon, Artocarpus altilis, the bettlenut palm Areca, and Bombax ceiba for kapok (tree-fiber, like cotton). These tend to be indicators of previous home sites in long abandoned areas. These were noted in abandoned village sites along the Minajerwi River, and in regrowth areas around Timika in south Papua. Pioneer tree species become established within the scrub and develop into a woodland or forest. Some of these trees grow from seeds, while others sprout from tree roots which were left in the ground during land clearing and site preparation. Dominant trees on the alluvial fans were Macaranga, Mallotus, Ficus, and Artocarpus. Reynders (1962) described the vegetation in the Star Mountains following disturbance by humans. In the absence of fire or cultivation, the light-demanding shrubs and trees are gradually replaced by species that are more shade-tolerant, and thin woody climbers replace the herbaceous creepers. There are generally few pandans, palms, and epiphytes. Over time, the forest becomes more varied in tree height and diameter. Larger woody climbers, rattan, and other palms appear and increase in frequency. Epiphytes become more common and varied. A ground layer of shade-loving herbs develops. In the process, the secondary forest develops the appearance and structure of the primary forest. However, long-living secondary forest tree species may continue to dominate the site. This may include species of Cryptocarya, Elmerrillia, Endospermum, Melicope (Euodia), Pimelodendron, and Sterculia. These stands have gone through an initial natural thinning stage as a result of species interactions. Community structure means that the seral stand is no longer composed of one age class, but has one age class in the overstory with some regeneration occurring that is in a very much younger age class. One such stand occurring north of Timika may be classified as seral Macaranga-Ficus-Anthocephalus-Artocarpus-dominated alluvial fans medium closed-forest community. Where stands are dominated by seral overmature overstory species the seral species (trees) composing the main upper (tree) canopy die and provide room for the next generation. The next generation may be composed of predominantly the same seral species, but in a different proportion, different species that are either more shade tolerant or competitively more successful, and/or climax species. Frequently, there is a secondary tree canopy that approaches a more even height class distribution. An evident gap usually exists between the dominant height class and the secondary tree canopy. Some individuals of the climax community may have grown to reach the main canopy by this time. Typically, stand biomass or volume declines because of mortality. One example, located along the PTFI road north of Timika, was classified as an overmature seral forest community dominated by Macaranga, Ficus, Anthocephalus, and Artocarpus. The stands experience natural thinning and develop a structure similar to that expected in the climatic climax community. Some remnants of seral stands may remain, but they should not have any effect on the density or structure of the stand. These stands lack the floristic richness of the climatic climax forest. It is expected to take hundreds of years for

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this floristic richness to be re-established and this depends on seed availability. Because of the relatively recent nature of settlements around Timika there are few examples of young and maturing climatic climax communities associated with shifting cultivation in the area. Repeated cutting and burning, plus cultivation or grazing, causes site-deterioration, including soil erosion. Instead of trees, stands of shrubs develop including Melastoma malabathricum, Lantana camara (which establishes in full light on compacted soil), Rhodamnia cinerea, Rhodomyrtus tomentosa, and locally Dillenia suffruticosa. The recently introduced Piper aduncum dominates sites in Papua New Guinea and some areas around Jayapura. Ultimately, disturbance leads to the development of open grassland of Imperata cylindrica (alang alang), which has underground diffuse rhizomes and can thereby survive fires. Pioneer species are most commonly found on disturbed roadside sites in south Papua.

natural secondary forest The most extensive natural stands of secondary forest are usually located in areas that have experienced high water flooding in the past. On these sites, an exceptional flood presumably resulted in the replacement of tall primary forest trees by species more representative of late secondary forests. Such forests occur east of the road near Mile 38 and Mile 39 Camps, and between the Ajkwa and Otomona Rivers in south Papua. Sites opposite the Mile 39 Camp were sampled. The four dominant genera were Macaranga, Ficus, Diospyros maritima, and Gonystylus macrophyllus. Other important genera recorded from the sites were Canarium, Diospyros maritima, Calophyllum, and Atalaya papuana. The estimated total basal area for trees with a diameter greater than 10 cm dbh was 36.1 m2/ha and the total clear bole volume was 304.1 m3/ha.

vegetation of ultrabasic and serpentine areas The ultrabasic and serpentine vegetation and flora of New Guinea have not been studied in detail. Van Royen (1963), and Paijmans (1976), and Takeuchi (2003b) included general discussions on these floras. The predominant species in forests on ultrabasic soils in New Guinea is often Casuarina papuana though this species has a wider ecological range in the Solomon Islands. Van Royen (1976) described the vegetation and flora of Waigeo Island, including a list of the major plant species. He described forest dominated by Vatica-Dillenia with Agathis locally dominant on lateritic soils with underlying limestone or ultrabasics. Brooks (1987) gave a brief description of the serpentine floras in New Guinea, noting that only Rinorea bengalensis, collected from the Bowutu Mountains, is known to accumulate nickel. Takeuchi (2003a) described the ultrabasic vegetation from the Raja Ampat Islands and listed the species present there. He noted that the flora is depauperate, with many of the common lowland families poorly represented. Van Royen (1960) also described the forest from Waigeo Island. Xerophytic vegetation is particularly common to north of Waigeo and on smaller areas of ultrabasic soils around Mayalibit Bay. Several vegetation types occur in this area: grass-sedge

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vegetation; myrtaceous communities dominated by Myrtella, Decaspermum, and Baeckia; and stands dominated by Casuarina rumphiana. Some lowland areas support a xerophytic vegetation dominated by Buchanania papuana and Vitex. On the ultrabasic formations of the Makanoi Range the forest above 200 m is dominated by stands of Castanopsis acumminatissima and Hopea. Araucaria cunninghamii is scattered but prominent. Engelhardtia rigida and Gevuina papuana are very common, often with Kania eugenoides and Eucalyptopsis papauana. On higher ridge crests on the ultrabasics Nothofagus flaviramea and Dacrydium elatum frequently occur. Xerophytic vegetation (vegetation adapted to low moisture) is particularly common north of Waigeo and on smaller areas of ultrabasic soils around Mayalibit Bay (van Royen 1976). Several types have been identified: grass-sedge vegetation; myrtaceous vegetation (Myrtella, Decaspermum, Baeckia), and Casuarina rumphiana stands. A xerophytic vegetation dominated by Buchanania papuana and Vitex occurs in some lowland areas. In this forest the most common shrubs and subcanopy trees are Decaspermum fruticosum, Myrtella beccarii, and Dodonea viscosa, mixed with Casuarina, Wendlandia, and other taxa. A more complete list of the species is given in van Royen (1960).

Limestone Vegetation Cape Suadja (the promontory to the north of Humboldt Bay) is dominated by Pometia forest that covers the limestone (van Royen 1965). A detailed report by J.F.U. Zieck in 1960 covers Mt Meja. The forest grows on limestone at 150–175 m altitude. Some areas are young secondary forest. Two forest types occur in the reserve of 400 ha. Pometia-Sapotaceae occurs on drier areas and Pometia-Homalium-Celtis and Intsia in wetter areas. Takeuchi (2003a) provided detailed descriptions of the limestone flora and vegetation in the Raja Ampat District of Papua. Stunted, woody, wind sheared vegetation, is comprised of Stenocarpus moorei, Exocarpus latifolius, Polyscias, Wikstroemia androsaemifolia, and Calophyllum with many vines including Alyxia purpureclada. Dwarfed plants of Podocarpus polystachys, to 1 m tall on solid limestone, sprawl across ledges. Galubia costata is a common emergent on limestone. Many new species records were made from the region but the limestone flora remains poorly documented. Paijmans (1976) stated that there is no evidence that any tree species are confined to limestone. Further studies will be required to confirm this.

Lowland Grassland Communities Grassland communities are locally common, especially on the southern slopes of the Cyclops Mountains where they are regularly burned. The composition of the grasslands varies greatly according to the underlying geology. Cape Tanah Merah, which has a low grade ore of copper and nickel, supports an open grassland vege-

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tation which is often bare and subject to high rates of erosion. Swamp and aquatic vegetation occur throughout the area, particularly along the southern foothills of the Cyclops around Lake Sentani.

Acknowledgments We wish to express our sincere thanks to Dr W. Vink for his very constructive comments on this chapter.

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. Bull. Am. Mus. Nat. Hist. 79: 199–288. Brass, L.J. 1941. The 1938–1939 Expedition to the Snow Mountains, Netherlands New Guinea. J. Arnold Arbor. 22: 271–345. Brooks, R.R. 1987. Serpentine and Its Vegetation. A Multidisciplinary Approach. Croom Helm, London and Sydney. Dick, J. 1991. Forest land use, forest use, forest use zonation and deforestation in Indonesia: a summary and interpretation of existing information. Ministry of Population and Environment (KLH) and the Environmental Impact Management Agency, BAPEDAL, Jakarta. Go´mez-Pompa, A., T.C. Whitmore, and M. Hadley (eds.). 1991. Rain Forest Regeneration and Management. Man and the Biosphere Series Volume 6. FAO Paris, UNESCO, and the Parthenon Publishing Group, Paris. Johns, R.J. 1982. Plant zonation. Pp. 309–330 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Kluwer Academic Publishers, Dordrecht, The Netherlands. Johns, R.J. 1986. The instability of the tropical ecosystem in Papuasia. Blumea 31: 341–371. Johns, R.J. 1987. A provisional classification of the dipterocarp forests of Papua New Guinea. In Kostermans, A.J.G.H. (ed.) Proceedings of the 3rd Round Table Conference on Dipterocarps, Samarinda. Lam, H.J. 1934. Materials towards a study of the flora of the island of New Guinea. Blumea 1: 115–159. Lam, H.J. (trans. L.M. Perry). 1945. Fragmenta Papuana [Observations of a Naturalist in Netherlands New Guinea]. Sargentia 5: 1–196. Paijmans, K. 1967. Vegetation of the Safia-Pongani area. CSIRO Aust. Land Res. Series 17: 142–167. Paijmans, K. 1970. An analysis of four tropical rain forest sites in New Guinea. J. Ecol. 58: 77–101. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO in association with Australian National University, Canberra. RePPProT. 1986. Review of Phase I and II Results for Irian Jaya. Regional Physical Planning Programme for Transmigration (RePPProT), Ministry of Transmigration, Jakarta. RePPProT. 1990. The Land Resources of Indonesia. A National Overview. Regional Physical Planning Programme for Transmigration (RePPProT), Ministry of Transmigration and the U.K. Overseas Development Agency, Jakarta.

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Lowland Vegetation of Papua / 961 Reynders, J.H. 1962. Shifting cultivation in the Star Mountains area. Nova Guinea Anthropology 10 (3): 45–73. Ridley, H.N. 1916. Report on the botany of the Wollaston Expedition to Dutch New Guinea. Trans. Linn. Soc. ser. 2 Bot. 1–269. Ridsdale, C.E. 1968. Botanical results of the New Guinea border demarcation expedition, 1967. Trans. Papua New Guinea Sci. Soc. 9: 3–22. Robbins, R.G. 1968. Vegetation of the Wewak-Tari area. CSIRO Aust. Land Res. Series 22: 109–124. Shea, G.A., P. Puradyatmika, A. Maulensey, and D. Martindale. 1998. Biodiversity Surveys in the PT Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. 11 vols. PT Hatfindo Prima, Bogor, Indonesia. Takeuchi, W. 2003a. A community-level floristic reconnaissance of the Raja Ampat Islands in New Guinea. Sida 20: 1093–1144. Takeuchi, W. 2003b. Botanical summary of a lowland ultrabasic flora in Papua New Guinea. Sida 20: 1491–1559. van Royen, P. 1960. Vegetation of some parts of Waigeo Island. Nova Guinea Bot. Series 3: 25–62. van Royen, P. 1963. The Vegetation of the Island of New Guinea. Department of Forests, Division of Botany, Lae. van Royen, P. 1965. Sertulum Papuanum 14. An outline of the flora and vegetation of the Cycloop Mountains. Nova Guinea Bot. Series 21: 451–469. van Steenis, C.G.G.J. 1957. Outline of vegetation types in Indonesia and some adjacent regions. Proceedings of the 8th Pacific Science Congress 4: 61–69. 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: Irian Jaya Studies. Balkema, Rotterdam/Brookfield Whitmore, T.C. 1967. Studies of Macaranga, an easy genus of Malayan wayside trees. Malay. Nat. J. 20: 89–99. Whitmore, T.C. 1984a. Tropical Rainforests of the Far East. Oxford Science Publications. Clarendon Press, Oxford. Whitmore, T.C. 1984b. Vegetation map of Malesia, at scale 1:5 million. Journal of Biogeography 11: 461–471. Zieck, J.F.U. 1960. The Tafelberg hydrological forest reserve near Manokwari [in Dutch]. Unpublished report.

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5.9. Heath Vegetation of Papua garry a. shea, robert j. johns, willem vink, and pratito puradyatmika e a th f o r es t s occur throughout Papua but are generally limited in area. Dick (1991), based on RePPProT (1986), gave the following areas of heath forest in Indonesia: Sumatra, 493 km2; Kalimantan, 26,753 km2; Sulawesi, 792 km2; and Papua, 7,100 km2. The 7,100 km2 of heath forest in Papua that were mapped, and reported by RePPProT (1986), for Papua represents 20.2% of the total area of heath vegetation in Indonesia. The extent of heath forest in Papua is based on aerial photograph interpretation and has not been confirmed by ground truthing. The largest area of heath forests was on the dissected Pleistocene terraces in south Papua, south of Mt Jaya. Other areas of heath forest in Papua include a 2,000 ha area which was reported west from Etna Bay between Bintuni Bay and the tip of Arguni Bay (Syuga-Wagura) at 400 m altitude. Vink (1998) reports a distinct vegetation on very poor soils on white sands, interspersed with loam from the Teminabuan area, south of Beriat at 50 m altitude. In this stand emergent Agathis labillardieri are underrepresented probably because of the long-term tapping of resin and removal of seedlings for plantations (Vink 1932: 34). Beehler (pers. comm.) also observed heath forest below Holuwon Airstrip, in Yali territory just east of the Baliem Gorge, at an altitude of 950 m on the south side of the main range. The only large area of heath forest for which data are available in Papua occurs on the southern slopes of the central mountains in the PT Freeport Indonesia (PTFI) Project Area and, more extensively, in adjoining areas terraces of the Lorentz World Heritage Area. This heath forest has developed from outwash fans composed of finely ground calcareous glacial outwash mixed with coarser stones, the result of the extensive Pleistocene glaciation of the limestone peaks of the central mountain ranges. This chapter is based on the studies by Shea et al. (1998). In south Papua the terraces occur at an altitude of 100–650 (–700) m asl. At approximately 150 m asl in south Papua the lowland rain forest on the alluvial fans grades into heath forest on the terraces. A zone of transitional forest, described in the first section of this paper, blends the characteristics of both lowland rain forest and heath forest. Heath forest occurs on soils derived from siliceous parent materials which are inherently poor in bases, highly acidic, lack buffering capacity, and are also coarsely textured. These heath forests differ from lowland evergreen rainforest in floristics, structure, and physiognomy. Whitmore (1984) notes that in the heath forests in Malesia there are more trees with small leaves than in the lowland evergreen rain forest. Microphyll and

H

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

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mesophyll leaves are most common. Many leaves are distinctly sclerophyllous (that is, thick and leathery). Deciduous species are generally absent. On aerial photographs, the flat canopy is highly distinctive owing to the pale tone and its very fine texture, which results from tree-crown structure and small leaf sizes. Species with leaves held obliquely vertical may be common (for example, Tristania spp. and some Sapotaceae). Trees of large girth are rare; buttresses are smaller, but stilt roots are more common than in lowland evergreen rain forest. Big woody climbers (including climbing palms) are rare, but slender, wiry, independent climbers are common. Epiphytes occur frequently and photophytes occur nearer the ground than in lowland evergreen rainforest. Myrmecophytes are abundant especially in the more open and stunted heath forests. The insectivores Nepenthes spp. are common in open places. The ground often has a bryophyte cover. The streams draining areas of heath forest are tea-colored by transmitted light, and opaque black by reflected light due to the presence of organic colloids. The streams are usually acid (pH ⬍ 5.5), and with a low oxygen content (Johnson 1967). Heath forests and upper montane forests have many features in common. Whitmore (1984) summarized the similarities as follows: the forest canopy of both vegetation types is rather even, dense, and commonly with a high albedo (of pale color on aerial photographs). Trees have dense crowns. Microphyll is the predominant leaf size (Raunkier 1934). The leaves tend to be held obliquely vertical, often closely placed on the twigs. The plant communities of both vegetation types include facies of low biomass in comparison with other forests. Paths made by animals or human travelers remain open for a long time, and many species have hard dense wood—two factors which both suggest that the growth rate is slow. Big woody climbers are absent or rare in both forests. Some species are in common in both vegetation types, including species of Vaccinium and Rhododendron. These may occur as trees, shrubs or epiphytes. Many filmy ferns and bryophytes occur in both these communities.

Transitional Forests between Alluvial Fans and Terraces The transitional communities blend the characteristics of the adjacent forest: lowland rainforest and heath forest. Shea et al. (1998) sampled three transitional stands in south Papua: one near Mile 38, one near Mile 39, and one 1.5 km north of Mile 39. The Mile points refer to standard localities in the PTFI Contract of Work area in south Papua. The stand located east of Mile 38 is dominated by Alstonia, Buchanania macrocarpa, Calophyllum costatum, and Ficus. Other common trees include Diospyros maritima, Garcinia dulcis, Syzygium polyantha, Erythroxylon, and Elaeocarpus. The estimated total basal area for trees with a diameter greater than 10 cm dbh is 28.7 m2/ha. The second stand is dominated by Alstonia, Calophyllum, and Elaeocarpus. The stand located 1.5 km north of Mile 39 in south Papua has the dominant trees Endospermum moluccanum, Nauclea papuana, Sloanea archboldiana, and Alstonia scholaris. Other common trees include Prumnopitys amara, Diospyros maritima, Fagraea woodiana, Timonius sp., Austrobuxus

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nitidus, and Calophyllum costatum. The estimated total basal area for trees with a diameter greater than 10 cm dbh is 16.4 m2/ha and the total clear bole volume is 123.3 m3/ha. Transitional lowland-heath forest is dominated by Endospermum, Nauclea, Sloanea, and Alstonia.

Heath Forest The factors contributing to the curious structure and the xeromorphic physiognomy (being adapted to low moisture) of heath forests are still unknown. Because heath forest is restricted to soils developed on siliceous deposits which are podzolized to a varying degree, periodic water deficiency and low supply of nutrients may be contributing factors. Richards (1952) thought nutrient deficiency of the very acid soils was the more likely cause. Brunig (1970, 1971, 1974) carried out more detailed studies and concluded that several features of heath forest are adaptations to survival of the community in conditions of periodic drought. Whitmore (1984) noted that, in general, respiration and photosynthesis both increase with temperature up to about 25C for many species. At such temperatures, conditions are favorable for growth, and net photosynthesis occurs. At higher temperatures, photosynthesis declines while respiration rises sharply, conditions unfavorable for growth. Leaves are cooled by three processes: convection, latent heat absorbed by the evaporation of water transpired, and re-radiation of incident energy (Whitmore 1984). Drought periods represent a serious danger to plants, in that the water supply for transpirational cooling is then cut off. Periodic droughts occur in even the perhumid parts of Malesia. The amount of water available to plants depends also on the storage capacity of the soil. Heath forest soils are mostly porous and coarse. Roots do not penetrate into the hard soils. The water-storage capacity of heath forest soils is less than that of soils in other forest types, and water shortage is more likely to occur. Brunig (1970, 1971) demonstrated that several of the striking structural and physiognomic features of heath forest are likely either to minimize water loss, thereby conserving the water available to the forest, or to reduce the heat loads on the leaves, which could be vital to survival in drought periods. Whitmore (1984) summarized Brunig’s findings as follows: the canopy of the heath forest is generally not as rough as the canopy of the adjacent lowland evergreen rain forest. Reduced canopy roughness has two effects. First, the amount of incident radiation intercepted is decreased progressively as the sun declines from the zenith. Secondly, the aerodynamic roughness of the canopy decreases and consequently, so does the turbulence from free and forced convection. The result is a tendency to decrease the rate at which water vapor is lost from the canopy. Mean leaf size in heath forest is less than in evergreen rainforest. Steeply inclined leaves and twigs are frequent in heath forest and mostly on the driest sites. In heath forest, a number of features of structure and physiognomy can be considered to be adaptations to minimize the effects of periodic water stress, which

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perhaps occurs at heath forest sites more than at sites occupied by other lowland forest. Whitmore (1984) suggested that the adaptive significance of the peculiar characteristics of heath forest canopy, tree crowns, and leaves is to minimize heat load on those occasions when transpirational cooling is restricted (i.e., during droughts). Several features of heath forests suggest inorganic nutrient deficiency. Heath forest is easily degraded to a low scrub if burned or cultivated. Plants with supplementary means of mineral nutrition are common, namely, myremecophytes and insectivorous plants. Casuarina, which is common in the heath forest in south Papua, has nodules containing nitrogen-fixing bacteria on its roots. Vink (pers. comm.) notes that Casuarina is often dominant in secondary forests and suggests this could have contributed to larger than expected areas being mapped as heath forest. Mycorrhiza may also play an important role in heath forests. No data are available on rates of biomass production in heath forest in Papua, but the studies of forests in the Gunung Mulu National Park (Borneo) quantify some aspects of the nutrient cycle. The most notable discovery was that the nutrient pool in the Mulu heath forest is about as large as in the other forest types (Proctor et al. 1983; Whitmore 1984). The rate of litter fall was similar, while the annual rate of return of all measured inorganic nutrients in fine litter fall was less than in the lowland evergreen forest, especially for nitrogen and potassium. Whitmore (1984) speculated as to why heath forests are so easily degraded by logging and clearing. He noted that in heath forest humic podzols are of low clay content, and much of the cation exchange capacity lies in the organic matter. This is easily oxidized once the forest cover has been removed, especially from the surface where it is concentrated, and especially if the soil is disturbed. The small amount of clay present is likely to wash rapidly downwards through such coarsely textured soil. Thus, these soils are easily degraded to become bleached sands over a humic pan. This ‘‘fragility’’ of the soil may be the explanation for the notorious infertility of heath forest sites. Thus, heath forest sites are increasingly difficult to reforest, the more the original forest and the soil have been disturbed. Thus, the heath forest vegetation type occurs on sites that have a number of unfavorable characteristics. Its distinctive flora, structure, and physiognomy are adapted to cope with the unfavorable site conditions. The various morphological traits that appear to minimize heat load are therefore considered to be adaptive. Heath forest (kerangas) develops mainly over coarse siliceous deposits which give rise to podzolized soils. The greatest extent of heath forest in Malesia is in Borneo where it occurs around much of the coastline on raised terraces of poorly consolidated coarse, sandy, marine, and riverine sediments. Similar and less extensive terraces occur in Papua, Peninsular Malaysia, and on the east coast of Sumatra. Coastal heath forests have been extensively converted to secondary savanna maintained by grazing and burning (Whitmore 1984). In parts of Borneo, podzolic soils become temporarily, to more or less permanently, waterlogged because the hard pan in the soil forms an impermeable layer. The forest is here known as kerapah, an Iban term denoting swampy conditions. In structure and physiog-

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nomy this is heath forest, though some species are particularly common in these sites. Burnham (1984) described the soils as coarse, siliceous deposits which generally support heath forests in Malesia. Podzols (spodosols) occur in places where parent materials consist predominantly of quartz; for example, beach sand, sandstone, or quartzite, or much more locally on unusually siliceous volcanic rocks. These parent materials are low in clay and in bases and also in minerals that might weather into these products. The soils are readily permeable and are well drained, except that the influence of ground water may sometimes cause subsoils to be waterlogged. Podzolization is possible wherever rainfall substantially exceeds evapotranspiration. In Malesia, podzols are rare unless the mean rainfall exceeds 2,000 mm/year. The podzols of Malesia have the following characteristics; under a raw humus layer in undisturbed sites, or elsewhere under a dark gray sandy A horizon, lies a bleached E (formerly called A2) horizon, below which is a very dark or strongly colored B horizon enriched in colloidal organic mater (humus podzol or humod) or in organic matter and sesquioxides (humus-iron podzol or orthod). The constituents which have accumulated in the B horizon have been moved by percolating water from the upper part of the soil profile. Mobilized sesquioxides often originate from the chemical breakdown of clay, which commonly occurs to some extent in the very acid conditions characteristic of podzols. The most widespread environment where podzols are formed is on old beach deposits lying just inland in the form of low ridges or terraces. These sediments are in their second or third weathering cycle and are very deficient in bases. Lowland podzols further inland on consolidated rocks, usually sandstone but also acidic volcanics, quartzite, and conglomerates, are restricted mainly to Borneo. Podzols are also frequent among mountain soils. Brunig noted (in Ashton 1971) that the structure and physiognomy of heath (kerangas) forest in Sarawak differs from the lowland dipterocarp forest, not only because of the change in species but also because of the striking difference in the structure, texture, and the whole color of the forest. In the lowland dipterocarp forest the entire growing space is loosely and evenly filled with green foliage. In the heath forest, the story formed by large saplings and small poles predominates and is often difficult to penetrate. The canopy is low, uniform, and usually densely closed with no trace of layering. Single emergents may occur and usually indicate extreme site conditions. Brown and reddish colors prevail in the foliage of the upper part of the canopy.

lowland heath vegetation in new guinea In the area of the Star Mountains in central New Guinea, inland heath forests are all found on river terraces, some of which have impeded drainage, and some of which carry Agathis labillardierei forest (Reynders 1964; Whitmore 1984). Paijmans (1976) did not report or describe any extensive areas of inland heath forest for New Guinea. Specht and Womersley (1979) described the heathlands from Normanby Island, which are developed on hillsides with surface covering of

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weathered sandstone, as: ‘‘scattered trees of Xanthostemon sp. (Myrtaceae) dominate the scene but there is a rather open shrubland of Baeckea frutescens, Myrtella beccarii and occasional shrubby plants of Xanthomyrtus sp. (all Myrtaceae). Nepenthes sp. (probably N. mirabilis) is very abundant and shrub Osbeckia chinensis are common. The ground flora is sparse between medium to large sandstone blocks. Here Drosera sp. and several sedges are found. Perched as an epiphyte exposed to the sun and drying sea breezes is a white-flowered Dendrobium (sect. Ceratobium). This orchid is frequently very abundant, and is associated with the curious ‘anthouse’ plants (Hydnophytum sp. and Myrmecodia sp.) also growing as epiphytes or sometimes perched on large rocks. The whole community is very open and exposed to the sun.’’ Although the floristics are different, there is a degree of similarity with the kerangas vegetation of the sandstones of Sarawak. Trees, when occurring, tend to have a sparse crown. In Papua New Guinea, there are lowland heath-like communities of some extent in the Sepik Basin. Of the low altitude communities, those at Kunjengini and Green River are known in some detail (Specht and Womersley 1979). The heath vegetation at Green River in the upper Sepik Basin is more characteristic. Unlike Kunjengini, the narrow forest ribbons fringing drainage channels include species not found in the usual lowland forest. This forest includes herbs like Lycopodium carolinianum, Burmannia sp., Cladium undulata, Cyperus sp., Rhynchospora rugosa, Scleria sp., Tricostularia undulate, Drosera burmannii, Eriocaulon australe, Flagellaria sp., Nepenthes sp., Spiranthes sp., Eriachne pallescens, E. triseta, Garnotia mezii, Isachne confusa, I. globosa, I. myosotis, Ischaemum barbatum, Leersia hexandra, Salomonia sp., and Xyris companata, X. papuana, and Xyris sp. Woody plants, shrubs, or small trees include Elaeocarpus sepikanus, Rhododendron zoelleri, Litsea sp., Kibessia galeata, Baeckea frutescens, Metrosideros eugenioides, Xanthostemon sp., Dipodium pandanum, Mussaenda ferruginea, Euodia sp., and Sterculia dalrympleana var. schlechteri.

heath vegetation types Heath forests vary in structure and species composition from site to site. They are mainly medium height forest with occasional tall emergent trees. The heath forests are structurally similar to the heath forest that occurs in Kalimantan. They are extensive on dissected terraces of the PTFI COW Mining and Project Area in south Papua. Prior to the study by Shea et al. (1998), there were no detailed descriptions of these heath forests in south Papua. A number of heath forest vegetation types were identified and classified, both qualitatively and quantitatively, by Shea et al. (1998).

heath forest on the slope of terraces The terrace slopes in south Papua occur mainly between Km 56 and Km 68 along the PTFI road. The slope is dissected by rivers and streams. Two stands occurring on dissected terraces were studied; Dacrydium, Stemonurus, Calophyllum, and Ceratopetalum dominated at Km 56, and at Km 68 Dacrydium, Palaquium, Teijs-

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968 / r. j. johns, g. a . shea, w . v i n k, & p. puradyatmika

manniodendron, and Calophyllum dominated the forest community. Other important trees include Ternstroemia, Palaquium amboinensis, Casuarina papuana, Xanthophyllum scortechinii, Pandanus, Teijsmanniodendron bogoriense, Diospyros maritima, and Syzygium stipilaris (Table 5.9.1). The estimated total basal area for trees with a diameter greater than 10 cm dbh is 34.1 m2/ha and the total clear bole volume is 261.5 m3/ha. The second stand on the terrace slopes is located east of road at Km 64 along PTFI Road, south Papua. The dominant tree genera are Dacrydium, Palaquium amboinensis, Teijsmanniodendron bogoriense, and Calophyllum congestifolium. Other trees include Drimys piperita, Syzygium polyantha, S. papuasicum, Garcinia dulcis, and Pandanus sp. (Table 5.9.2). The estimated total basal area for trees with a diameter greater than 10 cm dbh is 17.4 m2/ha and the total clear bole volume is 126.4 m3/ha. The forest at the crest of the terrace slopes has a structure and physiognomy similar to that described for heath forests from Borneo (Brunig 1974). Two stands were studied by Shea et al. (1998). The first stand assessed by Shea et al. (1998) is located at Km 64 along PTFI Road. Common trees are Casuarina papuana, Syzygium polyantha, Dacrydium, Palaquium amboinensis, Pandanus, Drimys piperita, Madhuca, Xanthostemon, and Calophyllum papuana. The most important trees are Syzygium polyantha, Dacrydium, Palaquium amboinensis, Diospyros maritima, Pandanus, Drimys piperita, Xanthostemon, and Calophyllum papuana. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 21.8 m2/ha and the total clear bole volume is 130.9 m3/ha (Table 5.9.3). The second stand was located at Km 64 along PTFI Road, south Papua. Dominant trees in the stand are Syzygium polyantha, Tristania macrosperma, Pandanus, and an unidentified genus of trees. Other important trees are Distylium stellare, Buchanania macrophylla, Diospyros maritima, Ternstroemia, Calophyllum costatum, Prunus costata, and Garcinia (Table 5.9.4). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 21.5 m2/ha and the total clear bole volume was 148.5 m3/ha.

heath forest on terraces The terrace surfaces between Km 68 and Km 71 have a rolling topography. The trees are characterized by having a heavy mass of bryophytes on the trunks and in the tree crowns. The trees are generally taller than those occurring on the terrace slope and the crest of the terrace slope. Shea et al. (1998) described two stands occurring at Km 70. These are dominated by Dacrydium, Syzygium, Metrosideros pullei, and Decaspermum fruiticosum. Other trees are Calophyllum costatum, Diospyros maritima, and Sloanea archboldiana (Table 5.9.5). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 41.7 m2/ha and the total clear bole volume was 450.3 m3/ha. The dominant trees in the second site are Metrosideros parviflora, Calophyllum congestifolium, Sloanea archboldiana, and Pandanus. Other species are Alstonia spectabilis, Garcinia, Syzygium anomala, and

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PAGE 968

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8.06

Syzygium stipilaris

PS

Note: Stand located on a terrace slope. Source: Shea et al. (1998).

261.51

3.71

Ternstroemia sp.

Total

56.69

Stemonurus sp.

27.43

4.12

Calophyllum papuana

1.65

Podocarpus sp.

6.59

Diospyros maritima

Xanthophyllum scortechnii

2.88

Teijsmanniodendron bogoriense

1.47

21.66

Palaquium amboinensis

Timonius sp.

13.19

2.88

17.17

94.01

Ceratopetalum succirubrum

Pandanus sp.

Casuarina papuana

Dacrydium sp.

Species

Total clear bole volume (m3/ha)

833.3

33.3

166.7

66.7

33.3

33.3

33.3

33.3

33.3

33.3

66.7

33.3

33.3

33.3

20.0

Absolute density

100

4

20

8

4

4

4

4

4

4

8

4

4

4

24

Relative density (%)

34.08

0.59

7.72

2.68

1.05

0.59

0.26

1.05

0.59

0.26

1.90

2.35

0.59

1.64

12.82

Absolute dominance (m2/ha)

100.00

1.73

22.65

7.87

3.07

1.73

0.77

3.07

1.73

0.77

5.57

6.91

1.73

4.80

37.62

Relative dominance (%)

2.00

0.08

0.42

0.17

0.08

0.08

0.08

0.08

0.08

0.08

0.17

0.08

0.08

0.08

0.42

Absolute frequency

100.00

4.17

20.83

8.33

4.17

4.17

4.17

4.17

4.17

4.1

8.33

4.17

4.17

4.17

20.83

Relative frequency (%)

300.00

63.48

24.20

11.24

9.89

8.93

11.24

9.89

9.89

78.93

21.90

15.08

9.89

12.97

82.45

Importance value

Table 5.9.1. Transitional lowland/heath forest dominated by Dacrydium, Stemonurus, Calophyllum papuana, and Ceratopetalum succirubrum

Heath Vegetation of Papua / 969

PAGE 969

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8.06

Syzygium polyantha

PS

Note: Stand located on a terrace slope. Source: Shea et al. (1998).

Total

126.38

23.08

1.47

Palaquium amboinensis

4.53

Garcinia dulcis

19.78

14.61

Syzygium papuasicum

Teijsmanniodendron bogoriense

Calophyllum congestifolium

5.13

1.10

Pandanus sp.

Drimys piperita

48.63

Dacrydium sp.

Species

Total clear bole volume (m3/ha)

533.33

33.33

33.33

33.33

33.33

33.33

100.00

33.33

33.33

200.00

Absolute density

100.00

6.25

6.25

6.25

6.25

6.25

18.75

6.25

6.25

37.50

Relative density (%)

17.40

2.35

1.05

0.26

0.59

2.35

2.22

1.05

0.26

7.26

Absolute dominance (m2/ha)

100.00

13.53

6.02

1.50

3.38

13.53

12.78

6.02

1.50

41.73

Relative dominance (%)

1.08

0.08

0.08

0.08

0.08

0.08

0.25

0.08

0.08

0.25

Absolute frequency

100.00

7.69

7.69

7.69

7.69

7.69

23.08

7.69

7.69

23.08

Relative frequency (%)

300.00

27.48

19.96

15.45

17.33

27.48

54.61

19.96

15.45

102.31

Importance value

Table 5.9.2. Transitional lowland/heath forest dominated by Palaquium amboinensis, Teijsmanniodendron bogoriense, Calophyllum congestifolium, and Dacrydium

970 / r. j. johns, g. a . shea, w . v i n k, & p. puradyatmika

PAGE 970

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32.15

Syzygium polyantha

PS

Note: Stand located on the crest of terrace slopes. Source: Shea et al. (1998).

130.92

1.83

Sterculia sp.

Total

1.83

Ternstroemia sp.

9.52

2.2

Diospyros maritima

Palaquium amboinensis

1.1

Podocarpus neriifolius

3.3

3.71

Xanthostemon sp.

Calophyllum papuana

3.85

Madhuca sp. 1.1

4.35

Drimys piperita

Annonaceae

3.62

44.51

17.86

Pandanus sp.

Casuarina papuana

Dacrydium sp.

Species

Total clear bole volume (m3/ha)

1000.0

33.3

33.3

100

200

66.7

33.3

33.3

33.3

33.3

100

66.7

66.7

66.7

133.3

Absolute density

100.00

3.33

3.33

10.00

20.00

6.67

3.33

3.33

3.33

3.33

10.00

6.67

6.67

6.67

13.33

Relative density (%)

21.85

0.26

0.26

1.44

4.38

0.52

0.26

0.59

0.26

0.59

0.78

0.85

0.85

6.54

4.25

Absolute dominance (m2/ha)

100.00

1.20

1.20

6.59

20.06

2.40

1.20

2.69

1.20

2.69

3.59

3.89

3.89

29.94

19.46

Relative dominance (%)

2.17

0.08

0.08

0.17

0.42

0.17

0.08

0.08

0.08

0.08

0.25

0.08

0.08

0.17

0.33

Absolute frequency

100.00

3.85

3.85

7.69

19.23

7.69

3.85

3.85

3.85

3.85

11.542

3.85

3.85

7.69

15.38

Relative frequency (%)

300.00

8.38

8.38

24.28

59.29

16.75

8.38

9.87

8.38

9.87

5.13

14.41

14.41

44.3

48.18

Importance value

Table 5.9.3. Transitional lowland/heath forest dominated by Casuarina papuana, Syzygium polyantha, Dacrydium and Palaquium amboinensis

Heath Vegetation of Papua / 971

PAGE 971

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1.10 2.20 3.71

Drimys piperita

Prunus costata

Ternstroemia sp.

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PS

8.01 148.55

Note: Stand located on a crest above a terrace slope. Source: Shea et al. (1998).

Total

Indet. sp.

41.94

1.83

Podocarpus neriifolius

Syzygium polyantha

7.46

Buchanania macrophylla 6.87

3.71

Garcinia sp.

Diospyros maritima

2.20

11.45

Palaquium amboinensis

Distylium stellare

2.06

14.24

Pandanus sp.

Calophyllum costatum

41.76

Tristania macrosperma

Species

Total clear bole volume (m3/ha)

866.66

33.33

166.67

100.00

33.33

100.00

33.33

33.33

33.33

33.33

33.33

33.33

33.33

133.33

66.67

Absolute density

100.00

3.85

19.23

11.54

3.85

11.54

3.85

3.85

3.85

3.85

3.85

3.85

3.85

15.38

7.69

Relative density (%)

21.46

1.64

5.63

1.11

0.26

1.11

0.59

0.26

1.64

0.59

0.59

0.26

0.26

3.53

3.99

Absolute dominance (m2/ha)

100.00

7.62

26.22

5.18

1.22

5.18

2.74

1.22

7.62

2.74

2.74

1.22

1.22

16.46

18.60

Relative dominance (%)

2.08

0.08

0.42

0.25

0.08

0.17

0.08

0.08

0.08

0.08

0.08

0.08

0.08

0.33

0.17

Absolute frequency

100

4

20

12

4

8

4

4

4

4

4

4

4

16

8

Relative frequency (%)

Table 5.9.4. Heath forest dominated by Syzygium polyantha, Tristania macrosperma, Pandanus, and an unidentified tree

300.00

15.47

65.45

28.72

9.07

24.72

10.59

9.07

15.47

10.59

10.59

9.07

9.07

47.85

34.29

Importance value

972 / r. j. johns, g. a . shea, w . v i n k, & p. puradyatmika

PAGE 972

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3.30

Cryptocarya caloneura

PS

433.3

33.3

33.3

33.3

33.3

66.7

33.3

33.3

33.3

100.0

33.3

Absolute density

Note: Stand located at Km 70 along PTFI Road, south Papua. Source: Shea et al. (1998).

450.31

3.71

Myristica fatua

Total

6.59

Diospyros maritime

2.61 43.96

Metrosideros pullei

Sloanea archboldiana

152.58

2.20

Podocarpus sp.

Syzygium sp.

5.13

224.38

5.86

Calophyllum costatum

Dacrydium sp.

Decaspermum fruticosum

Species

Total clear bole volume (m3/ha)

100.00

7.69

7.69

7.69

7.69

15.38

7.69

7.69

7.69

23.08

7.69

Relative density (%)

41.67

0.59

0.59

1.05

4.19

0.85

12.82

0.26

1.05

19.23

1.05

Absolute dominance (m2/ha)

100.00

1.41

1.41

2.51

10.05

2.04

30.77

0.63

2.51

46.15

2.51

Relative dominance (%)

1.08

0.08

0.08

0.08

0.08

0.17

0.08

0.08

0.08

0.25

0.08

Absolute frequency

100.00

7.69

7.69

7.69

7.69

15.38

7.69

7.69

7.69

23.08

7.69

Relative frequency (%)

Table 5.9.5. Heath forest dominated by Dacrydium, Metrosideros pullei, Decaspermum fruiticosum, and Syzygium

300.00

16.80

16.80

17.90

25.43

32.81

46.15

16.01

17.90

92.31

17.90

Importance value Heath Vegetation of Papua / 973

PAGE 973

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13.19 2.47

Garcinia sp.

Timonius sp.

PS

145.89

333.33

33.33

33.33

33.33

33.33

33.33

66.67

33.33

33.33

33.33

Absolute density

Note: Stand located at Km 70 along PTFI Road, south Papua. Source: Shea et al. (1998).

Total

44.88

16.03

Alstonia spectabilis

Metrosideros parviflora

3.30

14.01

Sloanea archboldiana

Syzygium anomala

19.78

Palm tree

9.16 23.08

Calophyllum congestifolium

Pandanus sp.

Species

Total clear bole volume (m3/ha)

100

10

10

10

10

10

20

10

10

10

Relative density (%)

14.33

3.21

0.59

1.05

1.64

0.59

2.22

1.05

2.35

1.64

Absolute dominance (m2/ha)

100.00

22.37

4.11

7.31

11.42

4.11

15.53

7.31

16.44

11.42

Relative dominance (%)

0.83

0.08

0.08

0.08

0.08

0.08

0.17

0.08

0.08

0.08

Absolute frequency

100

10

10

10

10

10

20

10

10

10

Relative frequency (%)

300.00

42.37

24.11

27.31

31.42

24.11

55.53

27.31

36.44

31.42

Importance value

Table 5.9.6. Heath forest dominated by Metrosideros parviflora, Calophyllum congestifolium, Sloanea archboldiana and Pandanus

974 / r. j. johns, g. a . shea, w . v i n k, & p. puradyatmika

PAGE 974

Heath Vegetation of Papua / 975

Timonius sp., and a palm (Table 5.9.6). The estimated total basal area for trees with a diameter greater than 10 cm dbh was 14.3 m2/ha and the total clear bole volume was 145.9 m3/ha. Another stand was located at Km 70 along the PTFI Road. The dominant trees are Metrosideros parviflora, Calophyllum congestifolium, Sloanea archboldiana, and Pandanus. The other common trees are Alstonia spectabilis, palm tree, Garcinia, Syzygium anomala, and Timonius sp. (Table 5.9.6). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) was 14.3 m2/ha and the total clear bole volume was 145.9 m3/ha.

Literature Cited Ashton, P.S. 1971. The plants and vegetation of Bako National Park. Malay. Nat. J. 24: 151–162. Brunig, E.F. 1970. Stand structure, physiognomy and environmental factors in some lowland forests in Sarawak. Trop. Ecol. 11: 26–43. Brunig, E.F. 1971. On the ecological significance of drought in the equatorial wet evergreen (rain) forest of Sarawak (Borneo). Erdkunde 23: 127–133. Brunig, E.F. 1974. Ecological Studies in the Kerangas Forests of Sarawak and Brunei. Borneo Literature Bureau, Kuching. Burnham, C.P. 1984. The forest environment: soils. In Whitmore, T. C. (ed.) Tropical Rain Forests of the Far East. Clarendon Press, Oxford. Dick, J. 1991. Forest land use, forest use, forest use zonation and deforestation in Indonesia: a summary and interpretation of existing information. Ministry of Population and Environment (KLH) and the Environmental Impact Management Agency, BAPEDAL, Jakarta. FAO/UNESCO. 1974. Soil Map of the World. A: 5,000,000. I Legend, UNESCO, Paris. Haantjens in Robbins, R.G. (1968). Vegetation of the Wewak-Tari area. CSIRO Aust. Land Res. Series, 22. Johnson, D.S. (1967). Distributional patterns of Malayan freshwater fish. Ecology 48: 722–730. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO in association with Australian National University, Canberra. Proctor, J., J.M. Anderson, P. Chai, and H.W. Vallack. 1983. Ecological studies in four contrasting lowland rainforests in Gunung Mulu National Park, Sarawak. I. Forest environment, structure and floristics. Journal of Ecology 71: 237–260. RePPProT. 1986. Review of Phase I and II results for Irian Jaya. Regional Physical Planning Programme for Transmigration (RePPProT). Ministry of Transmigration, Jakarta. Reynders, V.J. 1964. A soil sequence in the tropics from sea level to eternal snow. Snow Mountains, Central New Guinea. Int. Congr. Soil Sci. 5: 733–739. Richards, P.W. 1952. The Tropical Rain Forest. Cambridge University Press, Cambridge. Shea, G.A., D. Martindale, P. Puradyatmika, and A. Mandessy. 1998. Biodiversity surveys in the PT Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. PT Freeport Indonesia. Specht, R.L., and J.S Womersley. 1979. Heathlands and related shrublands of Malesia (with particular reference to Borneo and New Guinea). Pp. 321–328 in Specht, R.L.

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976 / r. j. johns, g. a . shea, w . v i n k, & p. puradyatmika (ed.) Heathlands and Related Shrublands. Ecosystems of the World 9A. Elsevier Scientific Publishing, Amsterdam. Vink, A.L. 1932. Memorie (vervolg) van overgave van de Onderafdeeling West NievwGuinea. Reprinted in Miedema, J., and W.A.L. Stokhof (eds.) Irian Jaya Source Materials, Ser. V, No. 3 DSALCUL/IRIS, Leiden/Jakarta. 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: Irian Jaya Studies. Balkema, Rotterdam. Whitmore, T.C. 1984. Tropical Rainforests of the Far East. Oxford Science Publications. Clarendon Press, Oxford.

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5.10. Montane Vegetation of Papua robert j. johns, garry a. shea, willem vink, and pratito puradyatmika o n ta n e r ai n f o re s t covers the largest area of any of the forest zones in New Guinea. In this chapter we discuss the classification of these forest areas into types, and provide descriptions of their typical floristic composition. We outline the diversity of species and discuss the concentration of fern and orchid genera and species in the montane zone. The montane regions of New Guinea are poorly known and a final classification of the plant formations has not been satisfactorily developed. Montane forests occur over an altitudinal range from (300–) 650–3,200 m asl. Many species have a broad altitudinal range because they extend into secondary forest types both above and below their normal range. This further complicates the altitudinal patterns caused by the ‘‘Massenerhebung effect’’ (Backhuys 1968). Papua has 44,470 km2 of montane forest. This represents 36.2% of the total area of montane vegetation for Indonesia following the mapping done by RePPProT (1986, 1990). The areas of montane vegetation in the other islands of Indonesia are: Sumatra, 32,355 km2; Kalimantan, 22,742 km2; Sulawesi, 21,810 km2; Nusa Tenggara, 1,204 km2; and Maluku, 1,393 km2; with a total of 122,973 km2 for Malesia. The montane environment for Malesia was described by Whitmore (1984). In this environment, the midday sun is high in the sky all year round, and there is only slight variation through the year in day-length. At higher altitudes the diurnal range in temperature is usually greater than the annual range of the means (Hnatiuk, Smith, and McVean 1976). The Freeport Mine has established a series of climatic stations from near sea level to over 4,000 m altitude. Based on these temperature records (kept since ca 1980) the temperature lapse rate for the area between Tembagapura and the Mill was calculated to be 0.5C/100 m (Shea et al. 1998). Rainfall is often high and over 6,000 mm/yr has been recorded from some localities; all months tend to receive adequate rain for year-round plant growth. Rain shadow areas in the mountains can have a significantly reduced rainfall. Cloud is frequent in the mid- and upper montane zone at ground level, which has several direct effects. Light is greatly reduced and cloud blown through foliage is strained of much of its moisture (‘‘fog-stripping’’). Crowded foliage, dense crowns, and needle leaves may be adaptations to enhance the efficiency of the process. Midmontane and upper montane forests probably receive a considerable amount of moisture from this source. Fog stripping may be more important in saddles through which winds are channeled than on adjacent ridge crests. Fog will tend to leach nutrients from leaves unless the plants have protective impermeable

M

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

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cuticles. Despite the extreme wetness of atmosphere and soil during fog periods, many sites above the cloud line may be seasonally dry and the vegetation may suffer temporary water stress. Winds tend to travel up-valley by day and downvalley in the early morning due to cold air drainage. Winds are generally not strong and tend to do only localized damage. As noted by Whitmore (1984), soils change with elevation. In general, they become more humic when available mineral nutrients are reduced, especially where peat is present. Peat soils are very acid and perhaps toxic. These elevational changes are correlated with the increasingly cool and moist aerial environment. The tendency for organic matter to accumulate at high elevations is accentuated by wetness, because anaerobic conditions inhibit decay, and the thickness of the organic layer commonly increases dramatically at the cloud line (ca 1,500 m). Soils are typically waterlogged and anaerobic near the surface, becoming better drained at depth. Peaty podzols commonly form as iron becomes reduced to the soluble ferrous state, moves down the profile, and is re-precipitated as a pan. Sphagnum and other peat-forming mosses occur above the cloud line and increase peat formation. Thus, there are a number of self-reinforcing factors, accentuated by frequent fog, which lead to acid and oligotrophic (nutrient poor) conditions in mountain soils. Ridges, knolls, and summits at all elevations are stable sites in which the soil is not being continually rejuvenated by slip or creep. The only water received is from the atmosphere; the soils are continually leached and do not receive soil water from higher sites. It follows that the soils of these places will tend to be more oligotrophic and drier than those of hillsides and valleys. Peat formation may occur in such sites if the soil is poor in litter-decaying organisms, and, as outlined above, once started the process tends to become self-reinforcing.

Previous Studies of Montane Forest in New Guinea and Their Classification Field studies in New Guinea prior to 1940 resulted in a series of accounts in which the altitudinal zonation of the vegetation received attention. Gibbs (1917) published her detailed observations on the flora and vegetation of the Arfak Mountains in Papua. Gibbs (1917) recognized several vegetation types in the Arfak Mountains: lower montane forests dominated by Araucaria and Lithocarpus; a low mountain forest above 2,100 m altitude from Mt Koebre. Upper montane (mossy forest), dominated by Phyllocladus, Papuacedrus, and Dacrycarpus, was confined to the higher forested zones on the ridges. This work was followed by the papers of Lane-Poole who made the first attempt to classify the major plant communities of New Guinea as a whole. Lane-Poole (1925), in his discussion of the foothill forests (lower montane forest), noted that they lack character and, at times, they appeared only a degenerate form of the lowland rainforest. The characteristic tree species of this zone is Castanopsis acuminatissima, but it is associated with Lithocarpus, Araucaria, and Agathis and many distinct species of lowland rainforest genera. Lane-Poole (1925) considered his

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midmontane forests well defined, in that the lower limit corresponds with the ‘‘cloud belt.’’ In summary Lane-Poole (1925) recognized the following forest zones based on his studies of the mountain forests in Papua New Guinea: foothill forests (300–1,700 m); mid-mountain forests (1,700–2,250 m); mossy forests (2,250– 3,350 m), and high-mountain forests (over 3,350 m). These equate to the lower montane, midmontane, upper montane, and subalpine forests as used in this chapter. The 1938–1939 Third Archbold Expedition explored the Snow Mountains, the highest range on the island of New Guinea, with Mt Jaya (Mt Carstensz) rising to 4,886 m elevation (Archbold and Rand 1942; Brass 1941). Of the total 6,000 collections from the Third Archbold Expedition, 4,200 numbers were collected from above 1,000 m altitude, and 1,200 collections above 3,000 m. Prior to the Brass Expedition many collections had been made of plants in Papua (Chapter 1.2). In 1920, Lam accompanied the van Overeem expedition to Doorman Top, an outlying mountain with a distinct ultramafic vegetation. The early expeditions included Beccari’s trips to the Arfak Mountains, the Wollaston and Kloss Expeditions to Mt Jaya in 1912 and 1916, the 1936 expedition of Colijn, and the many Dutch Expeditions including the Lorentz-van Nouhuys expedition in 1909 and the Herderschee expedition in 1913. Plants had previously been collected along the Idenburg River on several expeditions, but mostly at low altitudes. Lam (1945) provided a summary of the vegetation of New Guinea based partly on his visit in 1919–1920 (the van Overeem expedition). He emphasized the difficulties in distinguishing the mountain forest (montane) from the lowland rainforest and also the lack of any sharp boundary between them. No attempt was made by Lam to distinguish altitudinal types within the montane forest. Van Steenis (1934) proposed the following altitudinal zones for the Malesian region, based on analysis of the existing collections of mountain plants (it should be noted that van Steenis never visited New Guinea). His major zones were: tropical zone (0–1,000 m); colline subzone (500–1,000 m); montane zone (1,000–2,400 m); submontane subzone (1,000–1,500 m); and the subalpine zone (2,400–4,000 m). This classification of altitudinal zonation is of little value in eastern Malesia; while it may reflect the zonation patterns in Java, the altiudinal patterns appear distinct in New Guinea. Brass (1941) proposed the following preliminary classification for the major plant communities of New Guinea: savannah and savannah forest (0–1,700 m); monsoon forest (0–450 m); rainforest (0–2,400 m); mid-mountain forest (480– 2,350 m); beech forest (850–3,100 m); mossy forest (1,500–3,200 m); subalpine forest (3,000–4,050 m); and alpine grassland (2,900–snow line). In 1958 Robbins presented a classification of the montane vegetation based on his studies in the Central Highlands of Papua New Guinea, which he adapted from the system proposed by Beard (1955). The terms used have been widely, but not consistently applied and have been the subject of much discussion. Lower montane rainforest (900–3,000 m), a fagaceous alliance in which he included Castanopsis-Lithocarpus (to 2,210 m) and Nothofagus (2,100–2,700 m) forest. His broadleaf-

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gymnosperm alliance included a montane cloud forest formation (2,700–3,330 m) and a high mountain formation described from Mt Wilhelm at 3,600 m. At higher altitudes he recognized an alpine shrubbery formation (3,300–3,600 m); an alpine grassland formation (3,300–3,600 m), the alpine bog formation (3,300–3,600 m) and an herbaceous swamp formation dominated by Phragmites or Cyperaceae. As Robbins (1958) observed, the lower level of the montane zone overlaps broadly with the lowland rainforest. On the Mt Wilhelm massif all forest zones were ca 300 m higher than elsewhere in the Central Highlands. The broadleaf-gymnosperm alliance of Robbins starts at 2,700 m. Below this zone are a variety of rainforest communities extending above ca (300–) 700 m, often containing a large component of montane forest dominants: Lithocarpus, Agathis, Araucaria, and podocarps, often in association with Lauraceae, Podocarpaceae, Elaeocarpaceae, and a host of distinct higher altitude species of common lowland genera. The classification of these rainforest communities is not even considered by Robbins (1958). The Sixth Archbold Expedition (1959) worked in two major areas of Papua New Guinea: Lae to Edie Creek and the Eastern Highlands, based at eight camps from 1,350 to 3,570 m altitude. Brass (1964: 208–211) used the term ‘‘mossy forest’’ for forests that Robbins (1958) called montane cloud forest; these are treated here as midmontane forest. Recent ecological studies in the montane zone of New Guinea have concentrated on the forests dominated by Araucaria and Nothofagus. The Araucaria forests of Papuasia form a distinct forest ranging in altitude from ca 700–1,200 m. The largest extant stands were in the Bulolo-Wau and Watut areas and the Okapa area. The distribution of smaller stands has been plotted by Gray (1973). Womersley (1958) concluded that the Araucaria component was part of the Gondwanan element in the tropical mountains and was being replaced due to invasion of the sites by a more aggressive forest of Malaysian dicotyledonous trees. Recent studies, however, have postulated that Araucaria regenerates in disturbed secondary conditions. A detailed study of the montane forests dominated by Araucaria cunninghamii and A. hunsteinii was conducted by Enright (1982), centered in the Bulolo Valley. He established several ecological plots which, together with data from 0.5 hectare plots previously established by Johns, formed the basis of his studies. The division of the montane vegetation into three zones, the lower montane, the midmontane, and the upper montane, was proposed by Johns (1976). The lower montane is essentially the area of the oak forests, dominated by Castanopsis and Lithocarpus, together with Araucaria, Agathis, and Eucalyptopsis (Johns 1976– 1978). It is the zone in which traditional gardening was practiced in the highlands of New Guinea. The midmontane zone is where Nothofagus often dominates. The lower level of early morning cloud usually marks the lower limit of this forest. The forest is densely covered with epiphytes, particularly orchids and ferns. Above this is the upper montane forest, dominated by Podocarpaceae and Cupressaceae, mixed with many other genera. Epiphytes are abundant but the species diversity is lower than in the midmontane zone. These zones often intergrade; Nothofagus

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often penetrates along ridges into forest areas dominated by Castanopsis and Lithocarpus. Many similar exceptions can be found but generally the zones can be distinguished on their physiognomy, structure, and floristics. These zones provide a simple basis for the classification of the montane vegetation; we describe each in detail below.

Lower Montane Vegetation Lower montane vegetation occurs between (300–) 650 and 1,500 (–1,800) m asl in Papua. The lower boundary is the boundary with lowland rain forest or, in south Papua, with heath forest on dissected Pleistocene terraces. The upper boundary at 1,500–1,700 m asl is generally the upper level of village cultivation of traditional crops, or in the highland valleys the elevation at which clouds tend to intercept the mountain slopes. The characteristic tree species of the lower montane zone is Castanopsis acuminatissima, associated with several species of Lithocarpus and Araucaria. Many rainforest genera reach their upper limit at 2,200–2,500 m altitude. It was noted that Castanopsis and Lithocarpus rarely occurred above the upper limits of cultivation: even at high elevations (2,350 m) small patches were markedly dry in appearance. At 1,700 m there is a transition to mossy midmontane forest characterized by Nothofagus forest. Families which are characteristic of the lowland zone are also common in the lower montane subzone, but become less frequent and less dominant with elevation and are usually represented by a distinct group of ‘‘montane’’ species. This group includes montane species of the lowland families: Anacardiaceae, Annonaceae, Burseraceae, Dipterocarpaceae, Ebenaceae, Euphorbiaceae, Guttiferae, Leguminosae, Meliaceae, Myristicaceae, Myrtaceae, Rubiaceae, Sapindaceae, Sapotaceae, and Sterculiaceae. There is an increasing importance of members of the families Cunoniaceae, Elaeocarpaceae, Fagaceae, Lauraceae, and Podocarpaceae with elevation. There is a considerable range of vegetation types within the lower montane subzone. The structure and floristic composition of these vegetation types is determined by a series of environmental factors. Brass (1941) noted that rainforest at 1,200 m included several species which were common at 850 m, and even some species which occurred in the lowland rainforest of the Idenburg River. Species from lower altitudes did not extend their range much above 1,200 m. Several tree and subcanopy species occupied the range from ca 850–1,200 m. In contrast, the bryophytes, ferns, and orchid epiphytes occurred in the mossy beech forests at higher altitudes. A feature of these forests was the prominence of Syzygium and also of members of the Lauraceae (Cryptocarya, Endiandra, Litsea) and Schizomeria. Podocarpus sensu stricto was locally common, as are many high altitude representatives of common lowland genera such as Myristica, Calophyllum, Elaeocarpus, Horsfieldia, Gordonia, Adinandra, and others. Galbulimima belgraveana is locally common. As noted by Johns (1982), these lower montane forests differ markedly from the forests in the lowland zone. The frequency of palms, lianas, and trees with

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buttress roots is much reduced. Tree ferns, filmy ferns, and bryophytes (mosses) are less common. The canopy of the forest, 20 to 25 m in height, is made up of a number of broadleaf species. There is a wide variation in the composition from area to area. Typical canopy genera are Elaeocarpus, Sloanea, Elmerrillia, Litsea, and Cryptocarya. In the subcanopy, common genera include Elaeocarpus, Weinmannia, Pullea, Myristica, Daphniphyllum, Diospyros, and Pittosporum. Tree ferns are less common than in the mid- and upper montane forests. Several species of climbers and scramblers are present, including species of the genera Parsonsia, Clematis, Smilax, and Hoya. Epiphytes present include Schefflera, orchids, and parasitic Loranthaceae. The lower montane forest is quite distinct from the two neighboring zones, particularly in regard to the physiognomy of the vegetation (Johns 1976). While many of the species occur in the midmontane subzone, they are rare and seldom dominate in this higher zone. In comparison to the midmontane zone, podocarps are less abundant in the lower montane. The dominant tree genera of the midmontane subzone are Nothofagus and Cunoniaceae; they are seldom found in the lower montane. The lower montane subzone is distinctly drier in aspect than the midmontane, with ferns and mosses being less abundant with fewer species. Pandanus occurs locally in wet areas, but never forms a regular component of the forest as it often does with Nothofagus. The species composition of these lower montane stands varies from site to site. A typical stand is dominated by Elaeocarpus, Sloanea, Elmerrillia, and Litsea. The lower montane forest is less rich in species than in the midmontane zone, and trees of the oak family (Castanopsis and Lithocarpus), the Elaeocarpaceae (Elaeocarpus and Sloanea) and the laurel family (Litsea and Cryptocarya) are prominent. Changes in structure and floristics are due to a combination of factors. The change in average air temperature with altitude (about 0.5C/100 m) affects the ability of plants to survive, reproduce, and compete with other plants. There are also indications that cloud cover may also be an important factor. Brass (1964), for example, observed that throughout Papua New Guinea the lower limit of Castanopsis-Lithocarpus forest coincides with the lower edges of the afternoon cloud body. In the season dominated by south-easterly winds this forms almost daily at remarkably constant and well-defined levels on the hill and mountain slopes. This is consistent with studies in other parts of the tropics. For example, in Ecuador, Grubb and Whitmore (1966) studied the changes from lowland rainforest to montane forest, and suggested that these are mainly controlled by the frequency of cloud cover close to the ground. As a result, Castanopsis and Lithocarpus may occur at low elevations, where rainfall is high and humidity is constantly high. Forest structure is often related to topography. For example, forests on steep and unstable slopes with thin soils often have open and irregular canopies. On such slopes, tree dimensions are smaller than in valley sites, and many trunks are leaning or bent at the base. Here, soil erosion may cause roots to become exposed, and some trees develop adventitious roots. In contrast, forests on gentle slopes,

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plateaus, and foot slopes tend to have above average height, tree girth, and crown size. Where drainage and moisture conditions are favorable, lower montane forests may approach lowland alluvial forests in structure. Ridge crests often have a denser forest than side slopes. Some trees, such as Araucaria, tend to be concentrated on crests. Other trees, such as Pometia, are more common on foot slopes than on mid- and upper slopes. Less obvious floristic variation correlated with topography can be detected by quantitative analyses and has been clearly demonstrated in forests in the Solomon Islands, Malaysia, and Borneo. The lower montane vegetation in high rainfall areas throughout Papua is different from the vegetation in lower rainfall areas. The areas studied by Shea et al. (1998) in south Papua are in a high rainfall area. In other parts of New Guinea, where annual rainfall is between about 1,200 and 1,800 mm and where there is a distinct dry season, the forest may have deciduous and semi-deciduous trees in the canopy. The lower stories, however, remain evergreen. Such forest is similar in structure to its evergreen counterpart except that the canopy is somewhat more open, and scrambling bamboo is a normal feature. Common deciduous trees are Garuga floribunda, Intsia bijuga, Anisoptera thurifera, Pterocarpus indicus, and Sterculia spp. In the shrub layer, Maniltoa, Lunasia amara, Cycas, and Desmodium ormocarpoides are markedly more frequent than in evergreen lower montane forest. Scattered grasses, sedges, and ferns form a sparse ground cover. In areas where the annual rainfall is less than about 1,200 mm and the dry season is long and severe, the forest has a low and open canopy dominated by deciduous trees. The tree story below the canopy consists of deciduous and evergreen trees, and in the undergrowth many shrubs are spiny and scrambling. Flagellaria and thin woody lianas, some of which have cork ribs or spines, abound, but epiphytes are scarce. Deciduous trees, in addition to those mentioned above, include Gyrocarpus americanus, Bombax ceiba, Brachychiton carruthersii, Adenanthera pavonina, and Erythrina sp. The severity of the dry season is probably the main factor controlling the frequency of deciduous trees, but altitude and local variation in topography and moisture conditions also play a part. In general, deciduous trees decrease in number with increasing altitude, so that forests approaching the midmontane zone are virtually evergreen. Soil appears to have less influence on the structure and floristic composition of lower montane forest than do altitude, topography, and climate. In New Guinea, however, evidence suggests that forest of below-average stem diameter and crown size is correlated with the presence of strongly weathered, acid clay soils, which are fairly common on gently to moderately sloping low foothills and lower mountain slopes. The soil itself does not appear to influence the floristic composition of the forest. However, any suspected correlation would be hard to prove because of the complex interaction among soil, topography, and climate, and because of other factors such as previous disturbance and chance establishment, which may override and mask any influence of soil. Quantitative analyses have shown that species composition is associated with soils on some sites (Austin, Ashton,

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and Greig-Smith 1972, cited in Whitmore 1984a), while other studies have shown no correlation between species distribution and soils. Rock type influences the vegetation through the soils and land forms developed on it. Ultramafic rocks (igneous heavy rocks with low silicon and relatively high magnesium and iron contents) and limestones are of special interest. In New Guinea, ultramafic rocks appear to be correlated with the presence of dense, thinstemmed, small-crowned and often open forests. Ultramafic rocks often have topography of steep, straight slopes with unstable, shallow and chemically poor soils, and hence have thin-stemmed forest. However, even on sites with gentle topography and deep soils, forests on ultramafics tend to be denser and smaller crowned than average hill forest. Shallow soils also predominate on limestone, but are less infertile and penetrate deeper into solution hollows in the rock. In New Guinea, there is no evidence that any tree species is confined to either ultramafics or limestone. Casuarina papuana is commonly predominant in forest on ultramafic rocks, and occasionally in forest on limestone, but this species has a wide ecological range and is not restricted to any particular rock type or soil.

mixed lower montane forest The mixed lower montane forest that occurs on the foothills and lower slopes of the mountains of Papua is structurally and physiognomically similar to alluvial lowland forests. According to Paijmans (1976), it differs from alluvial lowland forests by being lower in stature due to the less favorable conditions of steep slopes and shallow unstable soils. Main canopies are 25–30 m high, while emergents may reach 40 m or more in height. Trees with a very large girth and large buttresses are less common than in the lowland forests. However, there may be more trees with a girth over 30 cm dbh, and there may be more trees in the pole and sapling stages. The shrub layer, consisting mainly of slender saplings, has a lower cover, but the herb layer, though patchy, is generally somewhat denser. Thick woody lianas, rattan, tree- and shrub-palms, and fleshy climbers and climbing ferns on tree trunks are less common. Tall palms are a normal feature in the canopy and locally emerge above it, but they usually occur in small numbers only. Tree ferns are more common than in the lowland rainforests, as also are scrambling bamboos, especially on ridge crests. Like forest on alluvium, these forests are very rich in species and very mixed. Most trees present in alluvial forest also occur in hill forest, although in many cases in different proportions. Frequent canopy trees are Pometia, Canarium, Anisoptera, Cryptocarya, Terminalia, Syzygium, Ficus, Celtis, Dysoxylum, and Buchanania. Many others are also common. Some tree genera common in the understories are Garcinia, Syzygium, Diospyros, Myristica, Pimeleodendron, Microcos, and Gnetum. Lithocarpus celebicus first appeared at 120 m and became prominent above 350 m altitude in the Idenburg Valley. At altitudes around 1,200 m, Lithocarpus and Castanopsis formed pure ridge top stands on broader crests. The oaks are well spaced with a thin canopy, but despite the abundant illumination of the forest

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floor, substage trees and ground plants are uncommon. The forest has a distinctly dry appearance, with a few xeromorphic ferns such as Syngramma, some epiphytes such as Vittaria and Oleandra, and a few climbers including Dimorphanthera. This forest showed a distinct preference for crests; mixed rainforest dominated the slopes and the valley floors. In south Papua examples of the mixed lower montane forest were described by Shea et al. (1998) from the mountain slopes above the heath forest at Mile 50. Two closed forest stands were sampled. The first stand was located east of the Freeport Road on a slope above Mile 50. The dominant trees were Elaeocarpus, Tristania, Lithocarpus, and Hopea papuana. Other trees included Crypteronia, Palaquium, Syzygium, Cryptocarya, Sloanea papuana, and Elaeocarpus (Table 5.10.1). The estimated total basal area for trees with a diameter greater than 10 cm dbh is 36.6 m2/ha and the total clear bole volume is 440.1 m3/ha. The second stand is located on a spur east of the road above Mile 50 in south Papua. The dominant trees are Elaeocarpus, Lithocarpus, Syzygium, and Intsia. Other common trees include Cryptocarya, Anisoptera thurifera, and Castanopsis acuminatissima (Table 5.10.2). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 38.2 m2/ha and the total clear bole volume is 410.8 m3/ha. A third stand was sampled on steep upper slope of the Otomona River Gorge at Mile 50, south Papua. The dominant trees were Lithocarpus, Anisoptera thurifera, Castanopsis acuminatissima, and Syzygium. Other genera recorded were palm trees, Palaquium, Hopea papuana, Timonius, Cryptocarya, and Haplolobus floribundus (Table 5.10.3). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 43.9 m2/ha and the total clear bole volume is 436.8 m3/ha.

secondary forest An interesting secondary forest occurs on wet slope east of PT Freeport Indonesia Road at Km 71–72 at about 650–700 m asl in south Papua. The stand is dominated by pioneer and secondary forest species such as Casuarina, Ficus, Anthocephalus cadamba, Ilex, Pandanus, Diospyros, Gordonia, Timonius timon, and T. trichanthus (Table 5.10.4). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 36.9 m2/ha and the total clear bole volume is 181.1 m3/ha.

castanopsis and lithocarpus forests Robbins (1958) treats Castanopsis and Lithocarpus as a single association. Here we question this treatment. A common pattern in the Bulolo-Watut area is for the Castanopsis forest to form distinct stands on disturbed sites (often old garden sites), whereas species of Lithocarpus usually occur in more diverse forest, in association with rainforest species, or more accurately, higher altitude species of lowland rainforest genera. Brass (1941) concluded that in the mountains of New Guinea ‘‘man follows the oaks,’’ because it is in the cool, healthy montane valleys

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440.00

Total

450

25

25

25

25

25

25

25

50

50

50

25

25

25

Absolute density

Note: Stand located above Mile 50 along PTFI Road, south Papua. Source: Shea et al. (1998).

106.82

6.68

Tristania sp.

56.38

Cyptocarya sp.

11.11

Palaquium sp.

Lithocarpus sp.

1.11

Polyosma sp. 8.24

1.85

Xanthophyllum sp.

Sloanea papuana

7.50

Symplocos cocchinchinensis

166.91

10.87

Elaeocarpus sp.

13.47

Syzygium anomala

0.58

41.21

Crypteronia sp.

Syzygium lauterbachianum

Hopea papuana sp.

Species

Total clear bole volume (m3/ha)

100.00

5.56

5.56

5.56

5.56

5.56

5.56

5.56

11.11

11.11

11.11

5.56

5.56

5.56

Relative density (%)

36.60

9.54

1.06

4.24

0.73

1.44

0.26

0.26

0.72

11.98

1.21

1.44

0.12

2.94

Absolute dominance (m2/ha)

100.00

26.02

2.89

11.57

2.01

3.94

0.72

0.72

2.00

32.53

3.29

3.94

0.32

8.03

Relative dominance (%)

1.13

0.06

0.06

0.06

0.06

0.06

0.06

0.06

0.13

0.13

0.13

0.06

0.06

0.06

Absolute frequency

100.00

5.56

5.56

5.56

5.56

5.56

5.56

5.56

11.11

11.11

11.11

5.56

5.56

5.56

Relative frequency (%)

Table 5.10.1. Mixed lower montane forest dominated by Elaeocarpus, Tristania, Lithocarpus, and Sloanea papuana

300.00

37.14

14.00

22.68

13.12

15.05

11.83

11.83

24.20

54.75

25.52

15.05

11.43

19.14

Importance value

986 / r. j. johns , g . a . shea, w . v i n k, & p . puradyatmika

PAGE 986

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2.64

Hopea novoguineensis

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1075

25

25

25

25

125

25

25

200

25

25

25

25

25

25

25

25

Absolute density

100.00

2.33

2.33

2.33

2.33

11.63

2.33

2.33

18.50

2.33

2.33

2.33

2.33

2.33

2.33

2.33

2.33

Relative density (%)

Note: Stand located on a spur above Mile 50 along PTFI Road, south Papua. Source: Shea et al. (1998).

410.85

Total

2.23 111.27

Lithocarpus sp.

Engelhardtia glauca

Elaeocarpus sepikensis 11.13

84.40

Elaeocarpus sp.

Anisoptera thurifera

4.04 68.31

Vaccinium sp.

4.45

30.84

Cryptocarya sp.

Symplocos cochinchinensis

0.49

1.48

Polyosma sp.

Diospyros

4.12

15.15

Microcos florida

Syzygium argentea

0.41

4.29

Castanopsis acuminatissima

Euodia sp.

3.96

Syzygium aqueanum

Species

Total clear bole volume (m3/ha)

38.24

6.62

0.26

1.06

7.54

6.09

1.44

1.06

3.36

0.12

0.47

0.26

0.74

1.44

0.12

0.47

0.47

Absolute dominance (m2/ha)

100.00

17.32

0.69

2.77

19.71

15.94

3.77

2.77

8.80

0.31

1.23

0.69

1.92

3.77

0.31

1.23

1.23

Relative dominance (%)

2.44

0.06

0.06

0.06

0.06

0.25

0.06

0.06

0.31

0.06

0.06

0.06

0.06

0.06

0.06

0.06

0.06

Absolute frequency

Table 5.10.2. Mixed lower montane forest dominated by Elaeocarpus, Lithocarpus, Syzygium, and Intsia

100.00

2.56

2.56

2.56

2.56

10.26

2.56

2.56

12.8

2.56

2.56

2.56

2.56

2.56

2.56

2.56

2.56

Relative frequency (%)

300.00

22.21

5.58

7.66

24.60

37.82

8.66

7.66

40.20

5.20

6.12

5.58

6.81

8.66

5.20

6.12

6.12

Importance value

Montane Vegetation of Papua / 987

PAGE 987

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2.01

Xanthophyllum sp.

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13.74 436.76

1066.66

33.33

33.33

33.33

33.33

33.33

33.33

33.33

33.33

33.33

133.33

33.33

100.00

33.33

33.33

66.67

33.33

200.00

133.33

Absolute density

100.00

3.13

3.13

3.13

3.13

3.13

3.13

3.13

3.13

3.13

12.5

3.13

9.38

3.13

3.13

6.25

3.13

18.75

12.5

Relative density (%)

43.89

1.64

1.05

0.59

0.26

1.64

1.05

0.26

2.35

1.64

3.86

0.26

1.11

1.05

0.59

10.47

0.26

9.75

6.08

Absolute dominance (m2/ha)

Note: Stand located on slopes of the Otomona River Gorge at Mile 50 along the PTFI Road, south Papua. Source: Shea et al. (1998).

Total

Hopea papuana

9.52

Haplolobus floribundus

Palaquium sp. 3.30

14.88

Cryptocarya sp.

Elaeocarpus sp.

8.06

Sloanea sp.

1.47

1.47

Alstonia sp.

Euodia sp.

16.03 18.13

Araceae

28.99

7.28

Timonius sp.

Syzygium sp.

13.92

4.12

162.65

Nibun (Palm)

Engelhardia glauca

Lithocarpus sp.

1.83

80.68

Anisoptera thurifera

Myristica fatua

48.68

Castanopsis acuminatissima

Species

Total clear bole volume (m3/ha)

100.00

3.73

2.38

1.34

0.60

3.73

2.38

0.60

5.37

3.73

8.79

0.60

2.53

2.38

1.34

23.85

0.60

22.21

13.86

Relative dominance (%)

2.17

0.08

0.08

0.08

0.08

0.08

0.08

0.08

0.08

0.08

0.17

0.08

0.17

0.08

0.08

0.17

0.08

0.33

0.25

Absolute frequency

100.00

3.85

3.85

3.85

3.85

3.85

3.85

3.85

3.85

3.85

7.69

3.85

7.69

3.85

3.85

7.69

3.85

15.38

11.54

Relative frequency (%)

300.00

10.70

9.36

8.31

7.57

10.70

9.36

7.57

12.34

10.7

28.99

7.57

19.60

9.36

8.31

37.79

7.57

56.34

37.90

Importance value

Table 5.10.3. Lower montane forest dominated by Lithocarpus, Anisoptera thurifera, Castanopsis acuminatissima and Syzygium

988 / r. j. johns , g . a . shea, w . v i n k, & p . puradyatmika

PAGE 988

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84.85 20.88

Anthocephalus cadamba

Ficus sp.

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0.37 0.73

Gordonia sp.

Diospyros sp.

PS

1499.99

33.33

33.33

33.33

33.33

33.33

400.00

400.00

366.66

66.67

100.00

Absolute density

Note: Stand located at Km 71–72 along PTFI Road, south Papua. Source: Shea et al. (1998).

181.10

0.92

Ilex sp.

Total

1.24

Timonius trichanthus

5.13

55.18

Casuarina

Timonius timon

1.24

10.58

Pandanus

Pandanus sp.

Species

Total clear bole volume (m3/ha)

100.00

2.22

2.22

2.22

2.22

2.22

26.67

26.67

24.44

4.44

6.67

Relative density (%)

36.89

0.26

0.26

0.26

0.59

1.05

6.80

12.56

11.58

1.64

1.90

Absolute dominance (m2/ha)

100.00

0.71

0.71

0.71

1.60

2.84

18.44

34.04

31.38

4.43

5.14

Relative dominance (%)

Table 5.10.4. Secondary forest dominated by Anthocephalus, Casuarina, Timonius, and Ficus

1.83

0.08

0.08

0.08

0.08

0.08

0.58

0.42

0.08

0.17

0.17

Absolute frequency

100.00

4.55

4.55

4.55

4.55

4.55

31.82

22.73

4.55

9.09

9.09

Relative frequency (%)

300.00

7.48

7.48

7.48

8.36

9.60

76.92

83.44

60.37

17.97

20.90

Importance value Montane Vegetation of Papua / 989

PAGE 989

990 / r. j. johns , g . a . shea, w . v i n k, & p . puradyatmika

that the greatest concentration of people occurs. The large valley systems now form grassy montane valleys, where once the Castanopsis and/or Lithocarpus forest was dominant. These forests are referred to as ‘‘oak forests’’ because of the important role played by Castanopsis and Lithocarpus. Castanopsis acuminatissima and Lithocarpus spp. may dominate in ridge-top forest at various altitudes above 600 m but seldom occur in stands with Nothofagus. Other tree species mixed with the oaks are Anisoptera, Hopea, Intsia, Pometia, Calophyllum, and Syzygium, particularly on lower slopes. In the Idenburg Valley the lower montane vegetation was dominated by Lithocarpus rufovillosus, L. lauterbachii, and Castanopsis acuminatissima; these formed a canopy at ca 30 m (Brass 1941). Unlike many areas of oak forest in Papua New Guinea, the canopy was conspicuously mossed, thus supporting a very rich and abundant epiphytic flora. Ferns were particularly important, including species of Arthropteris, Meringium, Polypodium, Goniophlebium, Leucostegia, Nephrolepis, Asplenium, Lindsaea, Hymenophyllum, and Loxogramme. Medinilla, Ophiorrhiza, Elatostematoides, Rhododendron macgregoriae, and Nepenthes were also common. Along the streams a series of rheophytes occurs including Ficus spp., Syzygium, Actinophloeus (clump palm), and Osmoxylon. Common herbaceous plants and ferns are Dryopteris, Lindsaya, Selaginella, Hemigraphis, and the grasses Pogonatherum and Isachne, and the herb Impatiens in disturbed pioneer communities. These were not dominated by Macaranga but by species of Saurauia, Wendlandia, Schuurmansia, and Ficus dammaropsis, with Cyathea contaminans a prominent later species in the succession. The secondary stands, which reached 25 m in height, were dominated by Homolanthus and Albizzia. Ground flora was dominated by Nephrolepis, Dryopteris, Elatostema, and Spathoglottis. In the Baliem Valley most of the oak forest had been cleared for traditional agriculture. This occurs at ca 2,400 m asl. Patches of oak forest occur throughout the Baliem Valley up to 2,200 m altitude; Brass (1941) assumed that prior to settlement the valley would have supported an extensive fagaceous forest. Casuarina forms extensive stands in the Baliem Valley at ca 1,600 m, especially along rivers and in landslide areas. Old grass slopes contain a number of ‘‘fire resistant’’ shrubs: Vaccinium, Alphitonia, Grevillea, Timonius, Decaspermum, and Glochidion. On limestone knolls more xerophytic elements dominate: Grevillea, Rhamnus, Eurya, and some species of Schefflera are common. A species of Lithocarpus dominated the swampy ground in the valley, with an undergrowth of Calamus and young Pandanus. Casuarina sumatrana formed distinctive stands up to 1,200 m, both on steep ridges and sandy creekbanks. Agathis formed a distinctive ridge top forest (i.e., an open stand with a subcanopy of Metrosideros, Quintinia, and Daphniphyllum). Brass (1941) noted that Agathis canopies were covered in moss and a rich associated epiphyte flora, with many species in common with the Nothofagus forests at higher altitudes. Vink (pers. comm.) observed that the canopy of Agathis is emergent and its crowns are always very clean. Agathis reached its upper limit at ca 1,200 m.

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1. Results map from the Irian Jaya Biodiversity Conservation Priority Setting Workshop, held 7–12 January 1997. (CI)

2. The WWF Ecoregions for the island of New Guinea. (Conservation Science Program, WWF–US)

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3. Aerial view of Lake Kamaka. the largest of the Triton Lakes, near Kaimana. These support eight endemic fish species. (G. Allen)

4. Aerial view of Lake Sentani. Grasslands around the lake are a product of burning in this low rainfall area. (B. M. Beehler)

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5. A dense school of Sweetlips Plectorhinchus polytaenia in the Dampier Strait, indicative of Raja Ampat’s healthy fish stocks. (M. V. Erdmann)

6. Aerial view of the specatular Karst island formations of the Wayag Group north of Waigeo Island, in the Raja Ampat. (M. V. Erdmann)

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7. Rainbowfishes are a dominant group in New Guinea and Australia. The Irian Jaya Rainbowfish Melanotaenia irianjaya ranges throughout the southern Bird’s Head Peninsula. (G. Allen)

8. Bostrichthys strigogenys (12 cm total length), a freshwater member of the gudgeon family (Eleotridae), which occurs in the southeastern part of Papua (G. Allen)

9. Price’s Damselfish Chrysiptera pricei is known only from shallow coral reefs of Cenderawasih Bay. (G. Allen)

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10.Coral reefs of the Bird’s Head Seascape are among the richest on Earth for marine fishes, containing a host of endemic species, including the Cenderawasih Fairy Wrasse Cirrhalabrus cenderawasih. (G. Allen)

11. An undescribed species of epaulette shark Hemiscyllium sp. discovered during a recent biological survey in Cenderawasih Bay. (G. Allen)

12. Aerial view of the Mamberamo River. (J. B. Burnett)

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13. Aerial view of the tidal and freshwater swamp forests and rivers of the Asmat/ Lorentz area. (J. B. Burnett)

14. Typical treefern-dotted alpine grasslands of the central cordillera. (B. M. Beehler)

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15. Rugged alpine peaks of the Puncak Jaya Range of western Papua. (M. P. Moore)

16. Sphagnum bog at 1,650 m in the Foja Mountains, the site of the 2005 biodiversity survey of the Fojas. (S. J. Richards)

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17. Unbroken hill and montane forest of the Foja Mountains of northern Papua. (S. J. Richards)

18. Aerial view of two logging mills and logging roads in northern Papua. Selective logging is widespread in northern Papua west of Jayapura. (Scott Frazier/CI)

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19. Interior of wet montane mossy forest of the Foja Mountains. (S. J. Richards)

20. Moss-encrusted forest floor of the montane zone of the Foja Mountains. (S. J. Richards)

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21. Sphagnum bog at 1,650 m in the Foja Mountains. Site of the Bog Camp for the 2005 expedition to the Fojas. (S. J. Richards)

22. Summit glaciers of the Puncak Jaya Range. (M. P. Moore)

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23. Wapoga mountain forest view from helipad. (M. P. Moore)

24. Vista from high in the Wapoga Range. (M. P. Moore)

25. Bulbophyllum macneiceae Schuit. & de Vogel (Port Moresby Botanic Gardens). Species of Bulbophyllum sect. Epicrianthes have small but spectacular flowers. They are probably insect-pollinated but this has never been observed. Most are known only from very few specimens. This species is named in honor of Lady McNeice of Singapore, an ardent supporter of New Guinea orchid research, and a devoted nature conservationist. (E. F. de Vogel)

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26. Cycads in Wasur National Park, southeastern Papua. (J. B. Burnett)

27. Dendrobium alexandrae Schltr. Leiden cult. 20030942. Most species of Dendrobium section Latouria have long-lasting and spectacular-looking flowers and are much in demand for hybridizing and cultivation. This species is present in many horticultural collections, but only three collections are in herbaria. (E. F. de Vogel)

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28. Dendrobium chrysopterum Schuit. & de Vogel. With about 400 species, Dendrobium is the second largest genus in New Guinea. From a horticultural point of view, it is by far the most important genus in New Guinea. (E. F. de Vogel)

29. Nepenthes pitcher plant in a sphagnum bog at 1,650 m, Foja Mountains. (S. J. Richards)

30. Paphiopedilum wilhelminae L.O. Williams. Leiden cult. 20020344. New Guinea harbors four species of Paphiopedilum, all known from a few populations only. This genus is highly sought after by collectors, and the species are all seriously endangered. (E. F. de Vogel)

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31. Rhododendron cf. macgregoriae, Foja Mountains. (B. M. Beehler)

32. Asaroe sp. fungus. (M. P. Moore)

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33. A limacodid caterpillar feeding on Macaranga tsonane (Euphorbiaceae) in lowland rain forest, East Sepik Province, Papua New Guinea. (G. Weiblen)

34. A large cerambycid beetle. (M. P. Moore)

35. A recently emergent cicada. This group is abundant and diverse on the island of New Guinea. (M. P. Moore)

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36. Hercules moth Coscinocera hercules caterpillar feeding on Homalanthus. (M. Janda)

37. A cryptically patterned mantis. (M. P. Moore)

Montane Vegetation of Papua / 991

Brass (1941) described a typical pure oak forest. Single species ‘‘oak’’ forests in the lower montane are usually dominated by Castanopsis acuminatissima often regenerating by suckers, which grow to form pure stands, usually following subsistence agriculture or local cyclonic events. The oak forest with Lithocarpus schlechteri, L. rufovillosus, Castanopsis acuminatissima, and Engelhardtia rigida form distinct forests in the ravines to 50 m from the summits. Also occurring in these ravines are elements of the mixed forest: Syzygium, Garcinia, Elaeocarpus, and various Lauraceae such as Cryptocarya. These ravines are often very diverse in their fern flora; one studied by Brass had more than 60 species.

Castanopsis Forest Castanopsis is represented by a single species C. acuminatissima in Papuasia. Castanopsis forest occurs over an altitudinal band from 300–1,500 m, commonly forming pure ridge-top stands, particularly in sites prone to natural disturbance, especially caused by wind throw (Johns 1986). In areas of high human population pressure, this species often dominates in the regeneration following gardening (probably due to its ability to sucker from the trunks). Castanopsis forest is characteristic of disturbed sites throughout its range. The forest is typically dry in aspect with few ground plants or subcanopy trees, despite the apparent high levels of light penetration through the canopy. The trunks and branches are virtually free of epiphytes, other than species particularly adapted to drier conditions (such as Pyrrosia). In some areas ant plants are common (particularly Myrmecodia and Hydnophytum), but this group tends to be more abundant at higher altitudes. Castanopsis acuminatissima forms almost pure stands on ridge crests and upper slopes between about 500 m and 2,300 m altitude. Lower montane forest dominated by Castanopsis has a dense and even canopy, an open shrub layer, a very sparse ground layer of herbs, and a thick carpet of fallen leaves on the forest floor. Castanopsis coppices freely from the base of the trunk, so many trees have a ring of coppice shoots around the main trunk, or branch out into several stems at or near ground level. Johns (1976) includes data from Castanopsis stands in the Bulolo Valley in Papua New Guinea.

Lithocarpus Forest Lithocarpus species (especially Lithocarpus schlechteri and L. rufovillosus) may form single-species stands on some sites. Lower montane forest dominated by Lithocarpus rufovillosus is scattered throughout New Guinea and the species have been recorded in Papua on Yapen Island and Misool Island. L. rufovillosus forms pure stands on sites that were previously gardened on the slopes of Mt Michael in Papua New Guinea. Lithocarpus lauterbachii was locally common in dry mossless forest at ca 2,100 m in the Idenburg Valley (Brass 1941) with a very open undergrowth of herbaceous plants. Alpinia was common, mixed with Gleichenia linearis and Nephrolepis acuminata, and species of Freycinetia were common as epiphytes.

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992 / r. j. johns , g . a . shea, w . v i n k, & p . puradyatmika

mixed oak forest Mixed lower montane oak forest is dominated by Castanopsis acuminatissima, Lithocarpus schlechteri, Litsea, and Elaeocarpus. Brass (1941) described a typical pure oak forest dominated by Lithocarpus and Castanopsis as about 25 m high, with slightly spurred grayish trunks sometimes surrounded by coppice shoots. The oaks grew well apart and formed a rather thin canopy. In spite of the abundant illumination, secondary and sub-stage trees were few. Apart from the oaks, the dominant trees identified by Brass were Pittosporum ramiflorum and Calophyllum papuanum. Typical subcanopy species are Phyllocladus hypophyllus and Helicia amplifolia; common shrubs are species of Rhododendron and Vaccinium. Johns (1982) noted that compound leaves are rare in oak forest and the average leaf size is much reduced from that in lowland rainforest. Lianas are uncommon and buttressed trees rare. Tree ferns such as Dicksonia (rare) and Cyathea (many species) are found, and the ground cover consists of species of Hymenophyllum and mosses such as Polytrichum. At higher elevations (1,200–1,500 m asl), near the boundary with the midmontane subzone, ridges may be dominated by a mixture of Castanopsis, Lithocarpus, and Engelhardtia but are rarely mixed with Nothofagus. Mixed lower montane stands with Castanopsis acuminatissima and several species of Lithocarpus, including L. schlechteri, often occur in association with Litsea and Elaeocarpus. Johns (1976, 1982) noted that compound leaves are rare and the average leaf size is much reduced from that in lowland rainforest. Lianas are uncommon and buttressed trees rare. Tree ferns such as Dicksonia (rare) and Cyathea (many species) are found and the ground cover consists of many species of filmy ferns (Hymenophyllum, Trichomanes) and mosses such as Polytrichum. Near the boundary with the midmontane subzone, lower ridges may be dominated by a mixture of Castanopsis, Lithocarpus, and Engelhardtia, and more exposed ridges by Nothofagus. Although forest communities dominated by the two groups often appear contiguous, such as in the Kainantu-Okapa area in Papua New Guinea, detailed study indicates pronounced ecological differences. Where the genera are contiguous the usual pattern is for Nothofagus to dominate the ridge situations, adjacent lower ridges and upper valley sides are dominated by the Castanopsis and Lithocarpus.

araucaria forests Araucaria cunninghamii var. papuana has a discontinuous distribution in New Guinea from the Vogelkop in Papua to the Milne Bay Province in the southeast of Papua New Guinea. Most large stands occur in the Central Highlands of Papua New Guinea, the Bulolo Valley, and the Owen Stanley Range, all within the altitudinal zone from 500 m to 2,450 m. A. cunninghamii has been recorded from a number of different habitats. In the Bulolo area, where the tallest trees occur, it occurs on slopes and ridges in the hill and lower montane forest associated with many species. At Pai-awa, the trees are of variable morphology and occur at very low altitude on serpentine sites. At Telefomin, the trees appear stunted, and at Erave, the trees occur in a swamp.

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PAGE 992

Montane Vegetation of Papua / 993

In the Anggi basin, particularly to the southeast of Lake Anggi, Gibbs (1917) observed gregarious stands of Araucaria cunninghamii var. papuana (as A. beccarii). Araucaria formed mixed stands, the young stands on marshy ground having an impenetrable undergrowth, while on the ridges with older trees the understory was absent. The forest originally extended to the water’s edge but Gibbs (1917) reported that it had been burnt for some 10 m above the lake. Shea et al. (1998) reported stands on ridges from mountain areas in Lorentz National Park. A. cunninghamii is emergent over mixed forest species characteristic of the lower montane. The detailed description of Araucaria forest is based on a literature review, especially Enright (1982). Two species of Araucaria occur naturally in New Guinea: Hoop Pine (A. cunninghamii) and Klinkii Pine (A. hunsteinii). Regeneration patterns of the two Araucaria species differ markedly. A. hunsteinii does not regenerate successfully under a closed canopy where light intensity is low or the litter layer is very deep. Preferred areas are those of local disturbance (e.g., tree fall). In the center of such openings, often approximately 30 m wide, early secondary species such as those of Macaranga, Homalanthus, Mallotus, and Pipturus grow up rapidly, and A. hunsteinii seedlings are unable to compete successfully. They do succeed, however, along the margins of the gaps where light intensity has been increased substantially, but not enough to allow invasion of the fast growing species noted above. Here, the A. hunsteinii seedlings survive in large numbers and some are ultimately recruited to the canopy. In contrast, A. cunninghamii seedlings do not appear able to survive under low light conditions for long periods of time. This is borne out by the data from the Kebar Valley (Mangold 1959). A. cunninghamii occurs most commonly above 1,000 m in cooler, wetter forests than A. hunsteinii. These populations are growing in highly acidic (pH ⬍ 5.0), slightly podzolized soils low in nutrients. A. hunsteinii is found in dry Pouteria luzoniensis forests, where nutrient concentrations, especially of calcium, are high and pH is neutral to basic, and in moist Flindersia pimenteliana and Gnetum gnemom forests where nutrient concentrations are slightly lower and soils more acidic (pH 5.5 to 6.5). This species does not occur on highly acidic, nutrient-poor soils and is rarely present in large numbers at altitudes above 1,000 m. Analysis of quantitative data (density, basal area) supports the claim that there are major disjunctions between forest types on the altitudinal gradient between 700 and 1,500 m (Enright 1978). The main disjunction divides forests containing many lowland rainforest species from ‘‘oak’’ forest, which is characterized by the presence of the fagaceous genera Castanopsis and Lithocarpus. A. cunninghamii is more common than A. hunsteinii in oak forest.

Succession in Araucaria Forest The two Araucaria species have distinctive regeneration strategies. A. cunninghamii appears better adapted to stable conditions (‘‘K-selected’’). A. hunsteinii has fewer, larger seeds and a slower growth rate than A. cunninghamii, features of a ‘‘K-’’ rather than a ‘‘r-selected’’ tree species. The large seeds of A. hunsteinii have a low

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dispersal ability (Havel 1965) but because the frequency of small-scale disturbances such as tree fall is high, the species is able to take advantage of habitats suitable for regeneration close to parent stands. The importance of rare climatic events and the role of birds in extending the normal dispersal range of seeds, however, should not be underestimated. The large food store in the seed allows seedlings to reach a size at which they may survive in shaded habitats for several years while waiting for a gap to be created. Few survive in such habitats for longer than three years. The food store may also have a bearing on dispersal by birds. A longer reproductive life may provide A. hunsteinii with more opportunities to produce successful offspring and guard against the likelihood of local extinction, which may occur if the frequency of perturbations is once every several hundreds of years. The likelihood of regeneration success in A. cunninghamii, on the other hand, does not alter markedly from one year to the next. This fact, combined with its higher seed production, means that A. cunninghamii will produce a larger number of potential offspring than A. hunsteinii during its shorter reproductive life. The different regeneration strategies of the two species prescribe the types of communities in which they can occur. A. hunsteinii is most common in late secondary forest.

agathis forest The taxonomy of the genus Agathis is still the subject of much controversy but three species were accepted for New Guinea by de Laubenfels (1988). Only Agathis labillardierei occurs in Papua, where it is widely dispersed. This species also occurs in the Sepik River valley in Papua New Guinea. It grows on a variety of sites, from peat swamp near sea level up to lower montane communities at 1,350 (–1,800) m altitude. Reynders (1964) recorded it from sandy terraces, sometimes poorly drained. Vink (pers. comm., cited in Whitmore 1977: 11) observed that it is commonly associated with blackwater streams. It is widespread above 200 m altitude and is locally very common. It forms pure stands in Papua and the Sepik Valley where it was often tapped for copal resin. Agathis labillardierei is widespread in Papua and forms distinctive forests. At 750 m in the Idenburg Valley, the altitude at which cloud cover appears in the mid-afternoon, a change was observed, with Agathis labillardierei becoming a prominent feature in some forests (Brass 1941). Agathis stands occur on ultramafics on Biak Island from Bosnik (Vink 1960, cited in Whitmore 1997) and on Yapen Island (Dijk 1939, cited in Whitmore 1997). Van Royen (1963) recorded Agathis over limestone. Little information is available on the ecology of Agathis in Papua.

rheophytic communities along fast-flowing streams Characteristic vegetation consisting mainly of shrubs occurs throughout New Guinea on sandy and rocky banks and beds of streams, sites that are subject to sudden brief flooding by fast-running water. The shrubs have horizontal branches

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spreading in the direction of the stream, and narrow, willow-like leaves. They are flood-resistant, tough, and firmly anchored by a wide root system. Common genera are Ficus, Syzygium, and Neonauclea, but many more show the same habit. The type is not restricted to New Guinea, but occurs worldwide. Its members (rheophytes, van Steenis 1952), also include ferns, grasses, and several more or less permanently inundated aquatic herbs. Shea at al. (1998) studied a lower montane community occurring along the upper Minajerwi River dominated by Ficus and Neonauclea, a low to medium rheophytic shrubland. A characteristic rheophyte in southern Papua is Dipteris lobbiana, which is also found in Borneo. It grows along small streams in these communities on the southern slopes of the mountains in heath and lower montane vegetation.

shrubland and herbland communities Disturbed areas along the Freeport Road may be dominated by shrubs, grasses, sedges, forbs, and ferns. These are pioneer communities colonizing disturbed sites. These communities differ depending on the dominant life form and species.

succession on abandoned gardens A main cause of secondary forest in the lower montane area is the practice of shifting cultivation. Shifting cultivation involves the clearing of the primary forest, or older secondary forest, followed by burning and cultivation. Where there was an extensive reserve of suitable primary forest, the abandoned farms are allowed to return to a medium to tall forest which is structurally similar to the primary forest. Where suitable land is limited, farmers revert to a bush-fallow farming system where garden plots are allowed to recover to a seral vegetation cover of pioneer trees which shade out the garden weeds and improve the fertility of the soil. After a relatively short fallow period of a few years, the garden site is once again cleared and used for farming. In parts of the highlands of New Guinea Casuarina oligodon has been used for several hundred years to reduce fallow periods. This species is mycorrhizal and rapidly restores the nitrogen levels following gardening cycles. The timber of Casuarina was also used for making village houses and for firewood and, more recently, as a source of charcoal for cooking. Although unmeasured, the successional sequence from newly abandoned field to primary climax forest is estimated to take several hundreds of years, depending on the availability of nearby seed sources (Gomez-Pompa et al. 1991). Abandoned gardens are quickly overgrown by an herbaceous community of garden weeds and grasses, including Ageratum, Stachytarpheta, Arthraxon ciliaris, and Ischaemum polystachyum. In the absence of fire, the pioneer herbaceous community is invaded by the grasses, ferns, and fast-growing light-demanding shrubs. Miscanthus floribulus, a cane grass with sharp finely serrated leaves, is usually the dominant species. This grass grows either in pure stands or in mixture with Imperata cylindrica. The community is eventually transformed into a mixed species scrubland.

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Midmontane Vegetation

midmontane forest Midmontane forest, occurring between 1,500 and 2,800 m asl in Papua, is characterized by long slopes, spurs, and ridges leading up to the summits of the central mountain ranges. The lower boundary of this subzone at 1,500 m asl is the elevation at which clouds tend to intercept the mountain slopes and the elevation that is most frequently cloud covered. At this elevation, there is a transition to mossy midmontane forest. The upper boundary is less well defined and occurs at approximately 2,800 m asl. Nothofagus, which tends to dominate in the midmontane subzone, does not generally occur above this elevation. The midmontane subzone therefore represents the main range of distribution of Nothofagus. The mainly lowland families Anacardiaceae, Annonaceae, Burseraceae, Dipterocarpaceae, Ebenaceae, Guttiferae, Leguminosae, Meliaceae, Myristicaceae, Sapindaceae, Sapotaceae, and Sterculiaceae are far less frequent and less dominant in the midmontane subzone. They tend to be replaced by members of the families Cunoniaceae, Cupressaceae, Elaeocarpaceae, Fagaceae, Lauraceae, and Podocarpaceae. A wide range of vegetation types occurs within the midmontane zone. The structure and floristic composition of these vegetation types is determined by a number of environmental factors. According to Paijmans (1976), the main factors influencing midmontane forest are climate, altitude, topography, and, to a lesser extent, soil and rock type. The physiognomy of midmontane forest is affected by the frequency and duration of low cloud cover. It is common for forests to take on a midmontane character at lower than normal elevations in areas that are frequently covered by low clouds, and at higher than normal elevations where adjacent lowlands have a monsoonal or seasonal climate. There is some evidence that cloudiness may also affect floristic composition. For example, where low clouds are common, ‘‘mossy forest,’’ characterized by Nothofagus, Phyllocladus, and Astronia, may occur on ridges at 900 m altitude, even though it is more common for these genera to occur at higher altitudes. These are, however, exceptions. In general, the higher the altitude of the forest, the longer it is enveloped in cloud and the more pronounced are characteristic midmontane features, especially a general mossiness. This has led to such forest being known as ‘‘cloud forest’’ and ‘‘mossy forest.’’ The floristic composition of the forest changes with increasing altitude. The total number of species present tends to decrease with altitude, while the number of individuals in certain taxa may increase. Nothofagus, or southern beech, is prominent mainly between 1,500 and 2,600 m. Castanopsis and Lithocarpus, which are dominant in the lower montane subzone, are less common and often rare or absent in the midmontane zone. With increasing altitude in the midmontane zone, conifers, Myrtaceae, and Elaeocarpaceae become increasingly common. This pattern of changing predominance can be clearly seen from the air. Castanopsis shows up as yellowish-brown colors, Nothofagus-dominated forest as gray tones and patches of dead trees, and forests with conifers have increasingly darker tones

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as the abundance of conifers increases with altitude. The forest in the highest part of the zone is also characterized by the families Myrsinaceae and Ericaceae and, especially along the margins, by genera such as Drimys, Carpodetus, Olearia, and Schuurmansia. Woody parasites belonging to the families Loranthaceae and Santalaceae may be common in tree crowns, especially along borders with grasslands. Forests on the gentle slopes of plateaus and mountain saddles at altitudes over 2,000 m may reach a height of 40 m and have girths over 1.5 m. In contrast, the forest canopy on ridge crests may be 20 m or less high, even at altitudes below 1,500 m. Narrow ridge crests have steep slopes, shallow soils, are more exposed to wind, and are shrouded in mist more often and for longer periods than the sides of the ridges. The forests on these crests are commonly closely spaced, thin-stemmed, dwarfed, crooked, and gnarled. They grade into scrub towards the upper limit of the subzone. These forest or scrub formations have been called ‘‘elfin woodland’’ (Troll 1957; Grubb et al. 1963). As in the lower montane forest, forest on steep slopes tend to be irregular and open, with many trees leaning or bent at the base, and climbing bamboo usually abundant. The influence of soil, or the lack of it, is largely correlated with topography. The underlying rock type appears to have only a minor influence, except where ultramafics and limestone are present. On these rock types, the forest tends to be denser and smaller-crowned, and consequently to have a lower average girth than on other rock types. Conifers and Casuarina papuana tend to be more frequent on ultramafics, but no species are known to be restricted to this rock type. With increasing altitude in the Arfak Mountains there was a gradual transition to an intermediate mossy forest. The trunks and bases of smaller trees were covered with hepatics and mosses. Intermediate mossy forest grew on the southwest ridge in the Anggi Lakes area, dominated by Podocarpus rumphii, Dacrycarpus compactus, Phyllocladus hypophyllus, and Papuacedrus papuana var. arfakensis. Many genera of small trees associated with this forest include Drimys, Baeckia, Backhousia, Psychotria, and several myrtaceous plants. Epiphytes and lianas were common throughout the forest. In the basin of the Anggi Lakes the same species predominate, but the forest was taller, to 16 m. The forest was reportedly very rich in epiphytic mosses, lichens, ferns, and orchids. Gibbs (1917) lists the species collected in these forests.

Mixed Midmontane Forest Mixed midmontane forest has a canopy between 20 and 30 m high. It is smaller crowned and more even in height than the dryland evergreen rainforest in the lowland zone. Leaves are generally more dense, leathery, shiny, and dark green, and have entire or serrated margins. The average leaf size is smaller than in lowland forest. Tree density is often very high, but the average girth is smaller than in forest at lower altitudes. Many trunks are low-branched, and bent or leaning, and old trees have thick, crooked, and often dead branches. Stilt roots and adventitious roots are found in places, and heavy buttresses are all but absent. Frequent canopy trees belong to the families Fagaceae, Lauraceae, Cunoniaceae,

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Elaeocarpaceae, and Myrtaceae, the genera Ilex, Dryadodaphne, and Pouteria, and conifers (Cupressaceae and Podocarpaceae). Garcinia, Alstonia, Astronia, Polyosma, Symplocos, Sericolea, Drimys, Prunus, Pittosporum, and Araliaceae are common in the lower tree stories. Notably absent from higher altitudes are members of the families Meliaceae, Burseraceae, Annonaceae, Leguminosae, Dipterocarpaceae, and Ebenaceae—families that are well represented in the lowlands. The shrub layer is generally denser than in lowland forests, but the density varies greatly with its composition and with the height and density of the tree layers. In many places, the shrub layer consists of a great number of slender saplings, while in other places tall ferns and gingers are more common than woody undergrowth. Various species of Rubiaceae, Myrsinaceae, Melastomataceae, Eurya, Cyrtandra, Saurauia, and Piper are nearly always present. The ground flora, mainly consisting of mosses, ferns, herbs, lycopods, and seedlings, is also variable in density. In some places, mosses almost completely cover the ground and the fallen logs and branches, while in other places ferns or Elatostema form a dense layer. The tall-stemmed moss Dawsonia, shaped like a miniature pine tree, is conspicuous, and near forest borders and in glades Sphagnum cushions may be found. Lianas are less common than in lowland forest. Thin woody ones abound in places, but thick woody climbers are seldom present. Climbing rattan and other palms are rare, and are absent above 2,200 m. However, in many places near the forest edge, a thin-stemmed climbing bamboo of the genus Nastus forms dense tangles, and a scrambling Rubus often covers the ground and shrubs. Other climbers include the pandan Freycinetia, Gesneriaceae, Lycopodium, and ferns. Epiphytic and ground mosses become abundant with increasing altitude, as do tree ferns and epiphytic ferns. Orchids are usually quite common in tree crowns, low on the trunks, and on the ground. Stilt-rooted pandans, often very tall and reaching into the canopy, occur both singly and in groups. Some of these are spared during forest clearing and are also planted in gardens, as their oily seeds with carotene-rich outer layers are highly valued as food. The species composition of the midmontane forest varies from site to site. Some of the more common species mixtures recorded by Shea et al. (1998) along the slopes and ridges in south Papua are: Lithocarpus, Nothofagus, Cryptocarya, and Engelhardtia midmontane forest; Nothofagus, Castanopsis, Elaeocarpus, and Pittosporum dominated midmontane forest; Nothofagus, Podocarpus, Dacrydium, and Papuacedrus dominated midmontane forest; and Nothofagus, Phyllocladus, and Dacrycarpus dominated midmontane forest. Several stands were sampled by Shea et al. (1998). A midmontane forest stand located on slope near Tembagapura, south Papua was dominated by Elaeocarpus nubigenus, Saurauia calyptrata, Pittosporum sp., and Cyathea. Other important trees are Podocarpus, Schefflera hellwigiana, Polyosma integrifolia, Timonius, Homalanthus nervosa, and Symplocos cochinchinensis (Table 5.10.5). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 25.3 m2/ha and the total clear bole volume is 149.1 m3/ha. Near Tembagapura the dominant trees are Elaeocarpus nubigenus, Pittosporum

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9.2

Podocarpus sp.

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2.9

Symplocos cochinchinensis

Source: Shea et al. (1998).

149.1

4.1

Homalanthus nervosus

Total

1.6

53.9

Timonius sp.

Elaeocarpus nubigenus

6.0

16.5

Pittosporum sp.

Cyathea sp.

7.3 23.4

7.3

Rhododendron inundatum

Sauauria calyptrata

1.1

Prunus costata

Fagraea ceilanica

1.1

Turpinia pentandra

10.6

0.5

Polyosma ilicifolia

Schefflera hellwigiana

3.4

Polyosma integrifolia

Species

Total clear bole volume (m3/ha)

767

33

33

33

133

133

33

33

33

33

33

33

67

33

33

67

Absolute density

100.0

4.3

4.3

4.3

17.4

17.4

4.3

4.3

4.3

4.3

4.3

4.3

8.7

4.3

4.3

8.7

Relative density (%)

25.3

0.6

0.6

0.6

8.2

2.0

2.4

4.2

1.0

1.0

0.3

0.3

1.3

1.6

0.3

0.9

Absolute dominance (m2/ha)

100.0

2.3

2.3

2.3

32.6

8.0

9.3

16.6

4.1

4.1

1.0

1.0

5.2

6.5

1.0

3.4

Relative dominance (%)

1.92

0.08

0.08

0.08

0.33

0.33

0.08

0.08

0.08

0.08

0.08

0.08

0.17

0.08

0.08

0.17

Absolute frequency

100.0

4.3

4.3

4.3

17.4

17.4

4.3

4.3

4.3

4.3

4.3

4.3

8.7

4.3

4.3

8.7

Relative frequency (%)

300

11.0

11.0

11.0

67.4

42.8

18.0

25.3

12.8

12.8

9.7

9.7

22.6

15.2

9.7

20.8

Importance value

Table 5.10.5. Mixed midmontane forest dominated by Elaeocarpus nubigenus, Saurauia calyptrata, Pittosporum sp., and Cyathea spp.

Montane Vegetation of Papua / 999

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sp., Drimys piperita, and Cyathea (Table 5.10.6); the other common species is Rhododendron inundatum. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 46.2 m2/ha and the total clear bole volume is 410.1 m3/ha. Another stand was located on slope adjacent water tank above Ridge Camp in the Freeport Contract area. The dominant trees there are Prunus dolichobotrys, Podocarpus pilgeri, Pullea, and Homalanthus nervosa (Table 5.10.7). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 26.0 m2/ha and the total clear bole volume is 181.2 m3/ha.

Nothofagus Forest The genus Nothofagus is represented in New Guinea by 13 species (van Steenis 1972b). All are trees but vary from tall canopy trees forming distinctive forests 30–35 m tall, to small shrubs, some less than 1 m tall, where plants occur in more exposed habitats. The shrubs grow in exposed situations on limestone at high altitudes to 3,000 m asl. Hynes (1973, 1974) made the first detailed ecological study of the Nothofagus working mainly in the Highlands of Papua New Guinea. Species of Nothofagus are often dominant or co-dominant in the midmontane forests between 1,500 and 2,800 m asl. They play an important ecological role in these forests. Nothofagus is of scientific importance because of its interesting biogeography. Ash (1982) also made a detailed study of Nothofagus in Papua New Guinea. Ecology of Nothofagus in New Guinea Palynological records in New Guinea (Walker 1970; Bowler et al. 1976; Hope 1976a) indicate that during the glacial period Nothofagus was at about two-thirds of its present altitude but then migrated rapidly to higher elevations as the climate warmed. These extensive Nothofagus forests were then gradually invaded by other genera, which together now constitute the midmontane forest. The rates at which Nothofagus migrated generally exceed the rates of vegetative spread by suckers and therefore require establishment from seed. Nothofagus colonization of other forest types is rarely recorded in the absence of catastrophes, and grasslands previously associated with cold air drainage may have provided routes for migration. The increase in Nothofagus abundance at low altitudes during cooler period was probably the result of the spread of local populations rather than migration from high altitudes (Ash 1982). At altitudes above about 2,000 m, the floristics of forest with Nothofagus are generally similar to adjacent forest without Nothofagus (Saunders 1965, 1970; Kalkman and Vink 1970). At lower altitudes, the ridge top forests with Nothofagus typically include species and genera at the lower limits of their altitude range. Forest containing Nothofagus changes composition in relation to altitude, topography, location, and so on, and there are no species or genera always associated with Nothofagus, though some species are usually present in certain environments. At altitudes below 2,300 m, Nothofagus is often associated with Castanopsis acuminat-

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178.2

Pittosporum sp.

Source: Shea et al. (1998).

Total

410.1

12.6

7.7

Rhododendron inundatum

4.7

Elaeocarpus nubigenus

Drimys piperita

206.9

Species

Cyathea spp.

Total clear bole volume (m3/ha)

799

33

133

133

67

433

Absolute density

100.0

4.2

16.7

16.7

8.3

54.2

Relative density (%)

46.3

1.6

17.9

1.7

0.9

24.2

Absolute dominance (m2/ha)

100.0

3.5

38.6

3.7

1.8

52.3

Relative dominance (%)

1.41

0.08

0.25

0.33

0.17

0.58

Absolute frequency

100.0

5.9

17.6

23.5

11.8

41.2

Relative frequency (%)

300.0

13.6

72.9

43.9

21.9

147.7

Importance value

Table 5.10.6. Mixed midmontane forest dominated by Elaeocarpus nubigenus, Pittosporum sp., Drimys piperita, and Cyathea spp.

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Source: Shea et al. (1998).

Total

Homalanthus nervosus

Prunus dolichobotrys

Podocarpus pilgeri

Pullea sp.

Species

181.15

1.83

116.58

55.41

7.33

Total clear bole volume (m3/ha)

433.32

33.33

333.33

33.33

33.33

Absolute density

100.00

7.69

76.92

7.69

7.69

Relative density (%)

26.04

0.26

16.81

7.92

1.05

Absolute dominance (m2/ha)

100.00

1.01

64.57

30.40

4.02

Relative dominance (%)

0.83

0.08

0.58

0.08

0.08

Absolute frequency

100

10

70

10

10

Relative frequency (%)

300.00

18.70

211.50

48.09

21.71

Importance value

Table 5.10.7. Mixed midmontane forest dominated by Prunus dolichobotrys, Podocarpus pilgeri, Pullea, and Homalanthus nervosus

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Montane Vegetation of Papua / 1003

issima or Lithocarpus, but in sites studied in Papua New Guinea the Nothofagus species always occupied ridges and spurs. At high altitudes, there are associations with Phyllocladus hypophyllus, while Dacrydium and Papuacedrus occur in some wet and cold habitats. It is notable that these latter associates of Nothofagus have similar biogeographic occurrence. Nothofagus forests have often been cleared by local people for agriculture. Undisturbed stands adjacent to forest garden plots or at higher altitudes are used by local people for hunting and gathering timbers, barks, and fruits. Pandanus may be tended in Nothofagus forest, and this may involve both clearing undergrowth and ringbarking of canopy trees. Occasionally, Nothofagus seedlings are planted near villages. Nothofagus timber is hard and durable with a density of 650 kg/m3 and moderate strength. Local people use the wood for house construction and for making tools and household utensils. The timber is also used as a cooking or heating fuel. Timber yields in PNG average 45–60 m3/ha, which is comparable to lowland forests and higher than for most high altitude forests. In Papua New Guinea, villagers on the southern slopes of Mt Giluwe have a complex local taxonomy for Nothofagus. The generic name applied was karapeh, with individual species being called Karapeh karapeh, Karapeh pu, Karapeh yomba, Karapeh tomba, and Karapeh tombona. This closely reflected the traditional uses of the different species of Nothofagus. The effect of logging on Nothofagus forest depends upon several factors, including the number and proportions of different species remaining after logging, the disturbance to the forest floor, the availability of seed and seedlings (suckers), and subsequent activities such as cultivation and burning. If disturbance is not too great secondary forest and logged forests sometimes contain Nothofagus. However, where impacts are too destructive, grasslands and secondary forest may develop. The present trend is for Nothofagus forest to be reduced in area, through conversion to other land uses. The various species of Nothofagus have different altitudinal ranges. N. flaviramea, N. starkenborghii, N. rubra, N. carri, and N. crenata are all recorded below 900 m and their upper altitudes are above 2,200 m. N. brassii, N. perryi, and N. grandis are abundant over the range 1,500–2,500 m, while N. pullei extends its range to above 2,800 m. In any one locality, only a few of the species are present and it is unusual to find more than two or three species of Nothofagus forming a mixed forest. Fluctuations in climate such as those that have occurred during the Pleistocene have shifted the vegetation zones altitudinally and thereby created or removed barriers to horizontal migration. This is reflected in the present distribution of the various species. Nothofagus is generally associated with continually high precipitation and cloudiness. Clouds reduce light penetration by as much as 70%, though average values in highland New Guinea range between 30% in the morning and 50% in the afternoon. Under very cloudy and wet conditions, decreased light may be limiting tree growth (Ash 1982). The interception of radiation by clouds reduces both the temperature and the daily range of temperatures. To grow competitively,

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Nothofagus requires at least a short growing season each year with mean daily temperatures in excess of 10C. Some of the tropical Nothofagus of New Guinea will grow competitively in localities with mean daily temperatures as high as 23C. This is much higher than for any temperate species of the genus. In New Guinea, Nothofagus reaches a maximum altitude of 3,100 m, which is well below the tree line (above 4,170 m on Mt Jaya). Nothofagus does, however, occur within the range of frequent frosts (as low as 1,500 m on cloudless nights). Brown and Powell (1974) recorded severe frost damage to Nothofagus and other midmontane species following prolonged dry and cloudless periods in 1972. For successful competitive growth, Nothofagus requires a fairly continual supply of water derived from reasonably aerated soils. The annual precipitation required to meet this condition in New Guinea is 1,500 mm/yr in an exposed, ridge top community. Rainfall in New Guinea is therefore generally in excess of the requirements for Nothofagus, but water deficiencies may occur during drought periods and where there is a rain shadow effect. Nothofagus is generally absent from localities with regular and sustained water deficits. The coriaceous leaves of Nothofagus lose water relatively slowly compared to other midmontane species (Hynes 1973) which indicates a resistance to water deficit. New Guinea is generally very cloudy and certain topographic features such as ridges and passes regularly experience ground level cloud from which the vegetation filters water droplets to provide very high local precipitation combined with low temperatures and low light penetration. The ‘‘cloud forests’’ (also known as ‘‘moss forests’’ because of the abundant epiphytic bryophytes) often contain Nothofagus, but the genus is not restricted to these conditions. Nothofagus pollen and seeds are dispersed by wind. Wind speeds in New Guinea are generally rather low, but gusts occasionally cause local damage especially to trees on exposed slopes, ridges, and forest edges. The burning of vegetation depends upon several factors, including the morphology and chemistry of the various species, low humidity, and a source of ignition. Nothofagus forests are not very flammable. The burning of Nothofagus forests is associated with periods of prolonged drought (El Nin˜o events) and human activities. Many fires are ignited by humans and the frequency of forest fires has increased during the later Quaternary as the result of human activities. In New Guinea, disturbed Nothofagus forest bordering cultivated areas is frequently subjected to burning, but fires rarely penetrate undisturbed forest. Nothofagus regenerate poorly after fires, especially if the regrowth is also burned. Repeated burning may convert Nothofagus forests into grasslands. Exceptional burning of extensive areas of undisturbed Nothofagus forest was recorded in the Star Mountains and near Telefomin region following droughts in 1972. New Guinea experiences considerable tectonic activity. Minor earthquakes may be sufficient to damage tree roots and thereby encourage diseases. Major earthquakes may cause trees to fall, and may also trigger landslides which both destroy vegetation and provide bare surfaces for recolonization. Volcanic activity may damage vegetation through burning, toxic gases, and burial under ash or lava. It

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Montane Vegetation of Papua / 1005

may also provide a nutrient-rich substrate, and light falls of volcanic ash may improve soil fertility over large areas. Nothofagus forests may be subjected to all these catastrophic events and its response depends to a considerable extent upon whether regeneration from existing individuals is possible or whether the site must be colonized with seeds. In New Guinea, Nothofagus is rarely recorded from sites of recent landslides or volcanic activity. There are other genera with better seed dispersal and faster growth rates than Nothofagus, and these soon form a dense community on bare ground. Nothofagus forests in New Guinea grow on a diverse range of substrates, from ultramafic to acidic igneous rock, calcareous to siliceous sediments, and various metamorphics. Nothofagus growth is probably most rapid on volcanic substrates, and slowest on shallow soils over limestone where tree may be stunted. Rock type correlates poorly with Nothofagus abundance, which may result partly from the considerable amount of volcanic ash that has fallen throughout New Guinea. Some variation in Nothofagus abundance with substrate has however been recorded; for example, Nothofagus is locally absent from the sandstone plateau near Mt Jaya (Hope 1976). Some Nothofagus have been recorded growing on extreme sites. N. starkenborghii has been found growing on poor inorganic lowland soils, N. perryi has been found on shallow soils over limestone, and N. rubra has been found growing on wet peaty soils. Much of the local variation in the abundance of Nothofagus species is related to topography (Paijmans 1976; Ash 1982). At low altitudes in New Guinea Nothofagus is usually restricted to ridge crests, but at high altitudes various species occur on most types of topography. Many factors change along the topo-sequence catena away from a ridge crest. Microclimatic changes include cloudiness, humidity, solar radiation, and wind, while surface processes are dominated by the downslope movement of water, solutes, and particules. Slopes are subjected to soil movements which may cause trees to lean and fall while stable ridge crests may support much older trees. Animals (including humans) may preferentially travel, feed, disperse seeds, and ignite fires along ridges. Certain pathogens, especially soil fungi, may be associated with the wetter conditions on the lower slopes. Thus, with so many complex and interacting variables operating upon many tree genera, it is not possible to offer a simple explanation of the topographic distribution of Nothofagus (Ash 1982). Diverse fauna and variety of fungi are found in the midmontane forests of New Guinea, but little evidence that Nothofagus is particularly affected by these organisms. Several features indicate a resistance to herbivore activity, including the hard cupules, coriaceous leaves, hard and durable wood, and the presence of polyphenolic compounds, tannins, and triterpenes (Hegnauer 1966; Hillis and Inoue 1967; Gibbs 1974). Minor damage from insects and fungi has, however, been observed for cupules, seeds, leaves, and wood. Leaf litter in Nothofagus forest decays within a few years (Hynes 1973), and large logs decay within decades. This indicates an active saprophytic flora of fungi, bacteria, and algae. Dieback and the deaths of stands of Nothofagus have been recorded from many

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localities in New Guinea. Dieback is usually associated with a combination of weather, substrate, pathogens, and hosts at a suitable stage of development. Pathogenic soil fungi, including Phytophthora cinnamomi and P. cambivora, and wood boring beetles are associated with dieback of Nothofagus pullei on Mt Giluwe, but the causal relationship has not been fully established. Nothofagus pullei stands in south Papua (in the vicinity of Hidden Valley) have experienced some dieback of this type, but the cause has not been established. It is possible that lightning strikes may be a factor in the dead patches in Nothofagus. Nothofagus has ectotrophic mycorrhizae, and there is generally a substantial increase in growth rates associated with mycorrhizal fungi of this type. The successful colonization of new sites from seed may be limited in the absence of mycorrhizae. The humid environment of Nothofagus forest is favorable for epiphytes, and many trees support sooty molds, epiphyllous bryophytes, bacteria, and algae on their leaves, and bryophytes and vascular epiphytes on their branches and trunks. Different species have different strategies for dealing with these epiphytes; for example, some species have bark that flakes off along with any attached epiphytes. There is a tendency for seasonal flowering, but flowering may occur sporadically throughout the year. Fruiting and flowering occurs at an early age on trees growing in the open, but occur later on trees grown under a canopy. Nothofagus produces abundant wind dispersed binucleate pollen that is morphologically variable, but the species are not distinctive. In any locality, the greater part of the pollen will be of local origin but occasionally fertilization will occur with pollen from distant sources. Seed production varies from year to year, but has been recorded to be as high as 3 million per hectare for Nothofagus pullei (Dawson 1966). The number was much lower for N. grandis and N. rubra. It is postulated that unpredictable fluctuations in seed production may allow more seeds and seedlings to escape destruction by herbivores. Experimental germination of seeds of N. grandis, N. rubra, and N. pullei with moderate shading yielded less than 1% success. The frequency of seedlings beneath stands of large Nothofagus trees was recorded to be 50–1,000 seedlings per ha in PNG, which suggests that few seeds develop into established seedlings. The majority of seedlings grow upon mossy substrates, especially fallen logs. The advantage of the mossy log substrate probably relates to the reduction of competition with other vascular plants, but mycorrhizae, pathogens, and herbivores may also be involved (Ash 1982). The canopy of Nothofagus stands in New Guinea is generally rather open, with 5–8% daylight penetration. Under these conditions, seedlings will make rather slow growth and few survive to become saplings (Ash 1982). The gregarious nature of many Nothofagus populations and the presence of regeneration indicate that allelopathy is not very significant. Because of difficulties associated with seeds and seedlings, much of the regeneration of Nothofagus in New Guinea arises from vegetative reproduction (suckers). The suckers originate from several sources including the undersurface of fallen trunks and damaged or suppressed seedlings and saplings. Horizontal suckers may spread along the soil surface for several meters, producing several vertical shoot

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Montane Vegetation of Papua / 1007

and root systems. The rate of growth of seedlings or suckers is greatly increased by senescence of the tree canopy or the creation of canopy gaps. Compared to other genera, Nothofagus grows slowly under shaded subcanopy conditions but rapidly under more open conditions. Nothofagus Forest Types Nothofagus forests may be conveniently classified into those that are nearly pure Nothofagus stands versus those with a mixture of other genera, and also into those with a restricted range of size classes versus those with a complete range of sizes including regeneration. Nearly pure Nothofagus pullei forest with a mosaic of patches each dominated by a single size class occur on Mt Giluwe in Papua New Guinea. The forest is evidently characterized by a cycle of stand mortality followed by regeneration to form canopy trees, which may in turn suffer extensive mortality. There are many genera of competitors in the regrowth, but Nothofagus apparently grows faster and to a greater height, and thereby dominates the forest. Similar Nothofagus dieback has been recorded from many regions of New Guinea, including the area around Hidden Valley. Small populations of Nothofagus on ridges may have a single abundant size class of large canopy trees, but Nothofagus regeneration is often observed where a tree is senescent. The local patches of regeneration are probably sufficient to maintain these populations. Extensive stands of Nothofagus may have all size classes represented, but at a small scale the forest resembles that on ridges with local patches of regeneration. Forests in which Nothofagus forms only a minor portion of the canopy and is even less abundant in smaller size classes have also been recorded. The Nothofagus trees may be very large and senescent but the regeneration is mostly from other genera. This forest type is apparently successional from Nothofagus to a multigenera forest. In some localities, Nothofagus regeneration is recorded in multigenera forest, but it is difficult to estimate the stability of the populations from short periods of observation of the individuals. Some species of Nothofagus form canopy trees with a height of 45 m and trunk diameters of 80–180 cm. The lower trunk is often buttressed and the roots extend a few meters from the tree and to a depth of 20–40 cm. The bole is often half the height of the tree and gives rise to several branches which spread at angles of 10–30 from vertical to give a nearly level or domed canopy. Altitude has little effect on the maximum size of Nothofagus trees in New Guinea. Stunted trees are associated with poor growing conditions, such as shallow soils over limestone, waterlogged soils, or exposed cloudy reaches. Canopy leaves tend to be 4 to 10 cm in length. Leaves may be smaller in Nothofagus pullei, N. carrii, and, on some sites, N. rubra and N. pseudoresinosa. Information from tree growth rates, radiocarbon dating, and regular parenchyma rings suggests that Nothofagus trees in New Guinea reach a maximum age of 350–550 years. Brass (1941) recorded the changes of floristic composition of an altitudinal series of forests dominated by Nothofagus in the Idenburg area. At 900 m, on the ridges ‘‘a mossy forest’’ dominated by N. grandis with Dacrycarpus imbricatus,

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and Phyllocladus hypophyllus was associated with Metrosideros, Rhodamnia, and Endriandra. Agathis and Gulubia (Palmae) were scattered emergents. The subcanopy of this forest included abundant Pandanus leptocaulis, Dianella, and Mapania in addition to Cyathea biformis, Nepenthes, Freycinetia, and the bamboo Schizostachyum. At 1,800 m the forest is dominated by Nothofagus rubra. It forms a closed forest about 25 m tall on the major ridges, descending to 1,550 m on the south facing (colder) spurs. Other canopy trees included Phyllocladus hypophyllus, Calophyllum congestifolium, and Weinmannia. The subcanopy of the forest has several Cunoniaceae which are prominent, and in addition but less common, Archboldiodendron, Metrosideros, and Fagraea. Freycinetia, Pothos, Tecomanthe, and the climbing Cyathea biformis were common on the tree trunks. The thin undergrowth consisted of the small tree ferns Cyathea perpelvigera and Cyathea melanoclada, and species of Medinilla, Argostemma, Burmannia, Lindsaea, Grammitis, and the saprophyte Corsia. Woody epiphytes were common in the middle and upper levels of the canopy. These included species of Melastomataceae, Vaccinium, Rhododendron, Drimys, Rapanea, Sericolea, the ant plants Hydnophytum and Myrmecodia, Psychotria, Alyxia, and Nepenthes, which grows as a scrambling plant in canopy. Epiphytic ferns were highly diverse and abundant, as were the many species of Zingiberaceae. Small openings caused by landslides and windthrows (or clearings) support a variety of plants. Riedelia, Dianella, Eurya, Medinilla, Lycopodium cernuum, Sticherus, Rapanea, Drimys, and Rhodomyrtus often regenerate freely in these openings. In larger gaps Breynia, Homalanthus, and Pandanus actinobotrys are often found. At 2,300 m on an exposed, ridge top community, Nothofagus rubra forms a small (2 to 3 m tall) tree, associated with Xanthomyrtus and Mearnsia. It also contains abundant secondary species of Rapanea, Drimys, Sericolea, Ternstroemia, and Kania. Papuacedrus formed small stands to 15 m tall, on edges of rockslides and in forest on very steep rocky ground. In the Bele River (Brass 1941) Nothofagus dominated from 2,200 to 3,100 m altitude. Three distinct species occur; Nothofagus starkenborghii dominated below 2,400 m and above that altitude Nothofagus brassii was dominant. At 2,300 m another species, Nothofagus crenata, was plentiful. Lauraceae, Elaeocarpaceae, Syzygium, Cunoniaceae, and Podocarpus were the other dominant trees. The forests are characterized by a mossy ground cover and differ markedly from the Nothofagus forests southwest of Bernhard Camp. Around 2,800 m the forest was dominated by two species of Nothofagus: N. brassii at slightly lower and N. rubra at upper altitudes. The dominants formed majestic trees to 35–40 m tall and up to 1.5 m diameter above their spurred bases. Rainforest species, where present, were restricted to moist bottom and lower slopes of valley systems. This forest has a rich associated flora on moist flats, consisting of Pilea, Piper, Cyrtandra, Marattia, Cyathea, and Dicksonia. Small clearings were dominated by Saurauia, Homalanthus, and Melicope.

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Montane Vegetation of Papua / 1009

Discrete patches of Nothofagus forest within mixed midmontane forest have been recorded in New Guinea (Robbins and Pullen 1965; Robbins 1970; Kalkman and Vink 1970; Walker 1966). The boundary of these patches is usually sharp, and Nothofagus regeneration scarcely extends beyond the canopy of the Nothofagus trees (Walker 1966; Kalkman and Vink 1970). It has been suggested that these patches are relics of more widespread Nothofagus forest that has been encroached upon by other genera. The Nothofagus stands are therefore in habitats that favor Nothofagus, such as ridges, or in localities where other genera have not become established for a variety of reasons, including their rates of migration. Excavations of ‘‘seedlings’’ in Nothofagus forest on Mt Kaindi in Papua New Guinea showed that the regeneration of the Nothofagus consisted entirely of root suckers (Johns, pers. obs.). Tall Nothofagus forest and closed-forest stands occurs on ridge crests and upper slopes, mainly between 1,500 and 2,800 m in Papua. These stands may reach 45 m in height, and rival lowland forests in terms of basal area and clear bole volume. Shea et al. (1998) sampled several stands of Nothofagus forest in south Papua. Stands can be dominated by Nothofagus pullei, Symplocos, and Pandanus, or by Nothofagus pullei, Pandanus, and Psychotria. On some sites, Nothofagus occurs in tall mixed stands with species of southern conifers. One such stand sampled by Shea et al. (1998) was dominated by Nothofagus, Podocarpus, Elaeocarpus, and Papuacedrus. Two stands of midmontane forest dominated by Nothofagus were sampled by Shea et al. (1998). The stand located adjacent to the housing estate at Hidden Valley above Tembagapura, at about 2,300 m asl, was dominated by Nothofagus pullei, Symplocos cochinchinensis, and Pandanus (Table 5.10.8). The estimated total basal area for trees with a diameter greater than 10 cm dbh is 45.1 m2/ha and the total clear bole volume is 331.6 m3/ha. The second stand was located adjacent to Tembagapura above Hidden Valley and Mile 66. The dominant trees were Nothofagus pullei, Pandanus, and Psychotria (Table 5.10.9). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 29.3 m2/ha and the total clear bole volume is 162.6 m3/ha.

midmontane swamp vegetation Montane swamp forest grows in small patches and bands fringing swampy intermontane basins occupied by grass or sedge swamp (Paijmans 1976). The forest usually has a low open canopy over a dense layer of small trees and shrubs, and a sparse herbaceous ground cover. Most trees grow on hummocks separated by deep pools of water. Common trees include Syzygium and other Myrtaceae, Garcinia, conifers, and locally Nothofagus sp. Some stands are dominated entirely by conifers, particularly the genus Dacrydium and Podocarpus. Such forests are not common but occur in isolated locations. Two types of midmontane swamp forest are recorded: Syzygium and Garcinia or those swamp forests dominated podocarps. In some areas swamp forests can also be dominated by Araucaria.

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Nothofagus pullei

331.7

Total

Source: Shea et al. (1998).

104.8

Symplocos cochinchinensis

7.3

219.6

Species

Pandanus sp.

Total clear bole volume (m3/ha)

867

300

100

467

Absolute density

100.0

34.6

11.5

53.8

Relative density (%)

45.1

27.5

2.8

44.8

Absolute dominance (m2/ha)

100.0

36.6

3.7

59.6

Relative dominance (%)

1.25

0.58

0.25

0.42

Absolute frequency

100.0

46.7

20.0

33.3

Relative frequency (%)

Table 5.10.8. Midmontane forest dominated by Nothofagus pullei, Symplocos cochinchinensis, and Pandanus

300.0

117.9

35.3

146.8

Importance value

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Source: Shea et al. (1998).

162.6

3.9

Total

9.1

Nothofagus pullei

Psychotria sp.

149.6

Species

Pandanus

Total clear bole volume (m3/ha)

200

30

40

130

Absolute density

100.0

15.0

20.0

65.0

Relative density (%)

29.3

0.8

6.5

22.0

Absolute dominance (m2/ha)

100.0

2.9

22.2

75.0

Relative dominance (%)

Table 5.10.9. Midmontane forest dominated by Nothofagus pullei, Pandanus, and Psychotria

0.45

0.05

0.10

0.30

Absolute frequency

100.0

11.1

22.2

66.7

Relative frequency (%)

300.0

29.0

64.4

206.6

Importance value

Montane Vegetation of Papua / 1011

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Dacrydium Swamp Forest Small patches of swamp forest dominated by Dacrydium cornwalliana are scattered throughout Papua forming pure stands of swamp forest from 1,340–2,300 m asl. The most extensive stand is on black peat near the Wissel Lakes. The forest was monospecific, the soil a knee-deep black mud. A small plot of 300 m2 contained 16 trees with diameters from 25 to 49 cm (mean: 35 cm) and top height from 28 to 34 m (mean: 30 m). Rejuvenation was only found in open spots caused by a fallen tree. In these spots the secondary vegetation of 1–3 m height included Euodia, Pittosporum pullifolium, Palmeria arfakiana, Schefflera, and Vaccinium, and a single 6 m high Pandanus. Climbers were an Asclepiadaceae and Geitonoplesium cymsoum. In the herb layer Juncus prismatocarpus, Scirpus inundatus, Zingiberaceae, Orchidaceae, ferns, lycopods, and Sphagnum were found. In the forest margin Dacrycarpus imbricatus occurred with only a few trees; this species must have been much more common, but its bark is used for walls in house building (Rappard and van Royen 1959). Dacrydium cornwalliana is rapidly disappearing because of clearcutting of these monospecific stands of good quality timber. What is left is a swampy low, open scrub with Melicope denhamii, Glochidion wisselense, Sloanea arfakensis, Rapanea communis, Medinilla fasciculifera, Vaccinium pullei, V. turfosum, Rhododendron macgregorii, Podocarpus rubens; rejuvenation of Dacrydium cornwalliana and Papuacedrus papuanus; herbs such as Polygonum strigosum, Oenanthe javanica, Gonostegia hirta; climbers and scramblers such as Rubus diclinis, Freycinetia pleurantha, Geitonoplesium cymosum, Nepenthex maxima; ferns such as Nephrolepis rosenstockii and Aglaomorpha novoguineensis; and a soil cover of Spagnum and other mosses, lichens, lycopods, and grasses (Zieck et al. 1960). Collections were made at these sites by Versteegh and Vink (photo in Flora Malesiana 10: p 366). Two plots were established in a stand of D. cornwalliana (Dacrydium nidulum var. araucarioides) in the Southern Highlands of Papua New Guinea (Johns 1980). The only associated tree species recorded was Nothofagus pullei, which was represented by a few scattered individuals. Several epiphytic ferns and the orchid Coelogyne fragrans were common. No data have been recorded from this forest type in Papua.

Midmontane Sedge-Grass Swamp Communities dominated by sedges and grasses commonly occur above about 1,800 m in swamps (Paijmans 1976). They occupy intermontane basins, local depressions in valley floors, and seepage slopes, where either standing or slowly moving water is permanently at or just above the surface. The sedges and grasses are usually low, but in places are over one meter high. They are interspersed with various swamp herbs, and scattered dwarf shrubs are locally present on little hummocks. Many different sedges are present, especially in stagnant swamps, and they commonly make up most of the ground cover. One of the most common sedges is Machaerina rubiginosa, which occurs mainly between 1,800 and 3,000 m, often in pure stands. Characteristic grasses are Arundinella furva and species of Isachne

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Montane Vegetation of Papua / 1013

and Dimeria, mainly between 1,800 and 3,000 m, and Agrostis reinwardtii from about 1,800 to well over 3,000 m.

midmontane grasslands Midmontane Phragmites Grasslands Phragmites karka may form pure stands in seepage areas on slopes and on valley floors. It is also associated with Miscanthus floridulus along river banks and swamp margins, and in very shallow swamps. Both grasses usually form large hummocks, rising well above water level.

Midmontane Grasslands and Herblands Small patches of midmontane grassland occur in intermontane valleys. Below 2,500 m, these grasslands are dominated by species such as Eulalia leptostachys, Ischaemum spp., Arthraxon ciliaris, and Imperata cylindrica. Grasslands above 2,500 m are usually lower in height. Common species are Danthonia archboldii, Deschampsia klossii, Agrostis reinwardtii, Dichelachne novoguineensis, Deyeuxia spp., Anthoxanthum angustum, Arundinella furva, and other species of Danthonia. Sedges (e.g., Gahnia and Machaerina rubiginosa) and herbs (e.g., Potentilla, Ranunculus, Gentiana, Anaphalis) may also be common. These grasslands are somewhat similar to grasslands that occur in the subalpine zone, and may represent the opportunistic colonization of a suitable site by species from higher altitudes. The community classification will depend on which species combinations are present on any particular site. Communities in midmontane zone of Papua can be dominated by a single grass species, such as Danthonia archboldii and Deschampsia klossii grassland. Others can be mixed-species grasslands, sedgeland communities, or herbland communities.

midmontane shrublands and scrub Exposed sites adjacent to grasslands above 1,500 m may be dominated by small trees and shrubs. Many of the shrubs also occur at higher elevations, including species of Rhododendron, Vaccinium, Styphelia, and Hypericum. The community classification depends on which species are actually dominant on the site.

regenerating vegetation Repeated cutting and burning, plus cultivation, causes site deterioration, including soil erosion. The secondary succession may be ‘‘deflected’’ to a fire climax or biotic climax. Instead of trees, stands of shrubs develop, including Melastoma malabathricum, Lantana camara, Rhodamnia cinerea, Rhodomyrtus tomentosa, and locally Dillenia suffruticosa. Ultimately, disturbance leads to the development of open grassland of Imperata cylindrica (locally known as alang alang), which has diffuse rhizomes and can thereby survive fires. Patches of this community type can be seen scattered throughout New Guinea. On the western slopes of Mt Giluwe the

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soils under these grasslands have a dense communities of epiphytic plants from the neighboring forest (R. Johns, pers. obs.).

primary succession on exposed rock and mineral soil Landslips in the montane zone tend to result in the removal of the soil and underlying till, exposing bare rock and unweathered till. Succession on the unweathered till tends to be more rapid than on the bare rock. The till has a rougher surface and the weathering process is more rapid. The vegetation will tend to undergo structural changes over time from closed-mossland, to short open-herbland, short herbland, low to medium shrubland, medium scrub, low open forest, to closed forest. Detailed studies were not carried out for these primary successions, and observations were limited. A number of interesting communities were observed by Shea et al. (1998). On exposed rock and mineral soil in the valley near Tembagapura (south Papua) the pioneer communities varied from crustose lichens to fructose lichen (Cladonia), foliose lichen (Stereocaulon), moss dominated communities, and liverwort (Marchantia) communities.

young seral forest Pioneer tree species become established within the scrub and develop into a woodland or forest. Some of these trees grow from seeds, while other sprout from tree roots that were left in the ground during land clearing and site preparation. The stand is young and even-aged, the canopy more or less of an even height. Early pioneer species tend to dominate heavily disturbed sites, while less disturbed sites are dominated by trees which are more characteristic of secondary forest and disturbed sites within the primary forest. Shea et al. (1998) established two plots in forests dominated by early pioneer tree species (above the PTFI road at Mile 67, south Papua). The dominant trees were Vitex pinnata, Homalanthus nervosus, Cyathea spp., and Mallotus trinervius. Other trees on the site were Ficus, Saurauia capitulata, and Macaranga rhizinoides (Table 5.10.10). The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 8.4 m2/ha and the total clear bole volume is 33.0 m3/ha. The second stand was dominated by Homalanthus nervosus, Timonius nitens, Symplocos cochinchinensis, and Cyathea (Table 5.10.11). The plant community is represented by the name of the four most dominant genera Homalanthus-Timonius-Symplocos-Cyathea. The most important species and genera, based on combined Importance Value (see Appendix 5.6.1) are Homalanthus nervosus, Timonius nitens, Symplocos, and Cyathea. The estimated total basal area for trees with a diameter greater than 10 cm dbh (or diameter above buttresses) is 36.6 m2/ha and the total clear bole volume is 265.7 m3/ha.

transition from midmontane to upper montane forest Upper montane forest usually has an abrupt lower boundary with midmontane forest. The reasons for this abrupt transition are as follows. Upper montane forest is frequently associated with special sites. If site conditions lead to peat formation,

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16.9 0.7 6.4 0.9 0.9 5.1 2.1 33.0

Species

Vitex pinnata

Ficus sp.

Homalanthus nervosus

Saurauria capitulata

Macaranga rhizinoides

Cyathea spp.

Mallotus trinervius

Total

Source: Shea et al. (1998).

Total clear bole volume (m3/ha)

566

33

133

33

33

167

67

100

Absolute density

100.0

5.9

23.5

5.9

5.9

29.4

11.8

17.6

Relative density (%)

8.5

0.6

1.4

0.3

0.3

2.3

0.5

3.1

Absolute dominance (m2/ha)

100.0

7.0

16.3

3.1

3.1

27.1

6.2

37.2

Relative dominance (%)

0.99

0.08

0.25

0.08

0.08

0.25

0.08

0.17

Absolute frequency

100.0

8.3

25

8.3

8.3

25

8.4

16.7

Relative frequency (%)

300.0

21.2

64.8

17.3

17.3

81.5

26.4

71.5

Importance value

Table 5.10.10. Young secondary forest dominated by Vitex pinnata, Homalanthus nervosus, Cyathea, and Mallotus trinervius Montane Vegetation of Papua / 1015

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Source: Shea et al. (1998).

265.6

7.1

Total

4.5

Symplocos cochinchinensis

230.8

23.2

Cyathea sp.

Homalanthus nervosus

Timonius nitens

Species

Total clear bole volume (m3/ha)

600

67

33

400

100

Absolute density

100.0

11.1

5.6

66.7

16.7

Relative density (%)

36.6

2.6

0.6

28.6

4.8

Absolute dominance (m2/ha)

100.0

7.1

1.6

78.0

13.2

Relative dominance (%)

1.00

0.08

0.08

0.58

0.25

Absolute frequency

100

8.3

8.3

58.3

25

Relative frequency (%)

300.0

26.6

15.5

203.0

54.9

Importance value

Table 5.10.11. Secondary forest dominated by Homalanthus nervosus, Timonius nitens, Symplocos cochinchinensis, and Cyathea

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Montane Vegetation of Papua / 1017

then certain upper montane tree species are favored (including members of the Coniferae and Myrtaceae) which are believed to facilitate peat formation by having slowly decomposing litter. If this does occur, the process of peat accumulation becomes self-reinforcing, tending to sharpen even further the boundary with the adjacent forest (Whitmore 1984). One factor that is likely to initiate peat development is waterlogged soil conditions, arising from frequent cloud (Whitmore 1984). The cloud cap tends to form at a particular level on any given mountain, so marked waterlogging will start rather abruptly at this elevation. Again, once peat has begun to form, the self-reinforcing process commences and a sharp boundary forms. Sphagnum frequently grows in these waterlogged places and also accentuates peat accumulation. Thus, there are several different reasons why the lower boundary of upper montane forest may be abrupt, and without examination it is not possible to say which factors are acting in any particular site (Whitmore 1984).

upper montane vegetation Whitmore (1984) described the upper montane forest in Malesia as varying in structure, physiognomy, and flora. The most dramatic change, partly so because it usually occurs sharply over a short distance, is from mesophyll-dominated forest with an uneven, billowing, canopy surface to microphyll-dominated forests with a lower, flattish canopy surface, the trees more slender, usually with gnarled limbs and very dense subcrowns. The upper montane rainforest is encountered first on knolls and narrow ridge crests with mesophyll forest occupying the valleys, saddles, and broader crests. Upwards it comes to clothe the entire landscape. It is as clearly distinctive on aerial photographs or from an airplane as to the traveler on foot. Upper montane rainforest is frequently only 10 m tall or less (range: 1.5–18 m), and its shorter facies are sometimes called elfin woodland. On outlying spurs and on isolated peaks upper montane forest occurs at lower elevations than on big mountain massifs. Upper montane forest reaches its highest altitudes on high mountains with long slopes. Knolls and sharp ridge crests commonly carry forest of lower stature. Knolls and mountain summits are oligotrophic sites that also are more prone to periodic water deficit than hillside, saddle, and valley sites. In New Guinea the upper montane forest generally lies between 2,800 and 3,200 m asl. The boundaries are not fixed and vary according to topography. For example, the lower boundary generally corresponds to the upper limit of distribution of Nothofagus but the boundary may be higher on long slopes, or may be lower on ridges and the crests of plateaus. On the Kemabu Plateau across the Zengillorong Valley and down the Ilorong Valley in south Papua, Nothofagus pullei forest (the beech forest) passes abruptly into lower subalpine forest at about 3,200 m asl. The beech trees at the transition were quite large (20 m) and relatively straight boled, but do not form a closed canopy everywhere (Hope 1976). Hope postulated that in this area, an upper montane forest may be more or less absent, with midmontane forest adjacent to subalpine forest. On the ridge crest at Tembagapura (south Papua), floristic studies suggest that

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1018 / r. j. johns , g. a . shea, w. v i n k, & p . puradyatmika

an open forest appears to occur at a lower elevation (2,600 m asl). Many components of this forest are similar to species collected at 3,200 m in the Star Mountains. The forest is low to medium in height, relatively open, and dominated by Papuacedrus papuana and Dacrycarpus cinctus. The open savanna has a distinctly subalpine savanna element but perhaps could be classified as upper montane or even subalpine. Whitmore (1984) noted that in New Guinea the upper montane flora is much poorer in species diversity than the midmontane or lower montane floras. Elaeocarpaceae are abundant and Lauraceae are distinctly rare, while ground-dwelling Ericaceae are fairly common. The canopy only reaches 12–18 m tall. Ground ferns are less common. Common genera include Zygogynum, Daphniphyllum, Dacrycarpus, Drimys, Elaeocarpus, Eurya, Papuacedrus, Macaranga, Pittosporum, Podocarpus, Quintinia, Rapanea, Saurauia, and Symplocos. Many of these genera also occur at higher elevations in the subalpine zone. Even when the same genera occur the upper montane forests are floristically less diverse than the lower montane forests. Several vegetation types are associated with disturbance, both natural and made by humans. The mixed forests were dominated by Papuacedrus, Saurauia, Symplocos, and Dacrycarpus.

upper montane coniferous forest In many places above 2,400 m conifers of the genera Podocarpus, Dacrycarpus, Papuacedrus, and Phyllocladus dominate the canopy and emergent tree layers (Johns 1982; Mangen 1993). Although generally smaller crowned than their broadleaved associates, many conifers reach a girth of well over 1.5 m, even at altitudes above 3,000 m (Paijmans 1976). Emergent trees of Papuacedrus papuana are easily recognized from a distance by their rather open crown with horizontal branches, hung with gray streaks of the lichen Usnea. Papuacedrus papuana has a wide ecological tolerance and is able to regenerate under dense forest, in the open, and on steep stony slopes. In New Guinea many larger trees are killed by bark stripping because the bark is especially favored for roofing by villagers. Belts of almost pure coniferous forest locally form transition zones on mountainside slopes between midmontane forest below and subalpine forest and grassland above. They also occur on the upper slopes of large dolines in limestone country. In such situations, the hardy conifers may be pioneering where broad-leaved forest is absent because of either fire or frost. In addition to being components of mixed species stands at high elevations, the conifers may form pure stands on some exposed sites. Shea et al. (1998) described two upper montane forest types on the ridge crest south of Tembagapura, south Papua. The first is a coniferous forest of mixed species composition located 1 km south of the military post on the crest before descending into Tembagapura. The dominant trees are Papuacedrus papuana, Saurauia trugal, Symplocos cochinchinensis, and Dacrycarpus imbricatus. Other trees are Cyathea spp., Podocarpus neriifolius, and Homalanthus nervosa (Table 5.10.12). Upper montane forest dominated by the gymnosperms Papuacedrus papuana and Dacrycarpus cinctus is adjacent (Table 5.10.13).

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PAGE 1018

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Source: Shea et al. (1998).

73.54

1.24

Homalanthus nervosus

Total

1.28 40.66

Papuacedrus papuana

2.29

Cyathea sp.

Podocarpus neriifolius

7.28

16.21

4.58

Symplocos cochinchinensis

Saurauia trugal

Dacrycarpus imbricatus

Species

Total clear bole volume (m3/ha)

400.00

33.33

66.67

33.33

33.33

66.67

133.33

33.33

Absolute density

100.00

8.33

16.67

8.33

8.33

16.67

33.33

8.33

Relative density (%)

21.00

0.00

9.75

0.26

1.64

3.47

4.25

1.64

Absolute dominance (m2/ha)

100.00

0.00

46.42

1.25

7.79

16.51

20.25

7.79

Relative dominance (%)

1.00

0.08

0.17

0.08

0.08

0.17

0.33

0.08

Absolute frequency

100.00

8.33

16.67

8.33

8.33

16.67

33.33

8.33

Relative frequency (%)

300.00

16.67

79.75

17.91

24.45

49.84

86.92

24.45

Importance value

Table 5.10.12. Upper montane forest dominated by Papuacedrus papuana, Saurauia trugal, Symplocos cochinchinensis, and Dacrycarpus imbricatus Montane Vegetation of Papua / 1019

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42.8 29.5 72.3

Species

Papuacedrus papuana

Dacrycarpus cinctus

Total

Source: Shea et al. (1998).

Total clear bole volume (m3/ha)

200

67

133

Absolute density

100.0

33.3

66.7

Relative density (%)

18.2

7.4

10.8

Absolute dominance (m2/ha)

100.0

40.6

59.4

Relative dominance (%)

0.50

0.17

0.33

Absolute frequency

100.0

33.3

66.7

Relative frequency (%)

Table 5.10.13. Upper montane forest dominated by the gymnosperms Papuacedrus papuana and Dacrycarpus cinctus

300.0

107.3

192.7

Importance value

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Montane Vegetation of Papua / 1021

Between 2,800 and 3,100 m the forests in the Idenburg area (Brass 1941) were dominated by Phyllocladus, Dacrycarpus compactus, Fagraea, and several species of Elaeocarpus. The subcanopy was dominated by Rapanea, Vaccinium, Litsea, Elaeocarpus, Rhododendron, Styphelia, and Pygeum. On heavily shaded slopes two species of tree ferns were common: Cyathea everta and C. bidentata. Several species of Loranthus grew as parasites. The exposed ridge crests had an undergrowth dominated by orchids, mosses, and some ferns, including Plagiogyria, Grammitis, Polypodium, and Meringium. Two species of Calamus were common at 2,200–2,300 and 2,650 m. Seral communities along streams changed markedly in species composition with small changes in altitude of only 150 to 200 m. Many contained elements of the subalpine grasslands: Viola, Tetramalopium, Anaphalis, Lycopodium, and Equisetum. At higher altitudes around Lake Habbema the forest was confined to the ridges, and in small patches associated with sinkholes. The forest was dominated by Papuacedrus, Phyllocladus, and Dacrycarpus compactus (Brass 1941).

similarities between upper montane forest and heath forest Upper montane and heath forest have many features of structure and physiognomy in common (Whitmore 1984). Some species also occur in both of these forest types. The forest canopy of both upper montane and heath forest is rather even, dense, and commonly with a high albedo (of pale color on aerial photographs). Trees have dense crowns, microphyll is the predominant leaf size, and the leaves tend to be held obliquely vertical, often closely placed on the twigs. The plant communities of both upper montane and heath forest include facies of low biomass in comparison with other forest types. Paths made by animals and human travelers remain open for a long time, and many species have hard, dense wood— two factors which suggest that growth rate is slow. Big woody climbers are absent or rare in both forest types.

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. Bull. Am. Mus. Nat. Hist. 79: 199–288. Ash, J. 1982. The Nothofagus Blume (Fagaceae) of New Guinea. Pp. 355–380 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr W. Junk Publisher, The Hague. Ashton, P.S. 1964. Ecological studies in the mixed Dipterocarp forest of Brunei State. Oxf. For. Mem. 25. Backhuys, W. 1968. Der Elevations-Effeckt bei einingen Alpenpflanzen der Schweiz. Blumea 16: 274–320. Bowler, J.M., G.S. Hope, H.N. Jennings, G. Singh, and D. Walker. 1976. Late Quaternary climates of Australia and New Guinea. Quaternary Research 6: 359–94. Brass, L.J. 1941. The 1938–1939 Expedition to the Snow Mountains, Netherlands New Guinea. J. Arnold Arbor. 22 (23): 271–345.

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1022 / r. j. johns , g. a . shea, w. v i n k, & p . puradyatmika Brass, L.J. 1964. Results of the Archbold Expeditions. No. 86. Summary of the sixth Archbold expedition to New Guinea: 1959. Bull. Am. Mus. Nat. Hist. 127 (4): 149–215. Bray, J.R., and E. Gorham. 1964. Litter production in forests of the world. Adv. Ecol. Res. 2: 101–157. Chappell, J.M.A. 1974. Geology of coral terraces, Huon Peninsula, New Guinea: a study of Quaternary tectonic movements and sea level changes. Bull. Am. Soc. Geol. 85: 553–570. Davis, T.A.W., and P.W. Richards. 1933. The vegetation of Morabali Creek, British Guiana; an ecological study of a limited area of tropical rainforest. J. Ecol. 21: 350–384. Edwards, P.J., and P.J. Grubb. 1977. Studies of mineral cycling in a montane rain forest in New Guinea. I. The distribution of organic matter in the vegetation and soil. J. Ecol. 65: 943–969. Enright, N.J. 1978. The ecology and population dynamics of Araucaria in Papua New Guinea. Ph.D. diss., A.N.U., Canberra. Enright, N.J. 1982. The Araucaria forests of New Guinea. Pp. 381–399 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr W. Junk Publishers, The Hague. Ewel, J. 1980. Tropical succession: manifold routes to maturity. Tropical Succession 2–7. Gibbs, L.S. 1917. A Contribution to the Phytogeography and Flora of the Arfak Mountains. Taylor and Francis, London. Gibbs, R. 1974. Fagaceae. Chemotaxonomy of Flowering Plants. 3: 1304–1309. McGillQueen’s Univ. Press, Montreal and London. Gray, B. 1973. Distribution of Araucaria in Papua New Guinea. Res. Bull. 1. Dept. of Forests, Port Moresby. Gray, B. 1975. Size composition and regeneration of Araucaria stands in New Guinea. J. Ecol. 63: 273–289. Gressitt, J.L. 1974. Insect biogeography. Ann. Rev. Entomol. 19: 293–322. Gressitt, J.L. (ed.). 1982. Biogeography and Ecology of New Guinea. Kluwer Academic Publishers Group, Dordrecht, The Netherlands. Griffiths, J.R., and R. Varne. 1972. Evolution of the Tasman Sea, Macquarie Ridge and Alpine Fault. Nature, Phys. Sci., London 235: 83–86. Grubb, P.J., J.R. Lloyd, T.D. Pennington, and T.C. Whitmore. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy and floristics. J. Ecol. 51: 567–601. Grubb, P.J., and T.C. Whitmore. 1966. A comparison of montane and lowland forest in Ecuador. II. The climate and its effects on the distribution and physiognomy of the forests. J. Ecol. 54: 303–333. Havel, J.J. 1965. Plantation establishment of Klinki Pine (A. hunsteinii) in New Guinea. Commonw. For. Rev. 44: 172–187. Havel, J.J. 1971. The Araucaria forests of New Guinea and their regenerative capacity. J. Ecol. 59: 203–214. Havel, J.J. 1972. New Guinea forests—structure, composition and management. Austr. For. 36: 24–37. Hegnauer, R. 1966. Chemataxonomie der Pflanzen. 4: 141–155. Biekhauser Verlag, Basel and Stuttgart. Hillis, W.E., and T. Inoue. 1967. The polyphenols of Nothofagus species 2. The heartwood of Nothofagus fusca. Phytochemistry 6: 59–67. Hnatiuk, R.J., J.M.B. Smith, and D.N. McVean. 1976. Mt Wilhelm Studies 2. The climatic of Mt Wilhelm. Publication BG/4. Research School of Pacific Studies, A.N.U., Canberra.

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Montane Vegetation of Papua / 1023 Hope, G.S. 1976. Vegetation. Pp. 112–172 in Hope, G.S., J.A. Peterson, I. Allison, and U. Radok (eds.) The Equatorial Glaciers of New Guinea.. Balkema, Rotterdam. Hynes, R.A. 1973. Ecology of Nothofagus forest in central New Guinea. Master’s thesis, Univ. Papua New Guinea. Hynes, R.A. 1974. Altitudinal zonation of forests in Malesia. Pp. 75–120 in Flenley, J.R. (ed.) Altitudinal Zonation of Forests in Malesia. Dept. Geogr., Univ. of Hull. Janzen, D.H. 1974. Epiphyte myrmecophytes in Sarawak: mutualism through feeding of plants by ants. Biotropica 237–259. Johns, R.J. 1976. A provisional classification of the montane vegetation of New Guinea. Science in New Guinea 4: 105–117. Johns, R.J. 1976–1978. Common Forest Trees of Papua New Guinea. 12 parts. PNG Office of Forests. Johns, R.J. 1980. Forest types of Papua New Guinea. The Dacrydium nidulum swamp forests of the Southern Highlands. Klinkii Johns, R.J. 1982. Plant zonation. Pp. 309–330 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Kluwer Academic Publishers, Dordrecht, The Netherlands. Johns, R.J. 1986 as 1976. The instability of the tropical ecosystem in Papuasia. Blumea 31: 341–371. Kalkman. C., and W. Vink. 1970. Botanical exploration in the Doma Peaks region, New Guinea. Blumea 18: 88–135. Lam, H.J. 1934. Materials towards a study of the flora of the island of New Guinea. Blumea 1: 115–159. Lam, H.J. (trans. A.M. Perry). 1945. Fragmenta Papuana [Observations of a Naturalist in Netherlands New Guinea]. Sargentia 5: 1–196. Lamprecht, H. 1972. Einige Strucktur Merkmale Naturlicher Tropenwaldtypen und Ihre Waldbauliche Bedeutung. Forstwiss. Cbl. 91: 270–277. Lane-Poole, C.E. 1925. The Forest Resources of the Territories of Papua and New Guinea. Govt. Printer, Melbourne. Mangen, J-M. 1993. Ecology and Vegetation of Mt. Trikora New Guinea (Irian Jaya/ Indonesia). Ministe`re des Affaires Culturelles. Travaux Scientifiques du Muse´e National d’Histoire Naturelle de Luxembourg. Paijmans, K. 1970. An analysis of four tropical rain forest sites in New Guinea. J. Ecol. 58: 77–101. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO in Association with Australian National University, Canberra. Rappard, F.W., and P. van Royen. 1959. Some notes on the vegetation in the Wissel Lakes area. Nova Guinea new ser., 10: 159–176. RePPProT. 1986. Review of Phase I and II results for Irian Jaya. Regional Physical Planning Programme for Transmigration (RePPProT), Ministry of Transmigration, Jakarta. RePPProT. 1990. The land resources of Indonesia. A national overview. Regional Physical Planning Programme for Transmigration (RePPProT), Ministry of Transmigration, Jakarta, and the U.K. Overseas Development Agency. Richards, P.W. 1952. The Tropical Rain Forest. Cambridge University Press, Cambridge. Robbins, R.G. 1958. Montane formations in the Central Highlands of New Guinea. Pp. 176–195 in Proc. Symposium on Humid Tropics Vegetation, Tijiawi, Indonesia. UNESCO. Robbins, R.G. 1970. Vegetation of the Goroka-Mt. Hagen area. Pp. 104–118 in Haantjens, H.A. (ed.) Lands of Goroka-Mt. Hagen Area. CSIRO, Land Res. Ser. 27. CSIRO, Melbourne.

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1024 / r. j. johns , g. a . shea, w. v i n k, & p . puradyatmika Robbins, R.G., and R. Pullen. 1965. Vegetation of the Wabag-Tari Area. Pp. 100–115 in Lands of the Wabag-Tari Area, Papua New Guinea. CSIRO Land Res. Ser. 15. CSIRO, Melbourne. Rodin, L.E., and N.I. Bazilevich (trans. Scripta Technica). 1967. Production and Mineral Cycling in Terrestrial Vegetation. Oliver and Boyd, London. Saunders, J.C. 1965. Forest resources of the Wabag-Tari Area. Pp. 116–124 in Lands of the Wabag-Tari Area, Papua New Guinea. CSIRO, Land Res. Ser. 15. CSIRO, Melbourne. Saunders, J.C. 1970. Forest resources of the Goroka-Mount Hagen area. Pp. 119–125 in Lands of the Goroka-Mount Hargen Area, Papua New Guinea. CSIRO, Land Res. Ser. 27. CSIRO, Melbourne. Schlinger, E.I. 1974. Continental drift, Nothofagus and some ecologically associated insects. Ann. Rev. Entomol. 19: 323–343. Shea, G.A., P. Puradyatmika, A. Maulensey, and D. Martindale. 1998. Biodiversity Surveys in the PT Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. 11 vols. PT Hatfindo Prima, Bogor, Indonesia. Troll, C. (1957). Tropical mountain vegetation. Pp. 37–46 in Proc. 9th Pacific Sci. Congress, Science Soc. of Thailand, Bangkok. van Steenis, C.G.G.J. 1934. On the origin of the Malaysian mountain flora I. Bull. Jard. Bot. Buitenz. ser. 3, 13: 135–262. van Steenis, C.G.G.J. 1950 or 1952. The delimitation of Malaysia and its main plant geographical divisions. Pp. ixx–xxv in van Steenis, C.G.G.J. (ed.) Flora Malesiana, Vol. 1(1). Noordhoff, Jakarta. van Steenis, C.G.G.J. 1971. Plant conservation in Malaysia. Bull. Jard. Bot. Nat. Belg. 41: 189–202. van Steenis, C.G.G.J. 1972a. The Mountain Flora of Java. Brill, Leiden. van Steenis, C.G.G.J. 1972b. Nothofagus. In Soepadmo, E., Fagaceae. Flora Malesiana ser. I, 7: 277–294. Vink, W. 1998. Notes on some lowland rainforests of the Bird’s Head peninsula, Irian Jaya. Pp. 90–109 in Barstra, G.-J. (ed.) Bird’s Head Approaches: Irian Jaya Studies. Balkema, Rotterdam. Walker, D. 1966. Vegetation of the Lake Ipea region, New Guinea Highlands. J. Ecology 57: 503–533. Walker, D. 1973. Highlands vegetation. Nat. Hist. 17: 410–414. Walker, D., and J.C. Guppy. 1976. Generic plant assemblages in the highland forests of Papua New Guinea. Aust. J. Ecol. 1: 203–212. Webb, L.J., and J.G. Tracey. 1967. An ecological guide to new planting areas and site potential for Hoop Pine. Aust. For. 31: 224–239. Whitmore, T.C. 1977. A first look at Agathis. Tropical Forestry Papers 11. Oxford. Whitmore, T.C. 1984. Tropical Rainforests of the Far East. Oxford Science Publications. Oxford: Clarendon Press. Whitton, A.J., M. Mustafa, and G.S. Henderson. 1987. The Ecology of Sulawesi. Gadjah Mada University Press, Yogyakarta. Womersley, J.S. 1958. The Araucaria forests of New Guinea. A unique vegetation type in Malaysia. Pp. 252–257 in Proc. Symp. Hum. Trop. Veg., Tjiawa. Zieck, J.F.U., J. Luitjes, and W. Vink. 1960. Tourneeversleg naar de noordelijke Wisselmerenstreken van 9 mei tot 1 juni 1960. Bosplanalogie, Manokwari: 17.

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5.11. Subalpine and Alpine Vegetation of Papua robert j. johns, garry a. shea, and pratito puradyatmika x t en s i v e a r e a s of high altitude vegetation span the mountain chain of New Guinea, forming ‘‘islands’’ of subalpine and alpine flora in a ‘‘sea’’ of montane and lowland forest. The subalpine zone occurs from ca 3,200 m to 4,170 m, and is subdivided into the lower subalpine zone from 3,200 m to 3,650 m and the upper subalpine zone from 3,650 m to 4,170 m. The subalpine zone is defined by the occurrence of subalpine forests, intermixed with other vegetation types, to 3,900 m. A tall shrubland occurs between 3,900 m and 4,170 m. This latter vegetation type does not occur on mountains in Papua New Guinea, and may indicate relatively milder conditions or more suitable soils than in Papua New Guinea. Uplift of the mountains in New Guinea, like New Zealand, occurred during late Tertiary and early Pleistocene times (Ollier 1986). In contrast, the Australian mountains, potentially an important source area for elements of the high altitude flora of New Guinea, are much older and were uplifted during Eocene/Miocene times. The earliest uplift of the mountains of Papua postdates that of Papua New Guinea, although ‘‘mountain building’’ continues throughout the island. During the Pleistocene the higher altitudes of Australasia were subjected to glacial and/or periglacial conditions. These factors have contributed to the unique high mountain vegetation and flora (Johns 1982; van Royen 1980). Next to Mt Jaya, only Mt Trikora (Mt Wilhelmina) has been studied (Mangen 1993). The following chapter refers almost exclusively to the work on Mt Jaya (Johns et al. 2006); detailed comparison was not made with the data from Mt Trikora. Areas of subalpine vegetation also occur along valley bottoms below the ca 3,000 m lower limit, as a consequence of various factors, including cold air drainage (Paton 1988), wet soils (Ashton and Hargraves 1983), exposure to strong winds (Ashton and Williams 1989), and periodic droughts and fire (Johns 1986). Patches of subalpine shrubbery and grassland on Mt Jaya occur along ridges below the forest limit particularly on mineral soils. Species that occur in these communities are also common as roadside weeds at midaltitudes and higher. Open subalpine savanna and shrubbery also occurs in areas of mineralization at lower altitudes (2,600–2,800 m) as observed on Mt Jaya to the southwest of the Army Camp (Johns et al. 2006). Casuarina oligophylla dominates communities on ultramafic rocks between 2,750 and 3,500 m elevation on Mt Doorman (Lam 1945).

E

Studies of High Altitude Vegetation in New Guinea Detailed studies on the ecology and vegetation history of the high mountains of New Guinea, were first made on Mt Wilhelm, which at 4,509 m, is the highest Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

1025

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1026 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

peak in Papua New Guinea. This research is summarized in a series of papers published by Walker (1968), Hnatiuk (1975), Hope (1976a, 1980), and Smith (1974). Wade and McVean (1969) recognized 27 vegetation communities above 3,110 m altitude, including three major types of forest on Mt Wilhelm. The first attempt to provide a detailed list of the high altitude species from Papua from was published by Hoogland (1958), largely based on their collections from an expedition in 1953 to Mt Wilhelm. A more comprehensive list of the vascular plants of Mt Wilhelm was published in 1972 by Johns and Stevens. Descriptions of the high altitude vegetation of Papua consisted mainly of observations compiled during several expeditions: Ridley (1916), Gibbs (1917), Colijn (1937), Brass (1941), Archbold, Rand, and Brass (1942), Lam (1945), Temple (1962), Cooper (1971), and Mangen (1993). Hope (1976b) collected plot data from the subalpine and alpine vegetation of Mt Jaya. When comparing the vegetation with other mountain ranges in New Guinea, Hope (1976b) emphasized the floristic variation between the different mountain ranges. G. S. Hope revisited Mt Trikora in early 1984; among his collections was Isoetes hopei, not previously collected from there. He found that the subalpine vegetation is different from that he has studied on Mt Jaya (Flora Malesiana Bulletin 9/2, 154, 1985). Hope (1976b) collected plot data using the releve´ method (described therein) and mapped the main formations occurring in the alpine and subalpine zones in and around the mine site on Mt Jaya (Figure 5.11.1). He also sampled a number of distinct communities. Figures 5.11.2 and 5.11.3 show the location of the plots (releve´s). In their study of the subalpine and alpine vegetation Shea et al. (1998) supplemented the vegetation data provided by Hope et al. (1976) in order to allow for the classification of the recognized community types. The following descriptions of the

Figure 5.11.1. Location map for the Mt Jaya study area. For the location of the releve´s, see Figures 5.11.2 and 5.11.3. Source: Hope (1976a).

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Subalpine and Alpine Vegetation of Papua / 1027

Figure 5.11.2. Location map for releve´s in and around the Carstensz Valley. Source: Hope (1976a).

vegetation types are based mainly on Hope (1976b), supplemented with observations made by Shea et al. (1998) and Johns et al. (2006).

Upper Montane-Subalpine Transition Forest In 1941, Brass described subalpine forest from Lake Habbema, dominated locally by Vaccinium dominans. Mangen (1993) described similar forest at 3,150 m on Mt Trikora that is dominated by Papuacedrus (Libocedrus) papuanus which forms a relatively open canopy with dense shrub and herb layers. Dacrycarpus compactus

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1028 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

Figure 5.11.3. Location map for releve´s at higher altitudes on Mt Jaya. Source: Hope (1976a).

and Phyllocladus hypophyllus are the other dominant trees in this community. The subcanopy, at 1–5 m tall, is composed of seedlings of Phyllocladus hypophyllus, Drimys piperita, Rapanea cucuminum, Prunus costata, Pittosporum pullifolium, Coprosma brassii, Xanthomyrtus compacta, Rhododendron brassii, R. versteegii, R. gaultherifolium, Styphelia suaveolens, and Trochocarpa nubicola. Common epiphytes are Phreatia, Dendrobium spp., Platanthera elliptica, and several small epiphytic ferns: Hymenophyllum rubellum, Humata pusilla, Selliguea plantaginea, and Grammitis fasciata. Myrmecodia cf. lamii is a common epiphyte with its tubers to 10–40 cm diameter and long branches up to 1 m long. This species also grows on the ground.

Subalpine Forest Subalpine forest is low to medium in height, with a closed main canopy, shrub layer, and ground cover. The canopy is 5–10 m in height, often dominated by the conifers Podocarpus, Dacrycarpus, and Papuacedrus, which sometimes emerge some 5 m above the canopy. Occasionally emergents occur to 15 m tall. The trees lack buttresses and stilt roots. No trees were observed with flowers and fruit on trunk or main branches (cauliflory or ramiflory). Dominant leaf-size classes are microphyll and nanophyll, but mesophyll and notophyll leaves may also be present; trees with compound leaves are rare and leaf drip-tips are absent. Large woody climbers are absent; small woody climbers are rare or absent. Vascular epiphytes and non-vascular epiphytes are abundant. Bryophytes in the ground cover are abundant. The following types of plants are absent: strangling figs, palms, bamboo, pandans and rattans. Tree ferns are occasional to common. The dominant families

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Subalpine and Alpine Vegetation of Papua / 1029

of trees and shrubs belong to the families Apiaceae, Araliaceae, Asteraceae, Cupressaceae, Epacridaceae, Ericaceae, Myrsinaceae, Myrtaceae, Podocarpaceae, Rubiaceae, Scrophulariaceae, and Winteraceae. On Mt Giluwe Pandanus occurs in the subalpine forest.

Lower Subalpine Forest On Mt Trikora the forests dominated by Papuacedrus are replaced by a dense evergreen scrub at 3,400 to 3,750 m altitude. Mangen (1993) established four plots in this type of forest on Mt Trikora. The crowns of the trees are often entangled, with sclerophyllous leaves and reduced leaf surfaces; most have simple leaves often with a dense indument on the lower surface or strongly revolute margins. Cyathea (3–4 species) have compound leaves. The trees form a dense canopy with a scattered undergrowth. Two species, Schefflera altigena and Saurauia alpicola, are emergent, reaching 7–8 m in height. Vaccinium dominans is abundant in the canopy, often with species such as Rhododendron correoides, R. gaultherifolia, Sericolea calophylla, Rapanea cacuminum, Olearia velutina, Drimys piperita, Coprosma brassii, and Xanthomyrtus compacta. Shrub layers are well developed, with Styphelia suaveolens, Trochocarpa nubicola, and grasses such as Agrostis rigidula var. remota, Oreobolus, and Ganhnia. Only three lianas have been collected: Rubus lorentzianus, Alyxia cacuminum, and Lycopodium clavatum. Epiphytes are common, including Cladomyza microphylla, Glomera sp., Octarrhena sp., Dendrobium dekockii, and three common ferns: Hymenophyllum foersteri, H. rubellum, and Selliguea sp. The stems, branches, and forest floor are covered with dense moss cushions. Established in these are a large number of ferns such as Plagiogyria tuberculata var. decrescens, Blechnum revolutum, Asplenium spp., and representatives of the Poaceae, Cyperaceae, and various dicotyledonous herbs. These forests are intermingled with small grassland patches that include many shrubs essentially the same as those in the subalpine forest. Dominant grasses are Deschampsia klossii, Agrostis rigidula var. remota, Danthonia oreoboloides, various Cyperaceae including Gahnia javaniva, Carex spp., and Oreobolus pumilio. Astelia papuana is often common in these grassland patches. Hope (1976b) established a plot (Plot 28) on the Kemabu Plateau, and also recorded the species in a stand near the Ertsberg mine. He noted the general similarity to analogous forest on other New Guinea mountains. Lower subalpine forest generally consists of a very dense canopy of low trees to about 10 m tall, with emergent more open crowned gymnosperms to 15 m tall. Characteristically, there is a thick epiphytic moss layer on branches and the ground, but relatively few epiphytic plants or woody climbers (lianas). Rapanea sp., Dacrycarpus compactus, and Papuacedrus papuana tend to be the dominant species. The dominant species recorded by Hope in Plot 28 were Rapanea sp., Dacrycarpus compactus, and Papuacedrus papuanus, that grow together with a large spiny Saurauia sp., and Rapanea to form the bulk of the lower tall scrub layer. Rhododendron culminicolum, Drimys piperita, Schefflera monticola, and Symplocos cochinchinensis var. orbic-

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1030 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

ularis are also common. These grow above a tangle of Coprosma brassii. The low shrub layer includes species such as Trochocarpa nubicola, Styphelia suaveolens, and various Rhododendron species. The herb layer is sparse but includes several ferns, especially Plagiogyria glauca, Pteris sp., and Blechnum sp. Some scrambling ground orchids, such as Glossoryncha sp. and Pedilochilus sp., form a tangle in places; Uncinia sp. and scattered Oreomyrrhis papuana are the only common herbs. Rubus spp., Parahebe albiflora, Sisyrinchium pulchellum, and Gleichenia bolanica are common in more open areas within the forest. The bryophytes form a luxuriant cover over the ground and on most tree branches, excluding most small vascular plants. Bright orange epiphytic orchids (Dendrobium sp.) provide the only source of vivid color on the olive or brown trunks and the somber green of the Dacrycarpus canopy. A prominent feature of the forests on the Kemabu Plateau is the widespread epiphytic and terrestrial Myrmecodia lamii, which grows up to 2 m in length. These were completely absent in the Ertsberg stand, which may be due to the high altitude (3,605 m), but have subsequently been collected at lower altitudes on mineral rich substrates on Mt Jaya. The forest patches on the Kemabu Plateau tend to occupy the moraine ridges or hill crests, and even here they are disturbed and penetrated by grassy glades. Dead trees and fire scars indicate that burning has been responsible for opening the forest, a feature common at higher altitudes in New Guinea due to El Nin˜o events (Johns 1986). The tangle of Coprosma brassii and the dense groves of tree ferns may have developed around the forest edge in response to this disturbance. The humus-rich soils usually average 30 cm deep, but can be very stony on some of the moraine areas. The trees become more stunted with increasing wetness and give way to mire communities on changes of slope. However, quite wet peaks can be colonized by forest, particularly where the water is moving through the soil profile. Hope (1976b) referred to this community as a Dacrycarypus compactus-Trochocarpa nubicola community. Shea et al. (1998) noted that the community is dominated by Rhododendron culminicolum, Papuacedrus papuana, and Saurauia sp., which all have a greater cover and dominance than Dacrycarpus compactus, while several shrubs have greater cover and dominance than Trochocarpa nubicola. The name appears inconsistent with the plot data. Hope (1976b) prepared a checklist of species for the Ertsberg site. He noted that work associated with the Ertsberg mine had resulted in the destruction of about 1.5 km2 of subalpine forest. Cuttings through the forest reveal that roots are concentrated in the topmost 40 cm of the organic rich soil, even where there is great variation in depth of sandy tilloidal sediments below the humus. Many roots appear to be active in the litter and moss-mat zone. The lower subalpine forest on the Kemabu Plateau is dominated by Dacrycarpus compactus, Papuacedrus papuana, Saurauia sp., and Schefflera monticola. Stand data are summarized in Tables 5.11.1 and 5.11.2. This stand is classified as Dacrycarpus compactus, Podocarpus brassii, Symplocos cochinchinensis, and Rapanea spp. dominated lower subalpine forest.

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PAGE 1030

1.10 46.16

Ryparosa sp.

Dacrycarpus compactus

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106.06

Source: Shea et al. (1998).

Total

0.53

12.83

Saurauia sp.

CH54

Rhododendron sp.

43.97

1.47

Total clear bole volume (m3/ha)

Papuacedrus papuana

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Schefflera monticola

Species

Total volume clear bole 0.02

1.38

0.05

0.51

0.74

0.52

Sum basal area 1.01

0.02

0.28

0.03

0.18

0.13

0.37

Total basal area (m2/ha) 24.46

0.52

9.42

0.52

4.58

8.37

1.05

Sum crown cover 32.19

63.59

58.52

120.10

78.93

15.11

Total crown cover (m2/ha) 12,147.9

1,072.8

2,119.5

1,884.0

4,003.5

2,564.3

503.7

121.3

10.7

21.2

18.8

40.0

25.6

5.0

Percentage cover (%)

Table 5.11.1. Lower subalpine forest, Kemabu Plateau, altitude 3,550 m

Absolute density 467

67

33

67

100

67

133

No. of subplots 2

1

2

2

2

4

Relative density (%) 100.00

14.29

7.14

14.27

21.43

14.29

28.57

Absolute dominance (m2/ha) 22.46

0.52

9.42

0.52

4.58

6.37

1.05

Relative dominance (%) 100.00

2.14

38.50

2.14

18.71

34.22

4.29

Absolute frequency 1.06

0.17

0.08

0.17

0.17

0.17

0.30

Relative frequency (%) 100.00

15.38

7.69

15.38

15.38

15.36

30.77

Importance value 300.00

31.81

53.33

31.61

55.53

63.89

63.62

Subalpine and Alpine Vegetation of Papua / 1031

PAGE 1031

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31.05 0.37 0.37 0.37 2.17

Dacrycarpus compactus

Rapanea spp.

Schefflera monticola

Rhododendron sp.

Saurauia spp.

Source: Shea et al. (1998).

52.10

15.71

Podocarpus brassi

Total

2.06

Total clear bole volume (m3/ha)

Symplocos cochinchinensis

Species

Total volume clear bole 0.011

0.011

0.011

0.931

0.471

0.062

Sum basal area 0.006

0.006

0.006

0.222

0.096

0.035

Total basal area (m2/ha) 13.59

1.03

0.26

0.26

0.26

7.39

3.21

1.18

Sum crown cover 12.56

3.14

12.56

113.83

50.24

47.89

Total crown cover (m2/ha) 8,659.0

654.2

418.7

104.6

416.7

3,794.2

1,674.6

1,596.2

Percentage cover (%) 88.6

6.5

4.2

1.1

4.2

37.9

18.8

16.0

Absolute density 300

33

33

33

33

67

33

67

No. of subplots 1

1

1

2

1

2

100.00

12.50

12.50

12.50

25.00

12.50

25.00

Relative density (%)

Table 5.11.2. Lower subalpine forest on a slope west of Carstensz Meadow, altitude 3,600 m Absolute dominance (m2/ha) 13.58

1.03

0.26

0.26

0.26

7.38

3.21

1.18

Relative dominance (%) 100.00

7.58

1.93

1.93

1.93

54.39

23.59

8.66

Absolute frequency 0.74

0.08

0.08

0.08

0.08

0.17

0.08

0.17

Relative frequency (%) 100.00

10.81

10.81

10.81

10.81

22.97

10.81

22.97

Importance value 300.00

18.39

25.24

25.24

25.24

102.36

45.90

58.63

1032 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

PAGE 1032

Subalpine and Alpine Vegetation of Papua / 1033

Upper Subalpine Forest The transition from lower subalpine forest to upper subalpine forest occurs on Mt Jaya at ca 3,650 m. Several tree species are rare or absent above this altitude, and the structure of those remaining become progressively lower and multi-trunked, with the exception of Dacrycarpus compactus, which persists as a straight-boled emergents even over 3,900 m. At the lower boundary, trees of Dacrycarpus compactus, Rapanea sp., and Drimys piperata are all more common than in the lower subalpine forest; Rhododendron culminicolum becomes co-dominant with Rapanea in most areas, particularly around 3,650–3,750 m. The forest is dense and tangled but interspersed with grassy glades and open patches where bare rock is exposed. The canopy can be 8–10 m high at lower altitudes, but the height decreases to about 6–8 m at 3,900 m. There is a shrubby understory into which many grassland species have penetrated, especially Styphelia suaveolens. However, upper subalpine forest also includes several forest species such as Xanthomyrtus linnaeifolia and Trochocarpa nubicola. The soils of the subalpine forests consist of a peaty litter and root mat often of little as 10 cm deep, overlying stony humus-rich loams or even rock and moraine. The taller forests in turn give way to a tall closed shrubland in some areas before the ‘‘treeline’’ (the limit of large woody plant growth) is reached along Discovery Valley and on the western and eastern slopes above the Carstensz Meadow. The tall shrubs and trees have locally different altitudinal limits along the track between Carstensz Meadow (3,650 m) and the lower Meren Valley (3,950). The following species reached their highest altitudes at: Glossorhyncha sp., 3,800 m; Rhododendron gaultheriifolium, 3,830 m; Schefflera monticola, 3,840 m; Podocarpus brassii, 3,840 m; Pittosporum pullifolium, 3,840 m; Dacrycarpus compactus, 3,850 m; and Xanthomyrtus linnaeifolia, 3,870 m. More studies are required to show the altitudinal limits of the species. Hope described a well-developed forest on the edge of the Dayak Meadow at 3,807 m (Plot 14B). The main canopy was 8–10 m tall, dominated by Dacrycarpus compactus, Rapanea sp., Drimys piperita, Symplocos cochinchinensis, and Rhododendron culminicolum. The latter was also common in the shrub layer along with Coprosma brassii. Table 5.11.3 describes the composition of the Dacrycarpus compactus, Podocarpus brassii, Rapanea spp., and Schefflera monticola dominated upper subalpine forest on the slope on the west side of Carstensz Meadow. Hope (1976a) referred to this community as a Coprosma brassii, Rapanea community. Rhododendron culminicolum has greater cover and dominance than either Coprosma brassii or Rapanea sp., and Shea et al. (1998) used the name Rhododendron culminicolum, Coprosma brassii, Rapanea sp., Dacrycarpus compactus upper subalpine low closedforest. This forest is documented in Table 5.11.4.

Subalpine Savanna A very distinctive area of open subalpine savanna occurs on Mt Jaya in an area of presumed high mineralization at 2,800–2,900 m altitude. The trees are broadly

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Source: Shea et al. (1998).

77.31

0.37

Total

Schefflera monticola

Podocarpus brassi 0.37

0.53 19.27

Rhododendron sp.

Cyathea sp.

1.10

55.67

Dacrycarpus compactus

Rapanea spp.

Total clear bole volume (m3/ha)

Species

Total basal area (m2/ha) 25.9

0.26

0.26

5.43

0.26

0.52

19.17

Total crown cover (m2/ha) 13,744.0

104.7

235.5

3,768.0

235.5

399.0

9,001.3

Percentage cover (%) 137.5

1.1

2.4

37.7

2.4

4.0

90.0

Absolute density 367

33

33

100

33

67

100

Relative density (%) 100.00

9.09

9.09

27.27

9.09

18.18

27.27

25.9

0.26

0.26

5.43

0.26

0.52

19.17

Absolute dominance

Table 5.11.3. Upper subalpine forest on a slope above Reclamation Nursery, altitude 3,750 m

Relative dominance (%) 100.00

1.01

1.01

20.96

1.01

2.02

73.99

Absolute frequency 0.91

0.08

0.08

0.25

0.08

0.17

0.25

Relative frequency (%) 100.00

9.09

9.09

27.27

9.09

18.18

27.27

Importance value 300.00

19.19

19.19

75.51

19.19

38.38

128.53

1034 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

PAGE 1034

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Source: Shea et al. (1998).

Total

Schefflera monticola 83.85

0.73

0.73

67.59

Dacrycarpus compactus

Cyathea sp.

12.96

Podocarpus brassi 1.47

0.37

Rhododendron spp.

Rapanea spp.

Total clear bole volume (m3/ha)

Species

Total basal area (m2/ha) 31.86

0.52

0.52

0.79

24.14

5.63

0.26

Total crown cover (m2/ha) 8,229.4

209.3

327.1

706.5

5,024.0

1,727.0

235.5

Percentage cover (%) 82.3

2.1

3.3

7.1

50.2

17.3

2.4

Absolute density 500

100

67

100

100

100

33

Relative density (%) 100.00

20.00

13.30

20.00

20.00

20.00

6.67

31.86

0.52

0.52

0.79

24.14

5.63

0.26

Absolute dominance

Table 5.11.4. Upper subalpine forest on a slope above Reclamation Nursery, altitude 3,800 m

Relative dominance (%) 100.00

1.64

1.64

2.46

75.77

17.66

0.82

Absolute frequency 1.25

0.25

0.17

0.25

0.25

0.25

0.08

Relative frequency (%) 100.00

20.00

13.30

20.00

20.00

20.00

6.67

Importance value 300.00

41.64

28.31

42.46

115.77

57.66

14.15

Subalpine and Alpine Vegetation of Papua / 1035

PAGE 1035

1036 / r o b e r t j o h n s, garry shea , a n d p r a t i t o p u r a d y a t m i k a

spaced, usually 5–8 m apart. Major species are Phyllocladus hypophyllus, Dacrydium sp., and Papuacedrus papuanus. The ground layer is very distinctive, being dominated by plants of the most unusual cyperaceous Mascherina, with its irislike foliage. Other unusual elements include Wittsteinia (Alseuosmiaceae), which was previously known from only a single collection in Papua New Guinea. Several epiphytic species grow on the branches and trunks of the canopy trees. Myrmecodia cf. lamii and clumps of Utricularia pulchra are scattered on the lower trunks throughout the forest. Mangen (1993) described a similar forest at 3,150 m on Mt Trikora dominated by Papuacedrus papuanus forming a relatively open canopy with a dense shrub and herb layer. Dacrycarpus compactus, sometimes to 50 cm dbh, and Phyllocladuus hypophyllous are the other dominant trees in this community. The subcanopy, at 1–5 m tall, is composed of Phyllocladus hypophyllus, Drimys piperita, Rapanea cucuminum, Prunus costata, Pittosporum pullifolium, Coprosma brassii, Xanthomyrtus compacta, Rhododendron brassii, R. gaultherifolium, R. versteegii, Styphelia suaveolens, and Trochocarpa nubicola. In 1941, Brass described a similar forest from Lake Habbema, which was characterized locally by Vaccinium dominans. Common epiphytes were Phreatia, Dendrobium spp., Platanthera elliptica, and several small epiphytic ferns: Hymenophyllum rubellum, Humata pusilla, Selliguea plantaginea, and Grammitis fasciata. Myrmecodia cf. lamii was a common epiphyte with its tubers to 10–40 cm diameter and long branches up to 1 m long. This species also grows on the ground.

Treeline Scrub and Shrublands Above 3,900 m, upper subalpine forest gives way to a scrub and shrublands. The loss of Dacrycarpus, reduction in frequency of Rhododendron culminicolum, and loss of the umbrella-like bunches of Schefflera foliage on their long thin trunks, gives this vegetation a distinct appearance. This is emphasized by the increasing frequency of the silvery tomentose Senecio carstenszensis. This tree-shrub daisy forms large clumps among the Rapanea sp. at 3,930 m together with the less pachycaulous Olearia velutina. With increasing altitude, Styphelia suaveolens increases in frequency inside the tall closed-scrub, and extends beyond it, with scattered Coprosma brassii bushes, in the shrub-rich grasslands at the tree line. The highest stand of tall closed-scrub was observed on the southern side of the lower Meren Valley where it reach an altitude of 4,170 m. The soils of the subalpine shrublands are similar to those in subalpine forests and consist of a peaty litter and root mat often as little as 10 cm deep, overlying stony humus-rich loams or even rock and moraine. The brittle surface of the limestone is eroded into a labyrinth of shallow knife-edged ridges (grikes). The limestone is often overgrown by a 4 m tall tangle of Coprosma brassii, which with its formidable rows of rigid, pungent pointed leaves makes some areas unpleasant to traverse. Plot 15 was established at 3,930 m in the lower Meren Valley within the tall closed-scrub (closed shrubland of Hope 1976b), about 100 m below the tree line. Here, Rapanea sp. and Drimys piperita occur as dense shrubs. Coprosma

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Subalpine and Alpine Vegetation of Papua / 1037

brassii, Olearia brassii, and Styphelia suaveolens are also common. Hope (1976a) classified this as a Coprosma brassii-Rapanea community. Again, this is inconsistent with the plot data, because Drimys piperita has the greatest cover and dominance in this vegetation. The cracks and pockets in the stone collect soil in which large shrubs and trees may grow and support shrublands on almost vertical cliffs. Hope (1976a) noted that the tangle of Coprosma brassii was virtually absent from the granitic Grasberg, and an increasingly open tall shrubland of Rapanea, Drimys, and Rhododendron correoides was replaced by shrub-rich grasslands at about 4,050 m. This vegetation has been completely removed to accommodate open-pit mining at the Grasberg, and the deposition of overburden.

Shrub-rich Grasslands Shea et al. (1998) established a plot in lower subalpine grassland and shrub grassland on a site located in the lower part of the subalpine zone. It is dominated by Deschampsia klossii, which is common on deep well-drained soil, with species of Monostachya, Poa, Festuca, and Danthonia, which dominate on shallow soils and poorly drained sites. This is an anthropogenic grassland resulting from repeated fires set by local people for hunting. In the absence of fire, the site will eventually revert to subalpine forest. Shrubs include species of Coprosma, Drimys, Olearia, Pittosporum, Rapanea, Rhododendron, and Vaccinium. The site and associated vegetation are described below. The low woodland or shrubland, generally has two layers, an open tree or shrub layer over mixed closed ground cover (grasses and forbs) with the height of trees and shrubs 1–5 m. Scattered emergent relict trees grow to 10 m tall. Buttresses and stilt roots are absent. No trees have flowers or fruit on trunk or main branches (cauliflory or ramiflory). The dominant leaf-size class is microphyll and nanophyll, but mesophyll and notophyll may also be present. Araliaceae trees with compound leaves are present. Leaf drip-tips are absent. Large and small woody climbers are absent. Vascular and non-vascular epiphytes are common on trees and larger shrubs; bryophytes are a common ground cover. Strangling figs, tree and shrub palms, bamboo, pandans, and rattans are all absent. Tree ferns are occasional to common. The dominant families are Apiaceae, Araliaceae, Asteraceae, Cupressaceae, Epacridaceae, Ericaceae, Myrsinaceae, Myrtaceae, Pittosporaceae, Podocarpaceae, Rubiaceae, Scrophulariaceae, and Winteraceae. Grasses and other herbs commonly belong to the families Asteraceae, Cyperaceae, Gentianaceae, Geraniaceae, Liliaceae, Plantaginaceae, Poaceae, Ranunculaceae, and Rosaceae. Hope (1976b) noted four shrub-rich grassland communities on Mt Jaya: forest edge grassland-shrublands; tree fern tussock grasslands; Coprosma brassii-Deschampsia klossii tussock grassland, and Gaultheria mundula-Poa nivicola tussock grasslands.

forest edge grassland-shrublands These communities occur in close proximity to forest and shrubland communities. Hope (1976b) gathered data from three plots. These plots may represent quite

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different communities, but each is growing near forest on very shallow soil or limestone, and the grasses have an opportunity to grow with the shrubs in conditions where a closed shrubland is unable to form and shade them out. Plot 30 was situated at 3,470 m on a limestone slope on the Kemabu Plateau, north of Carstensz Peaks, within an almost closed Cyathea sp. thicket. The tree ferns are about 2 m in height and interspersed with many large shrubs, so that grasses and clumps of Gahnia javanica make up only about 20 per cent of the cover. The nearest forest (lower subalpine forest) was at least 150 m distant and non-tree species occur in the stand. This site may have once supported forest, but soil erosion in the area has occurred, presumably after clearance, and reestablishment of trees is now unlikely. This community is classified as Coprosma brassii, Drimys piperita, Cyathea cf. pseudo-muelleri, and Rapanea sp. lower subalpine tall scrub. On the Kemabu Plateau, beyond the limits of the large Pleistocene moraines, the grasslands are dotted with mushroom shaded boulders, which testify to a removal or compaction of up to 1 m of former peaty soils, a process that may well postdate forest clearance there. The griking, karst development, and soil formation at the other two plot sites must postdate glacial retreat about 13,000 years ago. Solution is proceeding very rapidly, and soil accumulation is consequently very slow except in basins of deposition. Plot 14A (at 3,805 m) was only 5 m from subalpine forest on the Dayak Meadow and consisted of an open rock area with scattered low shrubs and numerous tuft grasses, principally Danthonia vestita and Poa egregria. This community is classified as Tetramolopium klossii, Danthonia vestita, Schoenus nitens, and Potentilla parvula lower subalpine openscrub. Plot 5 was situated on a steep limestone hill at the southern end of the Carstensz Meadow at 3,615 m. It also had tuft grasses growing in cracks in the limestone, with low bushes of Coprosma brassii and other shrubs growing in larger grikes. This community is classified as Styphelia suaveolens, Tetramolopium dekockii, Coprosma brassii, and Rhododendron spp. lower subalpine open scrub.

tree fern tussock grasslands This vegetation type has been variously classified as ‘‘shrub-rich grassland’’ by Wade and McVean (1969) and Hope (1976b) or as savanna by Paijmans (1976). Mangen (1993) follows the classification of Hope (1976a). Cyathea tomentosissima frequently dominates large areas on Mt Trikora between 3,000 and 3,500 m particularly in the Wamena-Baliem watershed. They form reasonably pure communities on Mt Trikora, the tree ferns growing to 2.5–5 m high. Common species are Rhododendron versteegii, Decaspermum nivalis, Sericolea calophylla, Rapanea cacuminum, Olearia velutina, Drimys piperita entity reducta, Coprosma brassii, and Xanthomyrtus compacta. Shrub layers are well developed with Styphelia suaveolens, Trochocarpa nubicola and grasses such as Agrostis rigidula var. remota, Oreobolus and Gahnia. Only three lianas have been collected: Rubus lorentzianus, Alyxia cacuminum, and Lycopodiuum clavatum. Epiphytes are common including Cladomyza microphylla, Glomera sp., Octarrhena sp., Dendrobium dekockii, and three common ferns, Hymenophyllum foersteri, H. rubellum, and Selliguea sp. The stems,

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branches, and forest floor are covered with dense moss cushions. Established in these are a large number of ferns such as Plagiogyria tuberculata var. decrescens, Blechnum revolutum, Asplenium spp., and representatives of Poaceae, Cyperaceae and various dicotyledonous herbs. They are mixed with grasses and smaller shrubs. In this vegetation type, Cyathea cf. pseudo-muelleri occurs as very open to dense groves with an almost continuous understory of tall tussock grasses, a rich herb flora and occasional low shrubs. The vegetation in the following plots is similar. Tree ferns, growing to 2–3 m in height, are found scattered through an open tussock layer of Deschampsia klossii, to about 40 cm in height, with numerous small tuft grasses growing in the gaps. On the ground are the creeping cushion grasses Monostachya oreoloides, Poa crassicaulis, and Poa sp. with Astelia papuana forming small cushions. Low bushes of Gaultheria spp. and Styphelia suaveolens occur within the grassland, generally outside the shade of the tree ferns. Both plots occupy well drained open sites, and the community is bordered by a pure stand of Deschampsia klossii tussock grassland or subalpine Astelia bog on wetter areas, and by forest on dry slopes. Plot 27 was located on a moraine crest on the Kemabu Plateau at 3,540 m. This tree fern-tussock grassland appears to be rather distinct from the very dense groves of shrubs and tree ferns along margins on the Kemabu Plateau, but might tend to change into the latter community in the absence of fire. The tree fern grasslands are one of the most extensive vegetation types in the area, covering at least one third (perhaps 80 km2) of the Kemabu Plateau. Based on cover and dominance data, this community is classified as Vaccinium amblyandrum-Danthonia vestitaCyathea cf. pseudo-muelleri-Deschampsia klossii-dominated lower subalpine medium to tall open-scrub. This community is referred to as ‘‘disclimax’’ because the community reflects the effects of fire, which has prevented the further development of this community into a lower subalpine forest. Plot 1 was located on a stream bank near the southern end of the Carstensz Meadow at 3,625 m asl. Here, the community occupied a 5 m strip between the boggy floor of the meadow and the forest edge. Shea (1998) refers to this community as Deschampsia klossii, Poa spp., Styphelia suaveolens, and Cyathea cf. pseudo-muelleri lower subalpine shrubland. This community reflects the effects of fire and cold air ponding, which has prevented the development of this community into lower subalpine forest.

coprosma brassii-deschampsia klossii tussock grassland This grassland is relatively shrub-free and consists of a complete cover of dense tussocks of Deschampsia klossii up to 1 m high, which almost completely exclude other species except for a few herbs and bryophytes that can occupy the wet tussock bases. Occasional stunted shrubs of Coprosma brassii and Styphelia suaveolens are present. This community occupies wet slopes, and gives way to short grass bog on wetter spots, and to a Gaultheria-Poa nivicola grassland on well drained slopes. This community is present in some form in all the New Guinea mountain grasslands. Two plots were recorded. Plot 2 was located at 3,620 m in Carstensz

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Meadow. Hope (1976b) referred to this community as Coprosma brassiiDeschampsia klossii tussock grassland. Deschampsia klossii is the dominant species but many other species have a higher cover and dominance than Coprosma brassii, including the shrubs Hypericum sp. and Vaccinium cf. wollastonii, the fern Belvisia sp., the grasses Bromus insignis, Hierochloe redolens, Carex brachyathera, and Carex gaudichaudiana, and a number of herbs. The community reflects the influence of repeated fires, which removed the original forest cover. Because of changes in the physical characteristics of the site and distance from the forest edge, succession back to forest will be likely to take a long time. Plot 21 was located at 3,985 m asl just north of Lake Larson. Hope (1976b) referred to this community as Coprosma brassii, Deschampsia klossii tussock grassland. The herbs Pilea sp. and Potentilla hooglandii have higher cover and dominance than Coprosma brassii. Several shrubs, grasses, and herbs have cover and dominance similar to that of Coprosma brassii. Based on cover and dominance the community was classified as Deschampsia klossii, Pilea sp., Potentilla hooglandii, and Coprosma brassii upper subalpine grassland.

gaultheria mundula-poa nivicola tussock grassland No plots were established in this community, but it closely resembled the Mt Wilhelm Coprosma divergens-Poa saruwagetica tussock grassland of Wade and McVean (1969) in its distribution and structure. It occurs on better drained moderate slopes, often as a relatively narrow band between forest and Deschampsia klossii, but can be extensive. Deschampsia klossii is a co-dominant with large tussocks up to 1 m in height interspersed with slightly smaller tussocks of Poa nivicola. Hierochloe redolens is always prominent, growing within the tussocks and up to 1.5 m tall. Shrubs are low, and vary in frequency from common to scattered. Coprosma brassii, Gaultheria mundula, and Styphelia suaveolens are the major species. This community can be classified as disclimax Deschampsia klossii-Poa nivicola-Hierochloe redolens-Gaultheria mundula-dominated lower and upper subalpine medium closedgrassland community. This community will slowly revert to forest. Wade and McVean (1969) felt that the Mt Wilhelm association was certainly secondary, replacing forest, but was now stable or only very slowly reverting to forest. Forest remnants and occasional tree ferns also occur in the Mt Jaya community so that it too may represent a former forest cover. Smith (1975) has noted an increase in size and number of shrubs in grasslands at 3,430–3,600 m on Mt Wilhelm, presumably associated with recent cessation of grass burning there.

Communities on Open Rocky Slopes A wide variety of communities become established on older landslides, rock fans, under rock overhangs, in the mouths of caves, and on the generally dry beds of streams. Hope (1976b, 1980) described four distinct communities from around the Carstensz Meadow at 3,680 m. Many communities occur at higher altitudes

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on the mountains and include several specialized plants that are often restricted to them. The crustose lichen communities are the initial colonizers of rock surfaces.

lichen field community Stereocaulon pseudomassartianum, a brittle thallose white lichen, forms an almost continuous layer about 5 cm deep. There are few associated species but moss tufts are occasional on larger boulders. This lichen is prominent on stream banks, but as a community it was mainly restricted to a granite-gravel fan on the western side of the Carstensz Meadow (Hope, 1976b: Plot 6: Carstensz Meadow at 3,685 m). The fresh fan surface consisted of loose deep gravel with virtually no fine fraction between the stones. This community is eventually replaced by tree ferns and shrubland, but the surfaces are continuously revered in gravel. Shea et al. (1998) called this a pioneer Stererocaulon pseudomassartianum, Campylopus archboldii, Campylopus tenuinervis, Selaginella dominated upper subalpine lichenland.

euphrasia lamii-tetramolopium distichum boulder community This community has ground orchids but no grasses. Mosses were mostly found growing in hollows on the boulders. Hope (1976b) described an interesting community (Plot 7: Carstensz Meadow at 3,670 m) from a large limestone boulder fan at the northeastern corner of the Carstensz Meadow. The boulder supply was limited and fine sediments filled the spaces between boulders, the fan was growing very slowly. The sparse field of herbs was dominated by scattered woody Euphrasia distichum (to 10 cm tall), which occupied most of the boulders. An unidentified gray crustose lichen colonizes the fresh limestone surfaces within about 30 years of exposure following the retreat of the glaciers. The herb component included scattered sedges. Shea et al. (1988) classified the community as a seral Tetramolopium distichum, Carex brachyathera, Schoenus maschalinus, Euphrasia lamii dominated upper subalpine shrubland.

rock overhang communities The limestone cliffs provide many natural overhangs, partially sheltered from the rain, in which heavily limed groundwater contributes to soil moisture. The vegetation consists of tuft grasses, particularly Danthonia vestita, Bromus insignis, and Brachypodium sylvaticum, together with dense patches of Cheilanthes papuana about 15 cm in height. Tetramolopium prostratum forms soft-leafed clumps among the scrambling Acaena anserinifolia (Hope, 1976b: Plot 4: Carstensz Meadow). The limestone wall is covered by a mat of Pilea spp., particularly the woody Pilea sp. (Hope herbarium collection, ANU 10893). Another urticaceous herb is Parietaria debilis. The vegetation is open in the dryer, most shaded parts of the floor of the overhang, with Parietaria debilis and Tetramolopium prostratum growing in the most protected areas. Bright orange and yellow crustose lichens (not identified) are present on the mineral efflorescences and sandy floor. Swiftlets have left guano in the soil in the driest areas. Shea et al. (1988) classified this community as seral

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Cheilanthes papuana, Acaena anserinifolia, Tetramolopium prostratum, and Poa cf. lunata lower subalpine short herbland.

crustose lichen communities on rock surfaces Although little studied, lichen communities are of great importance as there is a fairly complete lichen cover, even on cliffs, due to the moist climate. Gray, white, and black crustose lichens and several small thallose lichens are the initial colonizers of rock surfaces. The taxonomy of the species is little known (see Chapter 3.2). These communities were classified as pioneer crustose lichen subalpine open to closed-lichen land communities.

Transition Subalpine-Alpine Communities This vegetation type is located in the upper part of the subalpine zone near the transition to the alpine zone in the Dom Valley (Site 19). The site is dominated by subalpine grasslands and shrub-grasslands. The Deschampsia klossii at this site tends to be the viviparous variety, in which seeds germinate on the plant before being released, thus giving the plant an advantage at high elevations. The site and associated vegetation are described below. The medium to tall shrubland or grassland is generally two-layered: a tree or shrub layer over mixed closed ground cover (grasses and herbs), with the height of trees and shrubs to 1–2 m and scattered relict emergent trees to 5 m tall. Distinctive tree features: tree buttresses, stilt roots, breathing roots (pneumatophores), and trees with flowers and fruit on trunk or main branches (cauliflory or ramiflory) are all absent. The dominant leaf-size class: microphyll and nanophyll, but mesophyll and notophyll may also be present. Trees with compound leaves are present, mainly Araliaceae. Leaf drip-tips are absent. Large woody climbers and small woody climbers are absent. Vascular epiphytes and non-vascular epiphytes are common on trees and larger shrubs. Bryophytes are common ground cover. The following distinctive physiognomic features: strangling figs, palms, bamboos, pandans and rattans are absent. Tree ferns are occasional to common. The dominant families are Apiaceae, Araliaceae, Asteraceae, Cupressaceae, Epacridaceae, Ericaceae, Myrsinaceae, Myrtaceae, Pittosporaceae, Podocarpaceae, Rubiaceae, Scrophulariaceae, and Winteraceae. Grasses and other herbs commonly belong to the families Asteraceae, Cyperaceae, Gentianaceae, Geraniaceae, Liliaceae, Plantaginaceae, Poaceae, Ranunculaceae, and Rosaceae.

Subalpine and Alpine Mires and Bogs The high altitudes of Papua have large areas gently sloping country. Mire communities cover much of this in a mosaic, which presumably reflects differences in water supply and drainage. Six mire associations were recognized by Hope (1976b).

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poa lamii-vaccinium amblyandrum grass bog This community is widespread on level ground with impeded drainage but usually does not cover large areas. It consists of a dense or open sward of the tuft grass (Poa sp.) to 15 cm in height, with low shrubby Tetramolopium klossii scattered throughout. The ground layer is a dense mat of hard cushion plants such as Plantago stenophylla, Oreobolus pumilo, and Centrolepis philippinensis interspersed with soft cushions of hepatics (at Hamid Rock Shelter). Through this ground layer run the prostrate stems of the pink-fruited Vaccinium amblyandrum var. amblyandrum, which can form dense mats of up to 1 m2 in some places. Shea et al. (1998) includes data from a plot on Mt Jaya which indicate that the shrubs Tetramolopium klossii, Xanthomyrthus klossii, and Trochocarpa dekockii all have greater cover and dominance than Vaccinium amblyandrum. The monocots Agrostis reinwardtii, Eriocaulon sp., and Oreobolus pumilo, and the herbs Sagina papuana and Trachymene pulvilliforma all have greater cover and dominance than Vaccinium amblyandrum. Based on cover and dominance data this community is classified as Poa lamii, Tetramolopium klossii, Xanthomyrthus klossii, and Oreobolus pumilo lower subalpine low open scrub. A second plot was established by Hope (1976b: Plot 20) near the shore of Lake Larson at 3,980 m (Kemabu Plateau). Grass bog gives way abruptly to Deschampsia klossii on slopes, and to hard cushion bog in sites subject to frequent inundation. The soils are very deep acid peats with low mineral concentrations.

astelia papuana subalpine bog Astelia papuana forms hummocks up to 30 cm thick surrounded by tuft sedges, mats of mosses and hepatics, creeping Lycopodium spp., and scattered dwarfed shrubs of Styphelia suaveolens. Low tussocks of Deschampsia klossii may be present. Occasional larger bushes of Coprosma brassii may occur in the community, but except for the Astelia and thick hepatic cushions other plants tend to be low and ground hugging. A cover of lichens, especially Siphula thamnolioides, is common. The community is extensive on flat areas of granite gravel in the Carstensz Meadow, but also can occupy gentle slopes where surface runoff is high. The soils are peats or peaty gravels which appear to be fairly free-draining because the small puddles found between the cushions after rain disappear quickly. This community is analogous to the Astelia subalpine bog found on Mt Wilhelm and other New Guinea mountains but the subordinate species are very variable. Hope (1976b) established Plot 3 near the southern end of Carstensz Meadow at 3,605 m. This community is dominated by Astelia papuana, Styphelia suaveolens, Ranunculus tidens, and Epilobium detznerianum.

short grass bog This community resembles the Astelia subalpine bog in having a pronounced micro-relief of hummocks and water filled hollows, but always occupies deep, acid peats. The rises are occupied by clumps of Danthonia vestita and Deschampsia klossii interspersed with cushion grasses such as Monostachya oreoboloides, Poa

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pilata, and Poa sp. Many herbs, especially Plantago aundensis, Potentilla foersteriana, Gentiana ettingshausenii, and Ranunculus spp. also occur on the rises. The hollows are occupied by sedges, including Carex spp., and Scirpus cf. subtilissimus, and some hard cushion species. Drapetes ericoides also forms soft rounded hummocks. This community is very extensive to the north of Mt Jaya where it occupies flat but relatively well drained sites. In various forms the grass dominated bogs are common throughout the New Guinea mountains, and tend to border hard cushion bogs on wetter sites, and closed tussock grassland on slightly better drained areas. Two plots were established in this vegetation type by Hope (1976b). Plot 13 was located in the lower Meren Valley at 3,928 m. The dominant species in this community are Danthonia vestita, Carex spp., Plantago aundensis, and Monostachya oreoboloides. Plot 29 is located in Discovery Valley at 3,580 m asl. This site is part of a very large short grass bog that has formed on an old lake floor. This is now being incised by a deep stream channel, which probably allows the groundwater to drain away easily. The fine humus in the peats can be carried off by underground seepage, causing the characteristic hollows of a micro-pseudokarst. This process is probably accentuated by relatively faster growth on the rises tending to prevent surface drainage from the hollows.

hard cushion bog This community is found on wet, or even occasionally inundated, areas with little movement of surface or groundwater. Several species have adopted a hard spreading cushion habit. Some of these (e.g., Centrolepis philippinensis) tend to form a flat mat, but most can also form rounded cushions of varying sizes. One cushion of Rhododendron saxifragoides was 6 m in length and 40 cm high but most cushions are roughly circular and 5–30 cm in diameter and 5–10 cm in height. The deep green of Oreobolus spp. and Centrolepis philippinensis, the gray-green leafed Plantago polita, silvery Astelia papuana, and the lighter yellow greens of Monostachya oreoboloides, Potentilla brassii, and Eriocaulon spp. give a distinctive color to this community, in contest to the dark brown grasslands. Between the packed cushions are tufts of Carex spp. and Carpha alpina. The bog is absolutely firm to walk on, with little or no indentation. On cutting, a rigid mat more than 10 cm deep can be found. Hard cushion bog is widespread between 3,400 and 4,100 m on Mt Jaya, but usually forms a mosaic with other bog communities, particularly Poa lamii-Vaccinium bog and short grass bog. Plot 19A was located at the western end of Lake Larson at 3,980 m. The dominant species are Oreobolus pumilo, Monostachya oreoboloides, Carex spp., and Centrolepis philippinensis upper subalpine closed-grassland. Lake Larson Plot 19B is located at the western end of Lake Larson at 3,980 m. This community is classified as Oreobolus pumilo, Potentilla brassii, Centrolepis philippinensis, and Monostachya oreoboloides upper subalpine herbland or short closed-grassland community. Hard cushion bog is generally regarded as being confined to New Zealand, Tasmania, and Fuegia but is also known from the tropical Andes and Africa. Wade and McVean (1969) recognize a fragmentary community on Mt Giluwe and noted

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that short grass bog contains some hard cushions on Mt Wilhelm. Similar communities have been noted on Mt Albert Edward and Mt Scorpio, but on Mt Jaya the community is far more extensive. Both hard and soft cushion habits are far more common on Mt Jaya than have been noted elsewhere.

carpha alpina fen In this community, Carpha alpina forms an open sward to 10 cm with scattered cushion species, Gentiana sp., Ranunculus sp. and Potentilla brassii. The fen occupies the margins of small ponds, slight slopes, and ledges. It is a pioneer community on the calcite silt beaches of some tarns, provided that inundation is not constant. The Carpha alpina tufts can grow together to form hard round cushions, and the fen can thus grow to resemble short grass bog in structure and in the associated species. Carpha alpina fen often borders Carex gaudichaudiana fen and gives way to short grass bog or tussock grasslands. A very similar association is described on Mt Wilhelm and equivalents are also known from other New Guinea mountains. Plot 22 was recorded at the western side of Lake Larson at 4,210 m. The dominant species are Carpha alpina-Monostachya oreoboloides-Centrolepis philippinensis, and Plantago polita.

carex gaudichaudiana fen This fen community is a pioneer on bare peats and fluid muds. The sedge forms an open network across the surface of the sediments, often under 5–15 cm of water. Virtually no species are associated with the Carex. No plot was taken but the community is evidently identical to that described by Wade and McVean (1969). The fen was extensive where groundwater springs occurred in the Carstensz Meadow. Here mosses and a mat of Marchantia sp. grew over the sedge in the softest area, where the water was flowing sluggishly. Carpha alpina fen is usually the next stage in colonization. The dominant species are Carex gaudichaudiana and Marchantia sp.

Alpine Vegetation Alpine vegetation includes all the communities growing above the tall shrub limits (4,170 m). These are grasslands, heaths, and tundra. Some of them extend to lower altitudes in exposed or recently deglaciated areas, so that ‘‘alpine’’ cannot be defined in terms of all vegetation above a given altitude. Two grassland, two heath, and two tundra communities were examined. Plants grow on the highest points on which they can obtain a foothold. In addition to various bryophytes and lichens, Tetramolopium piloso-villosum, Geranium potentilloides var. alpestre, Scleranthus singuliflorus, and Epilobium detznerianun are found growing together at 4,595 m near the Meren Glacier. Ice cover and extreme steepness above that altitude prevent higher occurrences of herbs.

alpine tussock grassland Deschampsia klossii forms dense packed tussock grassland on deep well-drained soils from 4,000–4,500 m on Mt Jaya. The community differs from subalpine

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tussock grasslands in that the alpine tussocks are generally only 40 cm in height and there are no tall shrubs. Shrubs such as Styphelia suaveolens lie within the tussocks, together with many herbs, notably Papuzilla laeteviridis and the minute fern Cystopteris sp. Wade and McVean (1969) have noted a similar absence of alpine tussock grassland above 4,270 m on the cooler wetter aspects of Mt Wilhelm. Hope (1976a) has suggested that this is due to neoglacial disturbances rather than climatic limits. The closed alpine tussock grassland may well be the climax community for the alpine area. Given sufficient time for the development of deep peat soil, this grassland might eventually extend to higher altitudes. Neoglacial advances and retreats have prevented soil build up in most areas above 4,450 m. A sharp margin usually separates the grassland from the neoglacial advance and old avalanche areas. Hope (1976b) established Plot 23 located on the northern slopes of New Zealand Pass at 4,420 m asl. These grasslands follow this slope far to the west at high altitudes. This community is dominated by Deschampsia klossii, Potentilla foersteriana, Festuca crispato-pilosa, and Keysseria pinguiculiformis.

short alpine grassland This community is composed of scattered tuft grasses and occasional small tussocks, growing with scattered small shrubs to 40 cm tall. Plot 8 was established at 4,200 m in short alpine grassland on the exposed crest of the granite hill, the Grasberg. Here, there were scattered tuft grasses and occasional small tussocks. The main grasses were Agrostis reinwardtii, Deyeuxia brassii, Anthoxantium angustum, Monostachya oreoboloides, and Poa callosa. The ground was covered by bryophytes and lichens especially Rhacomitrium crispulum, Frullania reimersii, Cetraria spp., and Thamnolia vermicularis. Scattered shrubs were common, but never exceed 40 cm in height. The most common shrubs were Styphelia suaveolens, Tetraolopium ericoides, and Rhododendron correoides. This community was not found on equivalent areas on limestone, where shrubs tended to form an alpine heath. The peat soils were stony and shallow, and not very free-draining. Rock exposures were shattered and frost action was presumably intense. This community has subsequently been removed to make way for mining activities at Grasberg. The dominant species in this community were Agrostis reinwardtii, Deyeuxia brassii, Styphelia suaveolens, and Tetramolopium ericoides.

tetramolopium klossii-rhacomitrium heath This community had not been described before in New Guinea: its habitat is apparently almost entirely restricted to Mt Jaya. The areas covered by the last neoglacial advance are stony moraine fields that have been exposed by a steady ice retreat over the past 120 years. Tetramolopium klossii heath occupies the zone which has been ice free for more than about 30 years, and thus extends in the Meren Valley between 3,950 m and 4,200 m, and in New Zealand Pass (where the ice advance was restricted) between 4,250 and 4,450 m. Tetramolopium klossii grows as scattered low shrubs to 30 cm, rooted in a sparse moss carpet of Rhacomitrium

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crispulum, Bryum cf. rugicollum, and Distichum capillaceum. Styphelia suaveolens, and Vaccinium cf. coelorum grow as prostrate or creeping shrubs, with increasing frequency on the oldest moraines. Plot 9 was located on a moraine field in the upper Meren Valley at 4,210 m. Tetramolopium klossii and Vaccinium cf. coelorum grow along with a number of herbs and grasses, including Epilobium detznerianum, Gnaphalium breviscapum, and Scleranthus singuliflorus. These grow in a mat of mosses, including Rhacomitrium crispulum, Bryum cf. rugicollum, and Distichum capillaceum. Plot 10 is located in the lower Meren Valley at 4,005 m asl. Tetramolopium klossii, Vaccinium cf. coelorum, and Styphelia suaveolens grow along with herb and grasses, including Gnaphalium breviscapum, Anaphalis mariae, Geranium potentillioides var. alpestre, Plantago polita, Scleranthus singuliflorus, and Poa callosa. These grow in a mat of mosses. Bare ground makes up 30–70% of the area. Plot 11 is located in the lower Meren Valley at 4,050 m asl in one of the older moraine areas. Plot 11 contains occasional small shrubs of Coprosma brassii and patches of Tetramolopium distichum, Anaphalis mariae, Keysseria wollastonii, sedges, Pteris montis-wilhelminae, and Grammitis sp., Styphelia suaveolens grows as an erect shrub, and Rhododendron ultimum, with its very showy large red flowers, occurs as low bushes to 50 cm. Rounded cushions of Scleranthus singuliflorus and Sagina sp., tufts of Epilobium detznerianum, and individual plants of Deschampsia klossii are common. Bare ground makes up only 10–30% of the area. Plot 24 is located in the Yellow Valley at 4,278 m asl. Scattered Styphelia suaveolens and Tetramolopium klossii occur along with a number of herbs and shrubs, including Epilobium detzernerianum, Geranium potentillioides var. alpestre, Scleranthus singuliflorus, and Poa callosa. These grow in a mat of mosses, including Rhacomitrium crispulum, Bryum cf. rugicollum, and Distichum capillaceum.

dwarf shrub heath This community occupies ridge crests and slopes above 4,200 m and outside the area affected by neoglacial advance. It consists of a shrub mat up to 20 cm in depth, largely composed of Styphelia suaveolens with Tetramolopium klossii, Tetramolopium piloso-villosum, and very occasional stunted Coprosma brassii and sterile Senecio sp. shrubs. Deschampsia klossii and Monostachya oreoboloides occupy gaps in the heath, together with the ubiquitous Geranium cushions, Epilobium detznerianum and Parahebe vanderwateri. No plot data were obtained from this community, which was only observed among the limestone boulders and cliffs along the crest of the Midden Ridge, between the Yellow and Meren valleys. This community is analogous to the restricted dwarf shrub heath of Mt Wilhelm, but is not particularly extensive on Mt Jaya, despite conjecture to the contrary by Wade and McVean (1969). It is apparently the characteristic community on exposed crests with shallow soil, where tussock grassland cannot gain a foothold. Snowfall may be a factor in its distribution. It is bordered by alpine tussock grassland and also has a diffuse border with Tetramolopium klossii heath.

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dry alpine tundra The most recent moraines at 4,230–4,600 m asl have been exposed by steady ice retreat over the last thirty years and are being colonized by mosses and a very few herbaceous species able to grow in the alkaline mineral soil. As the snouts of the glaciers retreat, Epilobium detznerianum appears within a few months. After 12 months (based on extrapolation from long- and short-term glacier rates) small cushions of Distichum capillaceum, Bryum cf. rugicollum, and Scleranthus singuliflorus start to develop, initially on flat areas of fine sediments. Tufts of the purplish viviparous form of Deschampsia klossii can also become established. Although tussocks are not formed, the tufts become robust and common within a few years, and mats of Epilobium cf. prostratum and Rhacomitrium crispulum become extensive. Other herbs, such as Sagina sp., Keysseria wollastonii, and Pilea sp. start to appear in rock crevices. Poa wisselli forms common purple tufts to 15 cm. However, mosses continue to provide the major cover. This community is seral to Tetramolopium klossii heath, at least at altitudes of less than 4,500 m. Wade and McVean (1969) describe a similar community on Mt Wilhelm, but Hope (1976a) has shown that the alpine tundras are restricted to the area affected by an erosive episode which was probably linked with the last neoglacial period. However, bryophytes, very scattered tuft grasses, and the few other herbaceous species in these communities do provide a tundra-like vegetation which owes its origin not only to extremely cold, wet conditions but also to longterm variation in climate. It is possible that alpine tundras described from other parts of the world may also partly reflect a limited period of development. Plots 16, 17, and 18 represent sites ranging from an estimated 25 years to 15 and 10 years ice free, respectively. Plot 16 was located above the Northwall north of the upper Meren Valley at 4,480 m, on moraines that had been free of ice for 25 years. The most common herbs are Scleranthus singuliflorus, Keysseria wollastonii, Epilobium detznerianum, Pilea sp., and Sagina sp. Scattered grasses include Poa wisselii and Deschampsia klossii. Plot 17 was located in the upper Meren Valley at 4,245 m, on moraines that had been free of ice for 15 years. At this site, scattered Epilobium detznerianum and Scleranthus singuliflorus grow among patches of mosses, including Barbula wisselii, Bryum cf. rugicollum, and Distichium capillaceum. Bare ground covers a high percentage of the site. Plot 18 was located in the upper Meren Valley at 4,245 m on a moraine that had been free of ice for 10 years. Scleranthus singuliflorus, Epilobium detznerianum, and Epilobium cf. prostratum grow with scattered patches of Bryum cf. rugicollum.

wet alpine tundra The extremely free-draining limestone moraines at high altitudes provide little opportunity for the accumulation of shallow water, and surface streams are intermittent. The Yellow Valley has a very flat floor crossed by numerous low moraines. Behind some of these and in a few rock basins a continuous moss mat supports a few herbaceous species to give a community which appears analogous to the wet alpine tundra described by Wade and McVean (1969) from Mt Wilhelm. The

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major moss, which is tentatively identified as Breutelia aristifolia, is caked in limestone silts washed in periodically from the surrounding till. In it grow small cushions of Gnaphalium breviscapum, Geranium potentilloides var. alpestre, and Ranunculus spp. Sedges are scattered to common, with occasional stunted tufts of Deschampsia klossii. This community occurs at the relatively low altitude of 4,250 m, and is surrounded by Tetramolopium heath. Again its status as a true alpine tundra can be questioned. It can be regarded as a mire community. Plot 25 is located in the Yellow Valley at 4,280 m asl. The main species here is Carex brachyathera, with 30–50% cover. Associated herbs include Ranunculus sp., Plantago aundensis, Epilobium detzernianum, Gnaphalium breviscapum, and Sagina sp. Deschampsia klossii occurs as scattered tufts. Plot 26 is located in the Yellow Valley at 4,300 m asl. This site is dominated by the moss Breutelia aristifolia, which has a cover of over 75%. The herb Gnaphalium breviscapum and the grass Schoenus sp. are scattered in the site.

Cryovegetation The cryovegetation of Mt Jaya was described by Kol and Peterson (1976), while Peterson (1976) describes the englacial lakes where some cryoassociations occur. The cryovegetation is found growing on ice and snow in the nival zone. The following account is extracted from Kol and Peterson (1976) and Peterson (1976). The lower parts of the Meren and Carstensz glaciers appear rough and dirty to the casual observer, in contrast to the white snow and pale blue ice in the crevasses. The ice surface is pitted like rough concrete, especially on gentle slopes, and the pits and cracks are full of dark specks or small black flakes. Scattered across the ice are pools and englacial lakes similar to those noted by Colijn (1937). These vary from a few centimeters to 10 m across, are up to 4 m deep, and contain very clear water in which scattered black mats several centimeters in diameter rest on the ice-bottom or float across the top. The ice itself is clean and clear, except for a few particles of rock, because the tropical jungle and frequent rains at lower altitudes prevent much dust or other debris from rising in the atmosphere to be incorporated in the snow that forms the glacier. Observation of cryovegetation comprising many different species and exhibiting varied macroscopic characteristics have been reported from snow and ice in polar, subpolar, and temperate latitudes (Kol and Peterson 1976). The Mt Jaya ice supports several different cryophilic associations: black, red, and yellow-brown ice growths, black englacial-lake growths, and red and yellow snow growths. The only other equatorial cryohabitat studied at the time was the Pichincha volcano, Ecuador. Pichincha supports red snow growths, but ice communities have not previously been reported from other equatorial areas. Kol and Peterson (1976) found the following species of algae growing on the various ice and snow areas on Mt Jaya: Chlamydomonas antarcticus, Chlorosphaera antarctica, Scotiella antarctica, S. nivalis, S. norvegica var. carstenszis, Mesotaenium berggrenii, and Nostoc fuscescens var. carstenszis. The species composition of the

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cryoassociations determines their microscopic appearance and seems to reflect physical rather than chemical differences in the habitats. Table 5.11.5 lists the associations and their component species. The most extensive growth occurs in the ice ablation zones of the Meren and Carstensz glaciers and consists of the black ice and the yellow-brown ice associations. In general, the more horizontally-oriented surfaces support the black associations and the steeper slopes the yellow-brown associations. Thus, areas of ice ablation crust with relatively large ice crystal structure supported discrete, dense black colonies ranging in size from just visible up to 3–4 cm in diameter. Groups of these colonies formed the most common and extensive cryovegetation in the ablation zones. They are dominated by Nostoc fuscescens var. carstenszis, whose filaments are purple in transmitted light. Areas of the glacier with a fine to medium ice crystal structure supported diffuse yellow-brown communities. These yellow-brown discolorations were also extensive. They showed a distinct variation in color intensity, which reflected a variation in numbers rather than species. This association was dominated by Mesotaenium berggrenii. Towards the snow line smaller communities of diffuse pink and diffuse crimson ice existed. These were much less common, and relatively small in area, in comparison to the black and yellow-brown communities. Only the occasional steep

Table 5.11.5. Species composition of cryoassociations on Meren and Carstensz glaciers Black ice

Black lake

Association YellowRed brown ice ice

Yellow snow

Red snow

Chlamydomonas antarcticus













Chlorosphaera antarctica













Scotiella antarctica













Scotiella nivalis













Scotiella norvegica var. carstenszis nova var.













Mesotaenium berggrenii













Nostoc fuscescens var. carstenszis nova var.













Species

Note:  indicates a species in present in a particular ice type, indicates that the species is absent. Source: Kol and Peterson (1976).

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slope with a southeasterly aspect was completely free from biological discoloration in the ice-ablation zone. The englacial lakes of the Meren Glacier, which ranged in size from a few centimeters to several meters in depth and diameter, were universally colonized by large (15–30 cm diameter  3–7 cm depth), discrete, dense black colonies. Cryovegetation above the snow line was dominated by a diffuse yellow-brown community. However, small areas of crimson firn (granular snow) were not infrequent. Discrete black colonies were present in this region between the snow line and the firn line but they were always small and very scattered, never forming the extensive communities so characteristic of the lower slopes of the glaciers. According to Kol’s classification scheme for cryohabitats (Kol 1968), the Mt Jaya ice body would be classified as calcareous. However, pH values obtained from the ice, snow, and surface meltwater ranged between pH 4–5 and were thus lower than are usually encountered on permanent ice bodies lying over limestone bedrock in the northern hemisphere. Certainly, the species composition of the snow associations is more typical of a silicotrophic than a calcareous habitat. Total hardness measurements (made using the Pallin Test) on the ice, snow, and surface meltwater gave low readings of approximately 2,015 ppm. The concentrations of specific minerals were too low to be detected with the methods available. There were no consistent differences between the different communities in pH or total hardness characteristics. Rock fragments were a variable but an ever-present constituent of all the communities.

Englacial Lakes These lakes occupy enclosed depressions in the glacier surface, with the wider depressions generally containing the deeper lakes. Few of the lakes have glacial streamlets, and no more than half are fed directly by englacial streamlets. The surface of the glacier has a pronounced weathering crust that causes the depression walls to be whitened and toughened. These walls commonly hold fresh snow if nearer the firn line, and less commonly further down the glacier. Both snow and ice cryovegetation can be found on these walls, which slope outward in most cases, giving most depressions the shape of a funnel with a short, wide, rounded and closed-off outlet. The lakes are contained in the closed end and their steep walls are entirely in blue ice, there being, of course, no weathering crust beneath water level. In 1972, the largest englacial lakes occurred in the ablation zone on the south side of the glacier. They are also found directly down-glacier from the lowest part of the accumulation zone. The development of these lakes may be favored by lower ice velocities. An additional factor may be the greater compression of ice on the south side of the glacier, where a southward component in the glacier flow pushes ice against the Midden Ridge. Thus, the permeability of the ice is reduced and any tendency for transverse crevasses to develop is diminished. The algae were most prolific in hollows in the bottoms of the deeper lakes and

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temperatures inside the colonies were above 0C, and as much as 4C during the daytime. Absorption of radiation by the algae led to a marked daytime thermal gradient from the surface (0C) to the bottom (generally from 2–4C). This would have begun breaking down before sunset but only rarely did surface freezing occur in the higher lakes overnight. The heat generated is apparently dissipated into the meltwater and glacial ice, and a rather equable thermal region is established in which lakes remain above 0C at all times, even though meltwater from elsewhere on the glacier almost ceases during the cold nights. A very minor amount of mineral matter can be seen in the bottom of some of the lakes, but never enough to account for what we should now regard as the biological equivalent of cryoconite holes. These cryovegetation lakes are the largest reported. The particular factors that favor their formation in the Mt Jaya area are probably the optimum balance between incoming radiant energy, proportion of ultra-violet radiation, partial pressure of oxygen, suitable glacier physiography, and ice impermeability. By contrast, in high latitudes where most work on melt phenomena appears to have taken place, indirect radiation is important and cryovegetation (never as prolific as on Mt Jaya) is insignificant. Inorganic debris predominant at these high latitudes, and holes more frequently freeze over and meltwater is rare or absent. The cryovegetation of Mt Jaya add a new dimension to the study of cryoconite phenomena, and further detailed micrometeorological, glacioclimatic, and biological programs should be mounted while sections of the retreating glaciers still exist that are favorable to the formation of these lakes.

References 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. Bull. Am. Mus. Nat. Hist. 79: 199–288. Ashton, D.H., and G.R. Hargraves. 1983. Dynamics of subalpine vegetation at Echo Flat, Lake Mountain, Victoria. Proc. of the Ecol. Soc. of Australia 12: 35–60. Ashton, D.H., and R.J. Williams. 1989. Dynamics of the subalpine vegetation in the Victorian regions. In Good, R. (ed.) The Scientific Significance of the Australian Alps. Australian Alps National Parks Liaison Committee/Australian Academy of Science, Canberra. Brass, L.J. 1941. The 1938–1939 Expedition to the Snow Mountains, Netherlands New Guinea. J. Arnold Arb. 22: 271–342. Colijn, A.H. 1937. Naar de Eeuwige Sneeuw van Tropisch Nederland. Scheltens and Giltay, Amsterdam. Gibbs, L.S. 1917. A Contribution to the Phytogeography and Flora of the Arfak Mountains. Taylor and Francis, London. Hnatiuk, R. J. 1975. Aspects of the growth and climate of tussock grassland in montane New Guinea and sub-Antarctic islands. Ph.D. diss., Australian National University, Canberra. Hoogland, R.D. 1958. The alpine flora of Mt Wilhelm. Blumea, Suppl. 4: 220–238.

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Subalpine and Alpine Vegetation of Papua / 1053 Hope, G.S. 1976a. The vegetational history of Mt Wilhelm, Papua New Guinea. J. Ecol. 64: 627–663. Hope, G.S. 1976b. Vegetation. Pp. 112–172 in Hope, G.S., J.A. Peterson, I. Allison, and U. Radok (eds.) The Equatorial Glaciers of New Guinea. Balkema, Rotterdam. Hope, G.S. 1980. New Guinea mountain vegetation communities. Pp. 153–222 in van Royen, P. (ed.) The Alpine Flora of New Guinea. J. Cramer, Vaduz. Hope, G.S., J.A. Peterson, I. Allison, and U. Radok (eds.). 1976. The Equatorial Glaciers of New Guinea. Balkema, Rotterdam. Johns, R.J. 1982. Plant zonation. Pp. 309–330 in Gressitt, J.L. (ed.) Biogeography and Ecology of New Guinea. Dr. W. Junk Publishers, The Hague. Johns, R.J. 1986. The instability of the tropical ecosystem in Papuasia. Blumea 31: 341–371. Johns, R.J., P.J. Edwards, T.M.A. Litteridge, and H.C.F. Hopkins, 2006. A Guide to the Alpine and Subalpine Flora of Mount Jaya. Kew Publishing, Royal Botanic Gardens, Kew. Johns, R.J., and P.F. Stevens. 1972. Mt Wilhelm flora: a check list of the species. Botany Bulletin 6. Dept. of Forests, Papua New Guinea. Kalkman, C. 1963. Description of vegetation types in the Star Mountains region, West New Guinea. Nova Guinea Bot. 15: 247–261. Kol, E., and J.A. Peterson. 1976. Cryobiology. Pp. 81–91 in Hope, G.S., J.A. Peterson, I. Allison, and U. Radok (eds.) The Equatorial Glaciers of New Guinea. Balkema, Rotterdam. Lam, H.J. (trans. L.M. Perry). 1945. Fragmenta Papuana. Sargentia 5: 1–196. Mangen, J-M. 1993. Ecology and Vegetation of Mt Trikora New Guinea (Irian Jaya/ Indonesia). Ministe`re des Affaires Culturelles, Travaux Scientifiques du Muse´e National d’Histoire Naturelle de Luxembourg. Ollier, C.D. 1986. The origin of alpine landforms in Australasia. Pp. 2–36 in Barlow, B. (ed.) Flora and Fauna of Alpine Australasia. CSIRO/Brill, Melbourne. Paton, D.M. 1988. Genesis of an inverted tree line associated with a frost hollow in southeastern Australia. Aust. Journ. Bot. 36: 655–663. Paijmans, K. (ed.). 1976. New Guinea Vegetation. CSIRO in association with Australian National University, Canberra. Ridley, H.N. 1916. Botany of the Wollaston Expedition to Dutch New Guinea, 1912–1913. Trans. Linn. Soc. London, ser. 2, Bot., 9: 247–351. Shea, G.A., D. Martindale, P. Puradyatmika, and A. Mandessy. 1998. Biodiversity Surveys in the PT Freeport Indonesia Contract of Work Mining and Project Area, Irian Jaya, Indonesia. PT Freeport, Indonesia. Smith, J.M.B. 1974. Origins and ecology of the non-forest flora of Mt Wilhelm, New Guinea. Ph.D. diss., Australian National University, Canberra. Smith, J.M.B. 1975. Mountain grasslands of New Guinea. Journal of Biogeography 2: 87–101. van Royen, P. (ed.). 1980. The Alpine Flora of New Guinea. J. Cramer, Vaduz. Wade, L.K., and D.N. McVean. 1969. Mt Wilhelm Studies I: The Alpine and Subalpine Vegetation. Australian National University BG/1, Canberra. Walker, D. 1968. A reconnaissance of the non-arboreal vegetation of the Pindaunde Catchment, Mt Wilhelm, New Guinea. J. Ecol. 56: 455–466.

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5.12. Grassland and Savanna Ecosystems of the Trans-Fly, Southern Papua michele bowe, neil stronach, and renee bartolo n c on t r a st w i th most of Papua, the Trans-Fly region of the south-central bulge of the island of New Guinea has a distinctly monsoonal climate. Almost 75% of the annual 1,875 mm of rain falls in the December to May wet season, the remainder falling in a dry season from June to November (Paijmans et al. 1971). This has resulted in the development of vegetation types dominated by grassland and savanna, fringed by monsoon forests. With a maximum elevation of some 45 m, the landscape is unique in Papua for its open flatness as distinct from the rugged mountainous interior of the island, and it strongly resembles the coastal and adjacent areas of northern Australia. The Trans-Fly region straddles the international border with Papua New Guinea and the monsoonal climate affects a region of 101,274 km2, approximately half of which lies in Papua. Together with the climate, two major land forms influence the development of vegetation in this southern part of Papua: the coastal plain and the Oriomo plateau. These land forms were originally described from Papua New Guinea, following systematic surveys by the Commonwealth Scientific and Industrial Research Organisation (CSIRO) in the Morehead-Kiunga border region with Papua (Paijmans et al. 1971). The two land forms run southeast-northwest across the region. The coastal plain consists of low coastal beach ridges and intervening swales and riverine flats and, further inland, a back plain crossed by numerous smallmedium sized rivers. From west to east these include the Bian, Kumbe, and Maro rivers (in Papua) and the Torassi River, the mouth of which forms the international border between Indonesia and PNG. The coastal back plain varies in width from about 40 km close to the border with PNG to around 15 km further to the west. Generally the elevation does not exceed 5 m above mean sea level. The flats are inundated by freshwater in the wet season. Relict beach ridges are well drained and with sandy soils and the vegetation is dominated by littoral forest or Imperata grassland. The inland back plain becomes almost completely inundated in the wet season, which, together with the flooded coastal flats, gives rise to Papua’s most extensive wetlands. While the back plain largely dries out in the dry season, some limited areas of permanent swamp remain. These areas are of critical importance to the Trans-Fly’s large populations of resident and migratory waterfowl and waders. The vegetation of these temporarily flooded areas is low sedge grassland with scattered Nauclea and Pandanus trees. In the swampy areas delineated by old river channels and floodplains of smaller rivers, the vegetation consists of a mix of

I

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

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Phragmites and tall sedge grasslands in permanent swamp; low swamp grassland and semi-permanent to seasonal Melaleuca swamp forest; and Melaleuca savanna on higher ground (Paijmans et al. 1971). The Oriomo plateau is dissected by a number of smallish rivers with narrow catchments that drain southwards into the coastal plain. The plateau itself is low lying with a maximum elevation of about 55 m above sea level. It is crossed by the Bian, Kumbe, and Maro rivers which generally, with the exception of the Bian River, have narrow floodplains. The northern part of the plateau consists of watershed areas on slightly undulating terrain. The low permeability of the soils ensures seasonal inundation and the vegetation consists of Melaleuca savanna or low sedge grassland. The undulating ground with better-drained soils is generally not inundated and gives rise to tall mixed savanna and monsoon forests. There are local variations of vegetation in transitional areas, depending on inundation and soils with various mixtures of monsoon scrub and low mixed savannas. The vegetation has a long history of manipulation by humans through traditional subsistence activities, notably through the use of fire in grasslands and savanna, and through small-scale and local shifting cultivation. Although local drainage is the main factor that determines the type of vegetation, this annual burning often obscures the true relationship between habitat and vegetation. The savannas and grasslands of southern New Guinea have not been well studied. They are best known from various studies undertaken in Papua New Guinea (Brass, 1938; Rand and Brass 1940; Paijmans 1971) and in Wasur National Park by WWF (Bowe 1993, 1997; Stronach 1995, 2000, 2001; Bartolo et al. 2002). Van Royen surveyed the vegetation around the Maro, Kumbe, and Bian rivers in 1954 (van Royen 1963). Some limited work has been carried out on Yos Sudarso (Kimaam) (PHPA-AWB/Interwader 1990). The following description of grassland and savanna types is based on Paijmans et al. (1971) and Paijmans (1971).

Grassland and Savanna Types

low swamp grassland Low swamp grassland consists of dense mats of Pseudoraphis spinescens forming low swards in the dry season, but growing taller to maintain floating parts as wetseason floodwaters deepen. Slight differences in ground level favor the development of low swards of sedges such as Fimbristylis and Rhynchospora. Slight elevations such as levees are the habitat of Melaleuca viridiflora and M. cajeputi of small stature, while Dillenia alata, Barringtonia tetraptera, and Nauclea orientalis, common along the borders of the grass plains where they often form a line demarcating very slightly elevated ground. These grasslands are subject to prolonged inundation and occur close to the border with PNG in relict river channels such as the Tarl River and the edges of the Yauram area in the south-central part of Wasur National Park. This type of vegetation is becoming scarcer with large-scale encroachment of grasslands by Melaleuca (see section below on savanna and grassland dynamics).

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low to mid-height sedge-grass vegetation Low to mid-height sedge-grass vegetation consists of tussocky sedges and grasses of about 30–75 cm high in about equal proportions. The cover varies between from about 40–100%, and patches of bare ground are often visible. Dominant grasses are Ischaemum sp. on the coast and Eriachne further inland. Schoenus and Rhynchospora are common species of sedge. Herbs include Utricularia sp. In some localities, such as the plains behind Tomerau village in Wasur National Park and the fringes of the Bian River, the only tree is Pandanus sp., often in sufficient numbers to resemble savanna. Elsewhere scattered Melaleuca spp., Tristania, Nauclea orientalis, and Acacia sp. are found. Occasionally clumps of low trees and shrubs, especially Melaleuca spp. and Synoga lysicephala are found. The plains in the southeast corner of Wasur National Park are the habitat for this type of vegetation. They are subject to frequent burning, heavy grazing by introduced Rusa Deer (Cervus timorensis), and prolonged inundation. As with low swamp grassland above, this vegetation type is being colonized rapidly by Melaleuca spp., which is developing through woodland to forest with a progressively declining herb layer. The reasons for this change are not completely understood (see section below on savanna and grassland dynamics).

mid-height swamp grassland Mid-height swamp grassland occupies the edges of river banks and permanent swamps and forms dense floating mats. In areas of shallow gradient to dry land they can form extensive areas such as the fringes of Rawa Biru in Wasur National Park. The grass stands frequently include Echinochloa stagnina especially in deeper water with emergent parts up to 60–80 cm above the water level, Oryza sp., Leersia hexandra, and Hymenachne acutigluma. These latter can locally form pure stands. In some areas extensive mats of Hanguana malayana occur and in some places the pink lotus Nelumbo nucifera is abundant. In shallower swamps that may dry out, Ischaemum polystachyum is conspicuous, often together with Leersia hexandra, Eleocharis dulcis, and Phragmites karka. Areas of floating grass mats were once more extensive than at present, but have been reduced by the impact of grazing by introduced deer. Where deer grazing is heavy, the mats only survive in water that is too deep to allow deer to stand.

mid-height grassland In the coastal back plain, in the swales behind relict beach ridges, communities of mid-height grassland occur, consisting mainly of Imperata cylindrica up to 1 m tall. Due to frequent hot fires these communities tend to have only a few shrubs or trees that are moderately fire resistant, such as Melaleuca cajeputi, Alstonia actinophylla, and Livistona and Corypha elata palms. Where parts of the swales are slightly inundated and too wet for Imperata to grow densely, small herbs such as Crinum lilies often occur, which flower in the early wet season. These Imperata grasslands occur along the entire beach ridge system from the PNG border up to the Okaba area just west of the Bian River. The mid-height grasslands are main-

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tained anthropogenically by regular burning, in previous years largely for pig hunting. Where deer grazing of I. cylindrica regrowth is heavy, for instance after fire, the stands are replaced by a low sward of Chrysopogon aciculatus.

tall sedge-grass swamp Tall sedge-grass occurs on permanent, peaty swamps with a water depth of around 60 cm are dominated over large areas by single species or mixed stands of Phragmites karka and broad-leaved, robust sedges up to 2.5 m tall such as Scleria cf. poaeformis and Cyperus cf. platystylis. In some stands grasses and other sedges, such as Phragmites karka, Eleocharis spp., and Leersia hexandra are co-dominant. Scattered low Melaleuca and Acacia grow on elevations. Shallower margins are lined with low sedges and grasses and aquatic herbs such as Utricularia, often forming a distinct border and a fern-leaf pattern that is conspicuous on aerial photos. Open water is often covered by floating Azolla imbricata and Nymphaea indica. Where deer grazing is heavy, and especially in conjunction with burning after periodic drying, this vegetation type has disappeared and forms only relict stands, being replaced in many cases by Pseudoraphis spinescens grassland. Its decline is also accelerated by the drying out of the swamps.

melaleuca swamp savanna Melaleuca swamp savanna consists of seasonally inundated stands of Melaleuca cajeputi and M. leucodendron 10–15 m high, with a ground layer of mid-height to tall swamp grasses and sedges such as Phragmites, Leersia, or Elaeocharis. The tree canopy is open to moderately dense (20–60%). Sparse associated trees such as Barringtonia tetraptera and Nauclea orientalis, together with the composition of the herb layer, indicate that this vegetation type may result from encroachment of swamp grassland by Melaleuca. As with all permanently or seasonally dry sedge and grass communities, this is subject to periodic burning to which the trees are moderately resistant.

melaleuca savanna In Melaleuca savanna, Melaleuca viridiflora trees 8–14 m (up to 20 m) high form a moderately dense to open canopy over a herb layer of grasses and sedges up to 40 cm and is seasonally inundated to a lesser extent than is Melaleuca swamp savanna. The ground cover varies from 50–90% inversely with tree cover. Other tree species include Asteromyrtus symphocarpa (locally co-dominant), Acacia leptocarpa, and Dillenia alata, the latter often associated with eroding termitaria. Shrubs and small trees form occasional dense stands and include Pandanus sp. and Sinoga lysicephala. Herbs are moderately diverse and common grasses are Arundinella nepalensis, Eriachne triseta, and Ischaemum barbatum, and Schoenus spp. sedges are conspicuous.

low mixed savanna Low mixed savanna consists of trees with an average height of 13 m, some reaching 25 m, which form an irregular mosaic of open to moderately dense stands, the

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distribution of which reflect a combination of soil conditions and fire history. Seasonal shallow inundation or waterlogging is normal. Tree species are not diverse but always present or important are Tristania suaveolens, Asteromyrtus symphocarpa, Xanthostemon crenulatus, Alstonia scholaris, Grevillea glauca, and Banksia dentata. Frequent associates are Acacia spp., Dillenia alata, and Eucalyptus polycarpa. Epiphytic orchids and mistletoes are conspicuous. The shrub layer of up to 3.5 m high consists of scattered individuals and stands, the distribution and survival of which are determined by dry-season fires in the ground layer. Acacia leptocarpa is characteristic as are young individuals of the main tree species, often including Melastoma polyanthum, Rhodamnia, Acacia simsii, Sinoga lysicephala, and low Pandanus sp. The ground cover is general (80–100%), the distribution and composition being patchy and reflecting the variable and often poor soil drainage. As a consequence sedges are always common, including Schoenus sp. and Scleria (often in large, tall stands), and locally herbs include the insectivorous Drosera with Utricularia, Nepenthes, and Dianella. Grasses include poorly developed Imperata cylindrica, Eragrostis sp., Ischaemum barbatum, and Eriachne sp.

tall mixed savanna Trees in tall mixed savanna average 25–30 m in height, some reaching 40 m. As with low mixed savanna, canopy cover and height distribution are patchy. While seasonal inundation is unusual, the tendency to water logging depends on local soil conditions. The ground layer is dense, consisting of grasses up to 1.2 m (Imperata cylindrica mixed with Pseudopogonatherum irritans or Arundinella sp.) and sedges are occasionally common. The diversity of herbs is generally greater than that in low mixed savanna, although there are many species in common, though with few insectivorous species. Usually the dominant trees are Melaleuca cajeputi and Tristania suaveolens, often associated with Xanthostemon crenulatus, Acacia mangium, Alstonia scholaris, Melaleuca leucodendron, and Asteromyrtus symphocarpa. The shrub layer has a similar species composition to that in low mixed savanna.

Savanna and Grassland Dynamics The grasslands of southern New Guinea are at the center of an alarmingly rapid landscape change. Woody vegetation encroachment is clearly evident when comparing historical aerial photography from the 1940s to recent satellite imagery. The dominant woody species invading the grasslands are Melaleuca spp. and Nauclea orientalis. A significant reduction in the swamp reed Phragmites karka and other robust swamp grasses and sedges has also been observed (Bartolo et al. 2002). The primary disturbance agent in the wetlands is believed to be Rusa Deer (Cervus timorensis), which due to their intense grazing in the wetlands, has decimated some species of grasses. Where deer are or have in the past been abundant, the tall Phragmites and other robust grass and sedge swamps have largely disappeared. A combination of feral animal impacts and fire in conjunction with the drying

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out of swamp habitats is leading to a demise of grassland in the region, which is largely present in the protected areas of Wasur National Park in Papua and the adjacent Tonda Wildlife Management Area in PNG. These changes will ultimately result in a reduction in the region’s biodiversity. Factors that may also be contributing to a drop in the water table of the wetlands include changes in swamp hydrology due to the removal of grasses, an apparent reduction in wet season rainfall, and, locally, unsustainable water extraction from Rawa Biru swamp for use by the inhabitants of Merauke township. The Rusa Deer is not indigenous to Papua; its natural range encompasses much of the Indonesian archipelago to the west. It was introduced to Papua by the Dutch at Merauke in 1928, from where it has spread to most of the southern coastal lowlands of Papua and neighboring Papua New Guinea. Within Papua further introductions to Manokwari and Jayapura have given Rusa access to most of the potentially suitable habitat in the province. The deer were well adapted to their new environment in the grasslands and savannas of southern New Guinea. Not only was there an abundance of palatable grasses, but the swamp grasses were not heavily grazed by wallabies, the only large terrestrial grazing mammals indigenous to the region. The deer can graze out into swamps even in moderate depths of water, giving them access to the abundant swamp grasses. Conversely, wallaby populations are constrained by the availability of dry land when water levels are high. With the availability of so much food the deer bred successfully. There were few natural predators capable of killing deer and those few were unable to control their numbers. Initially the indigenous human inhabitants were not familiar with the deer and did not consider them to be an important food source. Under these circumstances the deer population grew rapidly and is still expanding its range. The problems that arise from the introduction of alien animals to new environments are well documented. Australasia, of which Papua is biogeographically a part, has a long and disastrous history of such introductions. It is now acknowledged that introduced herbivores have adversely affected the ecology of large parts of Australia. Introductions of several species of deer to New Zealand have also had regrettable ecological consequences. Within an ecological and biodiversity conservation context, the introduction of deer to Papua must be viewed similarly. According to the indigenous communities of Wasur National Park, the vegetation of the park has changed greatly since the deer were introduced. Three major habitat changes have been reported by the local people in Wasur, all of which appear to have taken place since the arrival of the deer. The main change has been a reduction in the tall swamp grasses, especially Hymenachne, which were once a major feature of the area. These formed a mat that floated on the water, reducing the extent of seasonal drying of the swamps and providing essential habitat for many wetland species. However, most species of swamp grasses were not resistant to heavy grazing by Rusa Deer. Consequently, the vegetation was greatly altered. The swamps are now considerably more seasonal in nature than formerly, reducing the habitat of crocodiles and nesting waterfowl. According to the local people, breeding of such

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species as the Australian Pelican (Pelecanus conspicillatus) has ceased and that of the Magpie Goose (Anseranas semipalmata) has declined in the park. A second major change has been the reduction of formerly abundant stands of Phragmites sp. This reed species is vulnerable to overgrazing by deer, which are thought to be responsible for the reduced extent and vigor of the stands. The reed beds are also an important refuge for the deer and other animals. In the 1980s and early 1990s the deer were commercially hunted in Wasur and the reed beds were regularly burned by the hunters to ease motorcycle access and prevent the deer from taking refuge. The resulting green-flush of new growth following burning quickly attracted the deer and also wallabies and this is likely to have further accelerated its decline. The presence of dense stands of Phragmites over much of the area would have slowed down the flow of water in the wet season considerably. The area would most probably have remained flooded for much longer, and the reduced cover of Phragmites today may explain why the swamps are neither as permanent nor extensive as they once were. The truth is that almost nothing is known in detail about the effects the deer have had on the flora and fauna, although we know that these effects must be substantial. Some of the deer-induced vegetation changes outlined above could be considered short-term and might be reversed if the deer population were to be reduced sufficiently (total removal of deer is likely not feasible, even if it were required). A third, more fundamental vegetation change in Wasur, which is likely to be due at least in part to the effects of grazing by deer, concerns the spread of trees (mostly species of Melaleuca) onto the open grasslands. The indigenous inhabitants of the area report that considerable areas of former grassland are now covered by Melaleuca forest or woodland. This process is continuing, as demonstrated by the large area of Melaleuca forest that has grown up within the past two or three decades. It is likely that the vegetation changes caused by the deer have provided Melaleuca with ideal conditions for regeneration. Recent work to quantify the extent and timing of the vegetation change has demonstrated the swift nature of the changes. The most alarming examples of encroachment of woody vegetation into open grassland habitat are occurring in both Wasur and Tonda. For example, at Umbal in southeastern Wasur, it appears that approximately 365 ha of Melaleuca viridiflora seedlings have established themselves (Bartolo et al. 2002). Of particular interest is that the density of M. viridiflora varies across the grassland in accordance with inundation regime. Where the water depth is greatest, the density of M. viridiflora seedlings is not as high when compared with areas where the water depth is less. Therefore, water depth and duration of inundation may be a determinant in the encroachment of Melaleuca into grasslands. There is also extensive localized Melaleuca encroachment on the boundary between the Melaleuca forest and open swamp in the region to the southeast of Yauram (Wasur National Park). The recent encroachment is estimated to be 150 hectares. The stages of encroachment are quite pronounced, evidenced by distinct age classes. In places there are stands of previously even-aged Melaleuca that were subject to fire in the late 1980s that have not regenerated. These patches should be

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examined closely as they may have been subjected to an appropriate fire regime to control encroachment. It is currently unclear whether this form of encroachment is a natural cyclical change. Analysis of historical aerial photography may aid in determining whether this boundary change reflects an episodic cyclical pulse or gradual directional change. We are currently using remotely-sensed data to quantify the conversion of grasslands to Melaleuca forest. Additionally, we are using different satellite-based data to examine the structure and density of the Melaleuca forests. Remote sensing provides a practical means of quantifying the temporal and spatial characteristics of Melaleuca encroachment, as the Trans-Fly region is remote and access to field sites is expensive and logistically difficult. The data that are being examined include black and white aerial photography acquired by the Dutch between 1944 and 1948. We are comparing these photographs to Landsat TM 5 imagery that was captured on 31 May 1997 that contains TM bands 2, 3, 4, and 5 (green, red, near-infrared, and mid-infrared, respectively). In Mblatar, a site within Wasur National Park, there has been extensive Melaleuca encroachment. In Figure 5.12.1a, the black-and-white aerial photograph captured in April 1948 shows a relatively open grassland environment characterized by a gray, smooth tone on the image. In Figure 5.12.1b, the 1997 satellite image, the grassland is characterized by pink tones and the Melaleuca encroachment is indicated by bright green tones. By comparing the two images it is clearly evident that there has been an extensive increase (approximately 1,450 hectares) in forest and woodland types at the expense of grasslands since 1948 (Bartolo et al. 2002). It is estimated that over 50% of the original extent of the grasslands has been lost or in the process of conversion to woodland. In addition to the effects of deer grazing described above, other factors may be contributing to the changes. Pigs also cause substantial damage to seasonal Eleocharis swamps. Field observations show large areas of ground dug over by pigs (Sus scrofa  elebensis; Stronach 2000). The bare damp soil resulting from these excavations provides ideal conditions for the germination of Melaleuca seedlings. The role of fire in the encroachment process is unclear. Late dry season fires that are typically hot have resulted in the death of some stands of Melaleuca. However, in most instances regeneration occurs in these stands. There is an element of chance in Melaleuca regeneration, in that if a late dry season fire burns a dry area (either grassland or forest) and the wet season rains are late or poor, then Melaleuca seedlings are likely to reach a size that can withstand subsequent flooding or fires. However, if a late dry season occurs followed immediately by a wet season with high rainfall, then the Melaleuca seedlings are likely to drown. Microtopography also is an important determinant of wetland vegetation. Small variations in topography, where the ground is slightly elevated, can result in Melaleuca seedlings surviving to maturity. The encroachment of Melaleuca onto grasslands is an extremely complex series of processes, and one that we are not close to understanding in all its aspects. The encroachment may even be part of a slow natural process that began many thou-

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A

B Figure 5.12.1. Change in grassland area in Mblatar area in Wasur National Park. Figure A is a black and white aerial photograph of the area in April, 1948. The grassland is this image is represented by the gray tone with a smooth texture. Figure B is a Landsat TM image from May, 1997. The remaining grassland area is labeled Mblatar and shown by a mid-gray tone.

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sands of years ago. It is clear that these changes in grassland-savanna dynamics have accelerated since the introduction of deer and key habitats are now threatened as a result.

Literature Cited Bartolo, R.E., M. Bowe, N. Stronach, and G.J.E. Hill. 2002. Landscape change and the threat to wetland biodiversity in Wasur National Park, West Papua (Irian Jaya). Pp. 11–19 in Ali, A., C.S.M. Rawi, M. Mansor, R. Nakamura, S. Ramakrishna, and T. Mundkur (eds.) Proceedings of the Asian Wetlands Symposium, 2001—Bringing partnerships into good wetland practices, 27–30 August 2001, Penang, Malaysia. Bowe, M. 1997. Turning a threat into an asset: an income generating scheme for community development and exotic species control in Wasur National Park, Irian Jaya, Indonesia. Case Study No. 8 in Claridge, G., and B. O’Callaghan (eds.) Community Involvement in Wetland Management: Lessons from the Field. Wetlands International, Kuala Lumpur. Bowe, M., B.T. Hariadi, and E. Futunanembun. 1993. Preliminary report on the Melaleuca scrub encroachment of the grasslands of Wasur National Park. WWF Project No. ID 0105. Brass, L.J. 1938. Botanical results of the Archbold Expeditions. IX. Notes on the vegetation of the Fly and Wassi Kussa Rivers, British New Guinea. J. Arnold Arbor. 19: 175–190. Paijmans, K. 1971. Vegetation, forest resources, and ecology of the Morehead-Kiunga area. Pp. 88–113 in CSIRO Land Resources of the Morehead-Kiunga Area, Territory of Papua and New Guinea. Land Research Series No 29: Commonwealth Scientific and Industrial Research Organisation, Melbourne. Paijmans, K., D.H. Blake, and P. Bleeker.1971. Summary description of the MoreheadKiunga Area. Pp. 12–17 in CSIRO Land Resources of the Morehead-Kiunga Area, Territory of Papua and New Guinea. Land Research Series No 29: Commonwealth Scientific and Industrial Research Organisation, Melbourne. Rand, A.L., and L.J. Brass. 1940. Results of the Archbold Expeditions. No. 29. Summary of the 1936–37 New Guinea Expedition. Bull. Am. Mus. Nat. Hist. 77: 341–380. Silvius, M. J., and A.W. Taufik. 1990. Conservation and Landuse of Pulau Kimaam, Irian Jaya. PHPA-AWB/Interwader report. Stronach, N., 1995. The Rusa Deer of Wasur National Park, Irian Jaya. Mimeo, WWF Project ID 0105. Stronach, N. 1997. Identification of the threats to biodiversity; Rusa deer as a threat; Outline of fire management practices; Vegetation changes and hydrology. In Kitchener, D. (ed.) Wasur National Park-Tonda Wildlife Management Area. Biodiversity Research Planning Workshop. WWF Indonesia Programme, Jakarta. Stronach, N. 2000. Fire in the TransFly Savanna, Irian Jaya/PNG. Pp. 90–94 in RussellSmith, J., G. Hill, S. Djoeroemana, and B. Myers (eds.) Fire and Sustainable Agriculture and Forestry Development in Eastern Indonesia and Northern Australia. Proceedings of an international workshop held at Northern Territory University, Darwin, Australia, 13–15 April 1999. ACIAR, Canberra. Stronach, N. 2001. Draft framework for research on vegetation change in the TransFly. WWF report. van Royen, P., 1963. Sertulum Papuanum 7, Notes on the vegetation in South New Guinea. In Nova Guinea—Contributions to the anthropology, botany, geology and zoology of the Papuan region Number 13. (Continuation of: Nova Guinea, new series, Vol. 10, 1959.)

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5.13. Caves of Papua louis deharveng, tony whitten, and philippe leclerc u r fa c e a nd s o i l e c o s ys t e m s are complex because they generally involve a lot of interacting species with important yearly and daily variations and strong connections with neighboring ecosystems. This makes their biodiversity difficult to inventory exhaustively, and their dynamics difficult to study. Cave ecosystems are much simpler. Their boundaries are discrete (Whitten, Soeriaatmadja, and Affif 1996), their species assemblages are not speciose, and their microhabitats much less diverse than those of outside ecosystems. They lack the primary producers and primary consumers that contribute the most to biodiversity in surface ecosystems (Gibert and Deharveng 2002). They do not exhibit marked seasonal or daily climatic changes. For these reasons, caves provide unique opportunities for testing basic hypotheses related to biodiversity dynamics and community ecology, and permit consideration of all the species of the assemblages under study. Caves are also natural windows into a huge underground environment, which would be otherwise largely inaccessible to humans. Caves are extremely diverse in morphology. They may be small or very large, they may develop long horizontal passages, deep vertical shafts or mazes, rivers may flow through them (i.e., active caves), or they may completely lack water (i.e., non-active or fossil caves). Except for the maze form, all other cave morphologies enumerated here have been reported from Papua, and wildlife was present in all caves that have been examined. In addition to natural cave passages, artificial galleries made by humans for various purposes also provide habitats suitable for subterranean life. Due to the stability and harshness of their environment, caves are places where evolution proceeds at a very different pace from most other ecosystems. The few animals that have adapted to subterranean conditions exhibit unique morphological and biological features, including loss of organs essential to surface species (eyes, pigment, wings for insects) and elongation of appendages (Christiansen 2004), loss of water regulation capacity, reduced metabolism, much longer life cycles, much lower reproductive rates, and frequent tendency to neoteny (the maintenance of larval characteristics into adulthood). Many of these traits tend to limit population dispersal between karst patches, which is reflected by extremely high level of endemism in subterranean fauna (Gibert and Deharveng 2002). In this context, Papua is a priori of exceptional interest because of the extent, the fragmentation, and the altitudinal range of its karsts, unmatched in any other region of Southeast Asia or Australasia. However, data are insufficient to derive

S

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

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any sound statement about the richness of the Papuan cave fauna. What is presented below is the first biospeleological research in Papua.

The Cave Environment

cave and speleothem formation Caves exist in volcanic rocks or in sandstone, but most are developed in limestone. They are generated by the slow dissolution of carbonate rocks by water. Rainwater dissolves carbon dioxide while percolating through soil, increasing its acidity. It reaches the bedrock with the ability to dissolve calcium carbonate (the main constituent of limestone), and progressively enlarges the small cracks, which ultimately may reach human-size passages that we call caves. Arriving in large voids, water suddenly looses its CO2 because of changes in atmospheric gas equilibrium, and precipitates its calcium carbonate, producing the cave-specific formations that we call stalactites (as well as a variety of other hanging formations). Remaining calcium carbonate in water dripping from stalactites precipitates at floor level, forming stalagmites and other formations. Any running or standing water in a cave may also deposit its calcium carbonate in a variety of crystallized forms. All of these formations, called speleothems, increase the surface area available to cave inhabitants.

karst hydrology Karst hydrology controls much of the community structure of aquatic, and less directly of terrestrial, cave organisms. The epikarst (upper part of the percolation zone just below the surface; Humphreys 2000) is more permeable and porous than the underlying massive bedrock, as microclimatic fluctuations, tree roots, and karst processes tend to fracture and disaggregate surface and subsurface rocks. An aquifer (a perched saturated zone) is often developed within this epikarstic zone, storing some of the infiltrated water, and hosting a peculiar interstitial fauna (Jones, Culver, and Herman 2004). Underneath, water circulates through very small pores that are evenly distributed in the rock or in fractures (10 ␮m to 10 mm) where flow is mostly laminar, and in larger conduits (the largest ones called caves) where flow is mainly turbulent. This last kind of waterway, used but also generated by karst waters, is specific to limestone, and allows rapid water transfer, while flow is slower in fractures and very slow in micropores (Bakalowicz 2005; White 2005). The most important feature determining the composition of aquatic cave fauna is the origin of the water available for life. Water that reaches the deep part of the karst by percolating through soil and bedrock in pores or fractures, as well as cave streamlets formed from percolating waters, often harbor specialized animals (called stygobites) restricted to these habitats. Underground streams that flow directly from large surface rivers, and cross the karst without passing through any filter (e.g., large boulders) often host only outside fauna, even kilometers into the cave. The lack of suitable waters in the caves that have been studied so far may

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explain the lack of observations of stygobitic fauna in Papua. However we assume that when exploration of additional Papuan caves is conducted such fauna will be found in caves that contain suitable water sources.

cave environments Cave environments contain a unique combination of features that are both favorable and unfavorable to living organisms. Favorable conditions include the very strong buffering of daily and seasonal fluctuations of temperature and, for terrestrial fauna, of humidity, and the fact that air movements are usually extremely reduced, at least in deep parts of the caves. Unfavorable conditions are absence of light and scarcity of food. A few animals called troglobites (terrestrial) or stygobites (aquatic) have adapted specially to these characteristics, and are now unable to survive in conditions found outside caves.

papuan caves Limestone covers large sections of Papua. Karst is found in extremely diverse ecological contexts, from sea level to the highest peaks of Indonesia above 4,500 m. In spite of this, speleological exploration of Papua is the least advanced of any sector in the Australasian region. While many big caves and shafts are known in several areas of Papua New Guinea, only a handful of caves have been explored in Indonesian Papua. Highland caves around Wamena and Mount Trikora have been visited by British cavers (White 1986; Willis 1990; Jones 1993), who explored more than two kilometers into the giant sink of the Baliem, one of the largest underground rivers in the world. A Dutch team surveyed several vertical systems in the Star Mountains (Severens et al. 1993). French teams explored large vertical shafts and caves in the Vogelkop, but results of these expeditions were never published. The Fakfak region was also visited by French teams (1992 expedition, reported in Lacas, Leclerc, and Mary 2001; further expeditions in 1994, 1999, 2001, 2002, without published report). Papuan caves are shown in Figures 5.13.1–3, and available information is summarized in Table 5.13.1.

Ecology Our knowledge of the cave ecology of Papua is extremely limited. However, some preliminary insights may be gained by examining the species distributions in the surveyed Papuan caves in a broader geographical and ecological context.

ecological classification of cave animals Cave animals are classically divided into three ecological groups, based on the strength of their ties to underground habitats (Humphreys 2000): troglobites, or obligate cave species, unable to survive outside subterranean environment; troglophiles, able to live and reproduce in caves but also found in similar dark, humid microhabitats outside caves; trogloxenes, species that either regularly enter caves for refuge but normally return to the outside environment to feed (e.g., bats and

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Figure 5.13.1. Karsts of Papua surveyed for caves. Open circles indicate no data on cave fauna; solid circles indicate that data on cave fauna are available. swiftlets), or accidentally wander into caves (sometimes referred as a fourth category, accidentals). These categories concern terrestrial species, while the terms stygobites, stygophiles, and stygoxenes are often used for aquatic species. Establishing the ecological status of cave invertebrates is, however, quite difficult, specially in tropics where outside fauna is often less known than cave fauna, and where many troglobites are non-troglomorphic (Deharveng and Mouret 1981). Troglomorphy is the best available indicator of troglobitism for Papua cave fauna. ‘‘Troglomorphic’’ qualifies a species supposed to be linked to cave life because it has characters that appear in many troglobitic and stygobitic species. Reduction of eyes and pigment, and the elongation of appendages (in Arthropods), are the most frequent troglomorphic features. With about six species recognized as troglobites and none as stygobites on this basis, the Papuan troglobitic fauna appears immensely underestimated and undersampled, especially in comparison to the surface diversity of the Papuan karst. In comparison, Papua New Guinea, which is one of the less-known cave fauna of Australasia, has at least 20 troglobites and 12 stygobites (Geoffroy 2001; Brehier, pers. comm.). Other ecological categories are also widely used to characterize species on other grounds than their link to cave conditions. The most important are anchialine stygobites, interstitial stygobites, and guanobites.

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Figure 5.13.2. The deepest shafts of the Vogelkop: Lomes Iono Besar (left) and Lomes Longmot (right). Source: Couturaud (1993).

Anchialine stygobites live in anchialine caves, where saltwater and freshwater mix. None has been reported from Papua or from Papua New Guinea, but they have never been searched for. Given that anchialine stygobites are well diversified in all littoral karsts of the tropics where they have been looked for, a rich fauna can be expected in Papua as well where littoral karsts are developed, such as Fakfak, Kokas, and Kaimana. Interstitial stygobites live in the phreatic zone (where voids in the rock or sediments are small and completely filled with water) and are rarely found in caves. They are more diversified than cave fauna where they have been investigated. They have never been sampled in Papua or Papua New Guinea. Guanobites, terrestrial animals adapted to feed on bat or swiftlet guano, or to

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Figure 5.13.3. Map of Nomonkendik Cave, one of the largest mapped caves of Papua. Source: Lacas et al. (2001).

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Wamena region Wikuda Cave* Danda* Kwalinga Cave* Wur Cave Baliem River Cave  Tinggina Cave*

Star Mountains Sibil Buk I* Adbon Buk*

Vogelkop Lomes Iono Besar* Lomes Longmot*

Region/Cave

1,717 1,100 603 ? 2,643

2,313 987

380 380

Length (m)

Table 5.13.1. Caves of Papua

0 20 229 ? 142

349 113

315 360

Depth (m)

none none none Rigal (unpubl.) none

none none

none none

Biological data

Willis (1990) White (1986) Willis (1990) none Checkley (1993)

Severens et al. (1993) Severens et al. (1993)

Couturaud (1993) Couturaud (1993)

Speleological data

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0

⬃250 ⬎ 500 ⬃70 ⬃70

Biak Japanese cave Mansapur Cave Yewnus Cave

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Note: * indicates caves that have been mapped, at least in part.

? 4 0

15

?

1

55 0 5 47

⬎300

⬃180

990

535 70 275 3,300

Misool Tulang-tulang Cave Maguei Cave Mapeingan Cave

Kaimana Batu Lubang Cave

Yahyah Cave Lubang Cave

Fakfak Peninsula Lubang Kayu Mera* Genuni Cave Giragandak Cave* Nomonkendik Cave*

none Leclerc (unpubl.) Leclerc (unpubl.)

Leclerc (unpubl.) Leclerc (unpubl.) Leclerc (unpubl.)

none

Lacas, Leclerc, and Mary (2001) Lacas, Leclerc, and Mary (2001) Lacas, Leclerc, and Mary (2001) Lacas, Leclerc, and Mary (2001); Leclerc (unpubl.) Leclerc (unpubl.) Leclerc (unpubl.)

(Tourist cave) Leclerc (unpubl.) Leclerc (unpubl.)

Leclerc (unpubl.) none Leclerc (unpubl.)

J. P. Mary (pers. comm.)

Lacas, Leclerc, and Mary (2001); Leclerc (unpubl.) Lacas, Leclerc, and Mary (2001) Lacas, Leclerc and Mary (2001) Lacas, Leclerc, and Mary (2001); J. P. Mary (pers. comm.) Leclerc (unpubl.) none

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prey on guano dwellers, may be obligate or non-obligate cave dwellers. They are linked to guano habitats more than to caves, and usually do not exhibit troglomorphic features. Many of the Papuan species characterized as troglophiles are probably guanobites.

food resources and food webs Absence of light, and hence of photosynthesis and primary production, is a universal character of subterranean ecosystems. It strongly constrains the functioning of living underground communities. Chemoautotrophic bacteria deriving their energy from inorganic compounds can reduce inorganic carbon and produce organic matter as a basic food resource for cave communities (Sarbu 2000). However, such a process is exceptional, and food supplies usually come from surface ecosystems. The pathways of organic matter input into caves include roots in shallow caves up to 30 meters deep (Jasinska and Knott 2000), various kinds of organic matter carried by water flow, flood debris along river streams, fine or dissolved organic matter in waters percolating from the surface, and many kinds of organic debris abandoned or brought in by humans. But in the tropics, the major sources of food are the bats and swiftlets, which roost and breed in caves but feed outside (Whitten, Soeriaatmadja, and Affif 1996). This food supply is mainly provided by their feces, called guano, where fungi, bacteria, guanobites, parasites of flying vertebrates and their predators proliferate. Molted hair, feathers, shed pieces of skin, and corpses may also be contained in guano, and may provide significant additional resources when bat or swiftlet colonies are large. The numerous parasites of flying vertebrates, often found on the guano, are preyed upon by cave predators, most of which are generalists. In Southeast Asia and probably Papua as well, large cave crickets feed on guano or flood debris, and act as the main disperser of food resources in terrestrial habitats. They are often abundant and wander far from guano towards cave walls and oligotrophic zones of the caves, where they leave isolated feces exploited as food resources by troglobites. Cave fauna is distributed according to the density rather than nature of food resources, with obligate subterranean species in the lowest resource areas and guanobites and troglophilic species in the high resource areas. Among these last two categories, many species remain on or very near the guano (floor community), while others, especially giant Arthropods, including adult crickets and their predators, tend to stay on cave walls, occasionally foraging near guano piles. Species interactions are still very poorly known, though it is generally assumed that decomposers as well as predators in caves are food generalists. The food web shown in Figure 5.13.4 provides an overview of our present knowledge. One of the most intriguing observations, which holds for Papua in spite of the low number of cave species recorded, is that diversity of predators (arachnids) is often substantially higher than diversity of potential prey among obligate cave species or regular troglophiles. Possibly the diet of troglobitic predators includes a large proportion

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Figure 5.13.4. Simplified cave food web. The groups reported from Papua caves are indicated in bold. Source: Modified from Deharveng (2004).

of non-troglobitic species, which are known to penetrate deeply into tropical caves.

cave fauna of papua Very little has been published on the subterranean fauna of Papua, where most karst areas have never been surveyed. The only significant information concerns the Fakfak caves (Lacas, Leclerc, and Mary 2001). Sporadic records exist for a few other regions (Figure 5.13.1). For each zoological group reported from Papua, we provide brief comments about cave species of Papua New Guinea, often based on the compilation of Geoffroy (2001). Though also poorly known, Papua New Guinea has been much better explored biologically than Papua. In particular, two big speleological expeditions including cave biologists (Chapman 1976; Smith 1980) collected a lot of fauna in the highland caves of Papua New Guinea, close to the border with Papua.

Invertebrates Gastropoda An aquatic species, probably not a stygobite, lives in Giragandak Cave and Nomonkendik Cave (Fakfak karst), while terrestrial snails occur in Mansapur Cave on Biak Island. In contrast, many families of aquatic snails are reported from Papua New Guinea, but no terrestrial ones.

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Hirudinea A pink leech, probably a bat parasite, is frequently found on cave walls of Giragandak Cave (Lacas, Leclerc, and Mary 2001). Similar species exist in Papua New Guinea (Selminum Tem: Chapman 1976; Mapos cave near Lae: Deharveng, unpubl.), and are found sporadically in caves of Sumatra (Deharveng and Bedos 2000) and Thailand (Lacas, Leclerc, and Mary 2001). Crustacea Oniscida (Terrestrial Woodlice) Woodlice have been observed in most caves, with a few exhibiting troglomorphic characters (Leclerc, unpubl.). Troglobitic woodlice are frequent and diversified in caves of Papua New Guinea. Crustacea (Aquatic) Crustacea are the most diverse group of aquatic cave animals in most ground waters of the world, but are very poorly known in Papua where no systematic sampling has ever been performed. While remarkable stygobitic isopods, shrimps, and crabs are known in Papua New Guinea, none have ever been cited from Papua. All Papuan species of aquatic cave Crustacea are troglophilic. They include two crabs and at least five shrimps, all but one of which have well developed eyes. Geelvinkia darnei is a crab described from a cave of the Lina massif near Manokwari. Nomonkendik Cave has another unidentified crab species, which also has large eyes. Among shrimps (L.B. Holtuis ident.), four are species widespread in tropical Asia or in Southeast Asia: Caridina typus, frequent in Lubang Kayu Mera near Fakfak; and Macrobrachium australe, M. lar, and Palaemon concinnus, known from caves in Papua (Hobbs 1998). Unidentified Palaeomonidae with large eyes (Leclerc, unpubl.) have also been observed in Yahyah Cave near Kokas, and in Mapeingan Cave on Misool. The Palaemonidae found in Nomonkendik and Giragandak caves (Lacas, Leclerc and Mary 2001) have slightly reduced eyes (Leclerc, unpubl.). In Nomonkendik Cave, some shrimps carried temnocephals (parasitic flatworms). Arachnida As is usual in the tropics, Arachnida are the most diversified group of terrestrial fauna in Papua, and the largest group of predators. Araneae are very common and highly diversified in Papuan caves, although estimates of their density and diversity are not yet available. In the Fakfak Peninsula caves species of Anapidae and Mysmenidae (Nomonkendik), Tetragnathidae and Uloboridae (Lubang Kayu Merah), and Nesticidae (Giragandak) were observed. Some are probably troglobites (Leclerc, unpubl.). Non-troglomorphic mygalomorph spiders are abundant in the fossil galleries of Nomonkendik Cave (Fakfak Peninsula). In some places, the entire surface (more than 100 m2) of the huge clay deposits on the river banks are spotted with spider holes, with their occupants distributed regularly every four to five square meters. Given the large size of a single specimen and the scarcity of potential prey (large crickets; Leclerc, unpubl.), the biomass of these spiders is surprisingly high. Papua New Guinea also

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has a large diversity of spiders, including Uloboridae and Nesticidae. Heteropoda, the large huntsman spiders so frequent in all Southeast Asia caves, are cited from Papua New Guinea but not from Papua caves. Opilionids are rare. The species found in Giragandak Cave (Fakfak) is troglophilic, like those cited from Papua New Guinea. Amblypygi of the genus Charon exist in Papua (Weygoldt 2000). Unidentified species, often of large size, were commonly seen in most caves of the Fakfak karst (Lacas, Leclerc, and Mary 2001). They are also common dwellers of entrance and high energy habitats in lowlands of Papua New Guinea, but are absent in colder caves of the highlands (Deharveng and Mouret 1981). Non-troglomorphic Schizomids (Lacas, Leclerc, and Mary 2001) and falsescorpions (Leclerc, unpubl.) have been encountered under stones in two caves of the Fakfak karst (Genuni Cave and Lubang Kayu Merah). Present as well in Papua New Guinea (Deharveng and Mouret 1981), they are regular predators on guano Arthropods in Southeast Asia, but have not yet been taxonomically investigated. One specimen of Palpigrad has been found in Giragandak Cave, Fakfak Peninsula (Lacas, Leclerc, and Mary 2001). These tiny Arachnids are widespread in Southeast Asia, most of them soil- rather than cave-dwellers. They have generally been overlooked by biospeleologists, and are not cited from Papua New Guinean caves. Myriapoda Chilopoda and Symphyla were rarely observed in Papuan caves, and records should be considered as accidental, while Diplopoda were frequent. According to Geoffroy (2001), three families of Diplopoda are found in the caves of Papua New Guinea: Metopidiotrichidae (one species), Doratodesmidae (three species), and Paradoxosomatidae (five species). This last family contains the only true troglobitic diplopod of Papua New Guinea, Selminosoma chapmani (Hoffman 1978). Undescribed material from Papua (Golovatch, pers. comm.) includes this last family (one species from a cave in Biak) and four other families not reported from Papua New Guinea: Haplodesmidae (probably Cylindrodesmus hirsutus) and Pyrgodesmidae from Fakfak caves, Opisotretidae from a cave on Misool, Cambalopsidae (Hypocambala sp., Figure 5.13.5) from a cave on Biak. The ecological status of these different forms is uncertain, but the species of Misool has a troglomorphic morphology. Carinated Cambalopsidae, widespread in Southeast Asian caves, have not been found so far in Papua or New Guinea caves. Collembola As in temperate caves, Collembola are normally the most abundant terrestrial decomposers in tropical caves. Seven species are recorded here from Papuan caves, of which three or four are possibly troglobites. Fakfak Peninsula hosts three cave Entomobryidae, each of them having been found in two caves of the karst: a blind but not troglomorphic Coecobrya sp., a genus common in Southeast Asia caves with several guanobitic species, a blind and slightly troglomorphic Pseudosinella

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Figure 5.13.5. Hypocambala sp. from Yewnus Cave in Biak. closely related to P. maros recently described from southern Sulawesi caves (Deharveng and Suhardjono 2004), and an Acrocyrtus with reduced eyes, possibly troglobitic. Acrocyrtus are large epigean Collembola (more than 2 mm) frequent in Southeast Asia. The Fakfak cave species is the first in this genus to exhibit strong eye reduction. Near Wamena, the Wur Cave (Rigal, unpubl.) has given four Collembola species, including a Coecobrya sp. and a Pseudosinella sp. quite similar to those of Fakfak caves, a Thalassaphorura (Onychiuridae) of uncertain ecological status, and Folsomia candida, a parthenogenetic and opportunist Isotomid that is widespread worldwide, especially in caves. In contrast to these Papuan species, several Collembola from Papua New Guinea are extremely troglomorphic: Coecoloba plumleyi and Pseudosinella sp. from Selminum Tem near Telefomin (Deharveng 1981), and a newly discovered Coecobrya sp. from New Britain (Brehier, pers. comm.). No doubt that similarly evolved species will be discovered in Papua when additional exploration has been conducted. Insecta Very little is known about cave insects in Papua. Surprisingly, no cave cockroach was found in Papua, and none is cited from Papua New Guinea, while they are among the most regular inhabitants of caves in Southeast Asia and Australia. Unidentified Orthoptera are present from entrances to dark zones, at least in eutrophic caves like Giragandak Cave near Kokas (Leclerc, pers. obs.). The only observed beetles were small carabids near organic deposits of the Nomonkendik

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underground river near Kokas. No troglobitic beetle has been recorded, but they should exist there, as several troglomorphic species are known in Papua New Guinea (Geoffroy 2001). Diptera are common in many caves but none has been identified so far. Parasitic flies have been seen on several bats.

Vertebrates Fish In Mapeingan Cave (Misool), slowly moving, whitish, small-eyed fishes were observed on the bottom of a small stream; they did not react to light. On Biak, many blue holes at the foot of the limestone cliffs just above sea level are said to host white fishes (Leclerc, unpubl.). These forms remain undescribed. The blind fish Oxyeleotris caeca of Papua New Guinea is the only cave fish described from New Guinea. It belongs to Eleotritidae, and seems to be closely related to O. fimbriata, one of the most widely distributed gudgeons in New Guinea. Eleotritidae have also cave representatives in Australia (one species) and Madagascar (two species). Cave fish have never been looked for in Papua, but the above observations suggest that they probably exist there. Birds Swiftlets (Apodidae) are frequent in Papuan caves. Collocalia fuciphaga, the most lucrative species in Southeast Asia, whose nests are made almost exclusively of saliva and are considered a delicacy, is absent from Papua. But five other species of cave swiftlets are present (Chantler and Droessens 1995), all placed in the genus Collocalia by these authors. With bats, they are by far the best studied cave animal group in the region, though many distributional and taxonomic problems persist. The Glossy Swiftlet, C. esculenta, widely distributed in Southeast Asia, is differentiated in three subspecies in Papua: a lowland form (C. e. nitens), a highlands form above 1,600 m (C. e. erwini), and the form C. e. numforensis restricted to an island of Geelvink Bay. C. esculenta nests in shallow caves and on rock cliffs and overhangs. The Mountain Swiftlet, C. hirundinacea, is also present with three forms: the typical form in the mainland and a few islands; C. h. baru in Yapen and Geelvink (now Cenderawasih) Bay; and C. h. excelsa above 1,600 m in the Central Range. They nest in dark or twilight zones of shafts. The Three-toed Swiftlet, C. papuensis, and the Bare-legged Swiftlet, C. nuditarsus, are rare species that live in the mainland mountains. No data are available on nesting and behavior. The Uniform Swiftlet, C. vanikorensis, widespread in Southeast Asia, is represented by three forms in Papua: C. v. waigeuensis which inhabits several islands of northern Maluku, and is recorded from Misool; C. v. steini on Biak and Numfoor islands; and C. v. granti in Papua lowlands and a few islands. They nest in the total darkness of caves and shafts.

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Mammals Except the Ground Cuscus (Phalanger gymnotis) which often rests in caves during the day (Flannery 1995), the only mammals regularly present in caves are bats. Their guano, especially that of insectivorous species, is at the basis of subterranean food webs. Large frugivorous cave species are often shot for their meat by local people (Lacas, Leclerc, and Mary 2001; Couturaud 1993). Twenty-three bat species have been recorded in Papua, but more are likely to exist, as bat distribution is much less well documented in Papua than in Papua New Guinea. This is suggested by the fact that six species that are recorded in Papua New Guinea close to the border with Papua have not yet been recorded in Papua (Table 5.13.2).

Gaps and Prospects In comparison with flying vertebrates or with invertebrates of any other habitat except soil, cave invertebrates of Papua are extremely poorly known. Very few papers deal with this fauna, and most of the data used in the present chapter have not been published. The number of obligate subterranean species is lower in Papua than in any region of comparable area in Southeast Asia. None of the troglobitic species encountered has been described so far. This reflects wide gaps in our faunal, ecological, and geographical knowledge of Papuan subterranean fauna. Faunal gaps are illustrated by the absence of many lineages, such as several orders of aquatic Crustacea and several families of Diplopoda, which are expected to exist in Papua because they are found in Papua New Guinea and other surrounding regions. Ecological gaps are reflected in the extremely poor data about aquatic habitat diversity, and the total lack of information about anchialine and interstitial fauna, which are rich wherever they have been documented in the tropics. While it has been shown that high elevation caves host the largest number of highly troglomorphic species in the tropics (Chapman 1976; Smith 1980; Geoffroy 2001), none has been so far biologically surveyed in Papua. None of the studied areas, except the Fakfak Peninsula, has been significantly sampled, and data are completely lacking for karsts as large as the Star Mountains, the Vogelkop, or the Kaimana karst, among others. We are therefore at the very beginning of the study of the Papuan cave fauna, with a huge field of investigation offered by the diversity of karsts in the province, which range from sea level to over 4,500 m in altitude.

Disturbance and Conservation At present, no case involving disturbance detrimental to cave fauna has been documented in Papua, but disturbance and associated threats to cave life are growing with deforestation and mining. Among a variety of potential disturbances, two are

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insects insects insects

Emballonuridae (9 species) Emballonura beccarii E. furax E. raffrayana insects insects insects insects insects insects insects insects insects insects

fruit fruit fruit

Pteropodidae (20 species) Aproteles bulmerae Dobsonia magna Roussettus amplexicaudatus

Hipposideridae (12 species) Aselliscus tricuspidatus Hipposideros ater H. calcaratus H. cervinus H. corynophyllus H. diadema H. edwardshilli H. maggietaylorae H. papua H. wollastoni

Diet category

Group/Taxon

Table 5.13.2. Cave-roosting bats known from Papua

58–360 55–1,700 40–350 3–1,360 1,500–1,800 20–1,210 240 10–360 100–300 400–2,000

0–1,300 120–360 0–1,320

1,760–2,400 0–2,700 0–2,200

Elevation range (m a.s.l.)

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Endemic to New Guinea (continued)

Roosts in small groups, in midsection of caves Also roosts outside of caves Endemic to New Guinea Widespread in Southeast Asia; also roosts outside of caves Rare; endemic of Sandaun Province, PNG, near Papua border Also roosts outside of caves

Sometimes found in large numbers

Biak, Yapen, and near border (Telefomin) Endemic to New Guinea Roosts in shallow caves

Near border with Papua Sometimes extremely large colonies

Location

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insects

insects insects insects insects insects insects insects aquatic insects and small fish insects

insects insects insects

Diet category

Note: The number of species per family refers to New Guinea mainland. Source: After Flannery (1995).

Pipistrellus angulatus

Phoniscus papuensis

Vespertilionidae (21 species) Kerivoula muscina Miniopterus australis M. macrocneme M. magnater M. medius M. propitristis M. schreibersii Myotis adversus

Rhinolophidae (4 species) Rhinolophus arcuatus R. euryotis R. megaphyllus

Group/Taxon

Table 5.13.2. (Continued)

0–990

20–1,210

20–1,600 0–1,500 0–3,200 0–260 360–1,360 0–1,600 0–2,120 49–260

360–1,600 165–1,720 260–360

Elevation range (m a.s.l.)

Not recorded from mainland Papua, but present in cave in Biak; usually roosts in trees Not recorded from mainland Papua, but present in Biak; also roosts outside of caves

Large clusters; distributed from Europe to Australia Also roosts outside of caves

Near border

Near border

Telefomin area near border Deep in caves Ok Menga near border

Location

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of particular concern for cave life in Papua: the indirect effects of deforestation, and the more direct effects of mining and quarrying. Organic matter input via rainwater that percolates through forest soils is an essential food supply for invertebrate communities, especially where guanoderived resources are absent (often in the deepest parts of caves). Changes in forest soil structure and chemical composition will modify the nature and amount of this vital input, with impacts on living subterranean communities. Deforestation also causes leeching of the upper soil layers, which induces the filling of bedrock cracks and modifications of hydrological circulation. Impacts on subterranean habitats are particularly difficult to quantify, but are highly predictable. Forest disturbance also modifies outside resources for bats, and may result in the abandonment of caves where they had been roosting for decades. Mining affects large areas in Papua, particularly at Grasberg, one of the world’s largest gold and copper mines, but it is not known to what extent karst rocks are directly affected. Mining directly threatens subterranean fauna through quarrying, which sometimes results in cave destruction and often in bat disturbance, with negative impacts on the whole cave fauna. Because of the very scattered available information, the limited human resources available for wildlife protection, and the low consideration given to the conservation of invertebrates, priority should be placed on monitoring and protecting large bat colonies wherever possible. Obligate cave invertebrates can cope with food scarcity. They can move through the cracks of the limestone massif to escape food shortage or disturbance. They are rarely at risk, except in case of pollution or extensive limestone quarrying. Conversely, the loss of guano resources is likely to lead to the complete extinction of the rich assemblages of guanobitic invertebrate species, which, contrary to bats, cannot fly and escape disturbance because most are wingless. These guanobites are therefore a priori more threatened than any other category of subterranean fauna. Protection of bat colonies would also secure the food supply of many troglobites and stygobites, which are ultimately and largely also dependent on guano.

Literature Cited Bakalowicz, M. 2005. Epikarst. Pp. 220–223 in Culver, D.C., and W.B. White (eds.) Encyclopedia of Caves. Elsevier Academic Press, Amsterdam. Chantler, P., and G. Driessens. 2000. Swifts. Pica Press, Tonbridge, Kent. Chapman, P. 1976. Speleobiology. In The British New Guinea Speleological Expedition 1975. Trans. British Cave Research Assoc. 3: 192–203. Checkley, D. 1993. Cave of Thunder: the exploration of the Baliem River Cave, Irian Jaya, Indonesia. International Caver 6: 11–17. Christiansen, K. 2004. Adaptation: morphological (external). Pp. 7–9 in Gunn, J. (ed.) Encyclopedia of Caves and Karst Science. Fitzroy Dearborn, London. Couturaud, A. unpublished. Irian 93 ou Pe´re´grinations spe´le´ologiques en Irian Jaya (Indone´sie). Fe´de´ration Franc¸aise de Spe´le´ologie (unpublished report).

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1082 / louis d eharveng , t o n y w h i t t e n, & p hil i pp e lec l erc Deharveng, L. 1981. The fauna of caves (Papua New-Guinea). Spelunca suppl. 3: 38–39. Deharveng, L. 2004. Asia, Southeast: Biospeleology. Pp. 109–112 in Gunn, J. (ed.) Encyclopedia of Caves and Karst Science. Fitzroy Dearborn, London. Deharveng, L., and A. Bedos. 2000. The cave fauna of Southeast Asia: origin, evolution and ecology. Pp. 609–638 in Wilkens, H., D.C. Culver, and W. Humphreys (eds.) Ecosystems of the World 30: Subterranean Ecosystems. Elsevier, Amsterdam. Deharveng, L., C. Mouret, and D. Berenguer. 1979. Speleological investigations in Papua New-Guinea. Niugini Caver 7 (1): 15–21. Deharveng, L., and Y. Suhardjono. 2004. Pseudosinella maros sp. n., a troglobitic Entomobryidae (Collembola) from Sulawesi Selatan, Indonesia. Rev. Suisse Zool. 111 (4): 979–984. Flannery, T. 1995. Mammals of New Guinea. Reed Books Australia. Geoffroy, J.J. 2001. Papouasie-Nouvelle-Guine´e. Pp. 2133–2146 in Juberthie, C., and V. Decu (eds.) Encyclopaedia Biospeologica Vol. III. Socie´te´ de Biospe´ologie, Moulis. Gibert, J., and L. Deharveng. 2002. Subterranean ecosystems: a truncated functional biodiversity. Bioscience 52 (6): 473–481. Hobbs, H.H., III, 1998. Decapoda (Caridea, Astacidea, Anomura). Pp. 891–911 in Juberthie, C., and V. Decu (eds.) Encyclopaedia Biospeologica Vol. II. Socie´te´ de Biospe´ologie, Moulis. Humphreys, W.F. 2000. Background and glossary. Pp. 3–14 in Wilkens, H., D.C. Culver, and W.F. Humphreys (eds.) Ecosystems of the World 30: Subterranean Ecosystems. Elsevier, Amsterdam. Jasinska, E.J., and B. Knott. 2000. Root-driven faunas in cave waters. Pp. 287–307 in Wilkens, H., D.C. Culver, and W.F. Humphreys (eds.) Ecosystems of the World 30: Subterranean Ecosystems. Elsevier, Amsterdam. Jones, S. 1993. 1992 Caves of Thunder Expedition. Sedbergh, Cumbria. Jones, W.K., D.C. Culver, and J.S. Herman (eds.). 2004. Epikarst. Karst Waters Institute Special Publication 9, Charles Town, West Virginia, U.S. Lacas, M., P. Leclerc, and J.P. Mary. 2001. 5. Reconnaissance spe´le´ologique du secteur de FakFak (Ouest Irian Jaya). Pp. 47–57 in Rigal, D. (ed.) Indone´sie 92, rapport spe´le´ologique. APS, Toulouse. Leclerc, P., L. Deharveng, P.K.L. Ng, C. Juberthie, and V. Decu. 2001. Indone´sie. Pp. 1805–1823 in Juberthie, C., and V. Decu (eds.) Encyclopaedia Biospeologica Vol. III. Socie´te´ de Biospe´ologie, Moulis. 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 21: 1–216. Ng, P.K.L., and D. Guinot. 1997. Geelvinkia darnei, a new species of cavernicolous crab (Crustacea, Decapoda, Brachyura, Parathelphusidae) from Irian Jaya, Indonesia. Me´moires de Biospe´ologie 24: 181–182. Sarbu, S. 2000. Movile cave: a chemoautotrophically based groundwater ecosystem. Pp. 319–343 in Wilkens, H., D.C. Culver, and W.F. Humphreys (eds.) Ecosystems of the World 30: Subterranean Ecosystems. Elsevier, Amsterdam. Severens, H., H. van Eck, C. van Rijswijk, and T. Schuurmans. 1993. Irian Jaya 1992, Nederlandse Speleologische Expeditie. Expeditieverslag. Smith, G.B. 1980. 20-Biospeleology. Pp. 121–129 in James, J.M., and H.J. Dyson (eds.) Caves and Karst of the Muller Range. Atea 78, Newtown. Weygoldt, P. 2000. Whip Spiders (Chelicerata: Amblypygi): Their Biology, Morphology and Systematics. Apollo Books, Stenstrup.

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Caves of Papua / 1083 White, T. 1986. The 1985 Indonesia Expedition. Cave Science 13 (1): 25–45. White, W.B. 2005. Hydrogeology of karst aquifers. Pp. 293–300 in Culver, D.C., and W.B. White (eds.) Encyclopedia of Caves. Elsevier Academic Press. Whitten, T., R.E. Soeriaatmadja, and S.A. Afiff. 1996. The Ecology of Java and Bali. Dalhousie University, Periplus Editions. Willis, D. 1990. The caves of the Cloud Mountains, Irian Jaya: the 1988 expedition. Cave Science 17 (1): 39–49.

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section six 

Human-Ecosystem Interactions

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6.1. The History of Human Impact on New Guinea geoffrey s. hope u m an s h a ve b e e n in the upland valleys of New Guinea for at least 30,000 years and presumably occupied the savanna plains that then connected the island to Australia for as much as 50,000 years or more. Through this immense stretch of time people have adapted and changed their environments until very few places on the island can be considered completely unaltered. In place of primary rainforests and seasonal forests they have created human landscapes such the grasslands, secondary forests, and coastal woodlands. Prograded estuaries, infill in valleys, and eroded slopes may be partially caused by human actions, in addition to the deliberate creation of terraced slopes and ditched plains. Some faunal species have become extinct and rare, offset by introductions through time. This chapter reviews the history of human–environment interactions in Papua in three poorly defined periods: pre-agriculture (ca 55,000–20,000 years ago), the spread of agriculture (20,000–5,000 years ago) and post-Austronesian changes (5,000 years ago to present). These periods span the whole prehistory of New Guinea, during which modern people arrived from Southeast Asia and became adapted to a new environment of strange animals and plants. Humans litter the landscape with the tools they use and their use of fire amplifies their effect on habitats. Evidence for humans’ more indirect effects comes from dating geomorphological features such as alluvial fans, buried surfaces, activated sand sheets, and peaty infills in basins. Attribution of these features to human-caused erosion usually depends on correlations with archaeological deposits and specific human-caused features such as ditches, earthworks, and quarries. The other main line of evidence comes from paleoecology, in which the vegetation and faunas are reconstructed from dated fossil sequences. In Papua the main effort has been to use pollen, but elsewhere in New Guinea some specialized swamp ditch systems have also been investigated (e.g., Denham et al. 2004; Haberle et al. 1991; Sullivan et al. 1987). Past fire is inferred from swamp and lake sediments by counting microscopic charcoal fragments, and by dating larger fragments in a range of other deposits (Haberle et al. 2001). When Europeans contacted the island they found the population was scattered along the coastal fringe with quite sparse and isolated groups in the rainforests and mountain slopes (Figure 6.1.1). Some areas, such as the Foja Mountains, seemed to be totally uninhabited. The much larger highlands populations in the large intermontane valleys of the Baliem and Paniai lakes were a major discovery of the twentieth century. Brookfield (1964) showed that this complex agriculture, based on root cropping, centered on the altitudes of 1,400–1,850 m is controlled by the local climate. The outer flanks of the mountains are perhumid, with precipitation more than double evaporation in almost all months. Under such misty

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

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Figure 6.1.1. Location of archaeological sites and paleoecological sites showing human impact. conditions crops do not thrive (Hanson et al. 2001). In the intermontane valleys the mountains cut off this regional rain and local circulations dominate in most seasons. Here air rises each day up the warmed slopes and descends over the valley, giving sunny conditions with adequate rainfall from afternoon thunderstorms. Away from the large highland basins, even small valleys may have this effect, and may thus support small hamlets.

Early Settlement Perhaps because of the dense populations and a greater concentration of research, evidence for the effects of early settlement in the highlands is slightly more abundant than in the lowlands. People may have been present by 32,000 bp (years before present) or earlier in the Baliem Valley (Hope 1998; Haberle et al. 2001) and are known at Kosipe (White et al. 1970) and near Nombe (Mountain 1993) in Papua New Guinea by this time (Figure 6.1.1). A waisted blade has been found

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in the Baliem Valley similar to those from Kosipe so it seems probable that some populations were present, hunting and gathering plant foods. The ice age climates of New Guinea were drier than present, and savanna probably extended right across the Torresian Plain. The climate at high altitudes was also colder, with ice caps and extensive alpine grasslands on many mountains along the central ranges. Pollen evidence from peat sections in the upper Baliem (Hope et al. 1993; Hope 1998) and Lake Sentani (Hope and Tulip 1994) provides records back 50,000 years or more. These suggest that closed tropical and montane forests have continuously occupied many areas from before the likely arrival times for people. The wet conditions that supported the rainforests seem to have been maintained all along, indicating that the warm tropical waters north of New Guinea named the Western Pacific Warm Pool have persisted since the late Pleistocene (Thunnell et al. 1994). However, it is likely that rain shadow effects north of the main range were strengthened. The Sentani and Sepik areas may have been drier than present, dominated by Nauclea woodlands and possibly experiencing natural fires. A record of fire and erosion dating from 32,000–26,000 bp has been obtained from fossil charcoals on Supulah Hill (Figure 6.1.2), north of Wamena (Hope 1998). While not directly associated with human activity the appearance of charcoal in a swamp section hints at early creation of clearings in beech forests, possibly associated with the gathering of Pandanus nuts. Large pieces of charcoal were found in sand sheets that washed from the hill 26,000 years ago, showing that the fires were extensive and resulted in erosion. Some sedimentary sequences, for

Figure 6.1.2. Supulah Hill in the Baliem Valley was burnt, cleared, and eroded about 33,000 years ago.

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example peat beds at Tari, PNG, and algal sediments from Lake Hordorli, in the Cyclops Mts near Jayapura, record no fires at all over tens of millennia in the Pleistocene. In such places the appearance of charcoal is probably an indicator of human activity (Haberle et al. 2001). The pollen record from the Tari Basin is the only continuous peat record from before 28,000 bp through to the present (Haberle 1998). Prior to 21,000 bp, forests dominated by Nothofagus, Castanopsis, and Myrtaceae covered the basin floor. Just before the onset of the last glacial maximum at around 21,000 bp we find the first evidence for burning that created a mosaic of grassland and forest (Figure 6.1.3). Although there is no direct archaeological evidence for humans in the basin at this time, the rapid increase in burning and the opening up of the vegetation is unprecedented in earlier glacial records from the basin, and is therefore considered to be a consequence of the arrival of humans in the region. This is at least 10,000 years later than the charcoal records from similar forests at Supulah Hill. The Tari record may represent a later occupation of a wetter site. The early human spread into the highlands may also have included hunting in the higher grasslands. For example fossils of a calf-sized diprotodontid, Maokopia ronaldii, and a robust kangaroo, Protemnodon hopeii, have been recovered from Kwiyawagi in the upper Baliem where it seems to have been adapted to extensive subalpine grasslands (Flannery 1992; Hope et al. 1993). The subalpine fauna seems

Figure 6.1.3. Charcoal histories through time from selected sites in western New Guinea.

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to have disappeared well before the warming climates of 14,000 years ago, at which time forest limits rose and the mountain grasslands diminished. Hence some other cause (which may include human hunting or disturbance) must be involved. A more direct case for human interaction is known from Nombe Cave in the Simbu of PNG (Flannery et al. 1983). Here the bones of an extinct fauna occur in the horizons just preceding human artifacts. The altitude of Nombe makes it likely that it was forested, hence hunting may have been a more gradual process than in open country. Human predation on grazing and browsing animals may have had significant effects on vegetation structure, allowing forest invasion of open areas to proceed more rapidly.

Clearance and the Start of Agriculture Evidence for the extension of human landscapes is apparent on sensitive ecological boundaries such as the savanna-rainforest ecotone in the lowlands and the alpine tree line during the period of climate transition about 12,000–10,000 years ago. In these locations forest encroachment has been resisted by fire and perhaps active clearance. As global temperatures warmed and glaciers retreated, the late glacial transition in the highlands was achieved in a two-phase warming sequence with an initial period of climatic instability between 14,500 and 12,000 bp, followed by a more persistent warming between 12,000 and 8,500 bp. At high altitude this led to an elevation in forest growth limits and a replacement of tree ferns and grasslands with a closed upper montane forest (Hope 1989). In the montane valleys the combination of increasing mean annual temperatures, high atmospheric CO2, and strengthening monsoon influence (Haberle et al. 2001) would be expected to result in expansion of forests into grassland habitat. While charcoal becomes common at that time, forests do generally expand. The earliest indications for the appearance and spread of ‘‘agriculture’’ in the highland valleys is seen at Kuk, PNG with the ditching within a mosaic of forest and grassland around 9,000 bp (Denham et al. 2003, 2004; Haberle 2003). This date accords remarkably well with the transition to ‘‘modern’’ Holocene climates, suggesting that expansion of clearing and plant manipulation was partly environmentally controlled. In Papua, the record of forest clearance from the Baliem is a little later than at Kuk (Haberle et al. 1991). By 7,000–6,000 years ago the lower parts of the major highland valleys had been cleared and crops such as taro and banana established (Figure 6.1.4). It is possible that the early Holocene was a time of more reliable climates when El Nin˜o–related drought and frost events were much rarer (Groves and Chappell 2000). Cores taken by Rien Dam at Anggi Lake show increased burning and grassland taxa after about 5,000 bp but fire is present from before 9,000 bp (S. Haberle, pers. comm.). Thus agriculture is broadly contemporaneous in good montane valleys. At high altitude, some subalpine lake records contain evidence of fire almost as soon as the ice retreats (Hope 1996) and pollen diagrams from Lake Habbema, an area north of Mt Trikora, show continuing disturbance to the present day (Figure

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Figure 6.1.4. The irrigated sweet potato fields of the Baliem Valley with scattered trees of Casuarina oligodon represent a high level of communal organization and hydraulic and arboricultural sophistication developed through the Holocene to tend taro. 6.1.3). At Mt Jaya burning started about 11,000 years ago and the Mapala rock shelter recorded hunting from 5,500 years ago that resulted in the extinction of a small wallaby (Thylogale christensenii) (Figure 6.1.2). Hope et al. (1993) speculate that pressure of hunting allowed the Copper Ringtail Possum, Pseudocheirops cupreus, to expand into the subalpine niche. This hunting probably postdates the development of large human populations at lower altitudes, so is indirect evidence of early Holocene expansion in the highlands. Clearance in isolated valleys and lower montane sites may have been much later, in the middle or late Holocene. Higher altitude sites, such as Kwiyawagi at 2,900 m, may only have been settled when cold-tolerant crops such as sweet potato, Ipomoea batatas, and potato, Solanum tuberosum, arrived in the last few centuries. Such altitudinal limits are still being tested, such as around 3,000 m at Iniuni south of Tiom, where subalpine forest is being cleared for cabbage gardens and pig grazing (Figure 6.1.5). In the lowlands the disturbance history resembles that of the highlands, which is curious considering the 50,000-year period of occupation of the coast and lowlands (O’Connor and Chappell 2004). Expansion of grasslands, marked by a charcoal record at Lake Hordorli, occurs in the Lake Sentani region at 11,200 bp (Hope and Tulip 1994; see Chapter 2.7) and it is likely that this reflects the development of early agriculture. Occupation of the area from before 25,000 bp is known from Lachitu rock shelter, about 60 km east of Jayapura (Gorecki et al. 1991) but this earlier occupation was not associated with extensive fire.

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Figure 6.1.5. Settlement at its limits: clearing primary forest above 3,000 m at Iniuni, East Baliem Valley. These fields are mostly planted with cabbage.

The Arrival and Effects of Austronesian-speaking Agriculturalists The Austronesian arrival around 4,000 years ago shows very little correlation with environmental change. However, the frequency of El Nin˜o events may have increased, ushering in the drought and frost events that have an effect on cropping and societies at all altitudes (Brookfield 1989; Brookfield and Allen 1989). These droughts may also have allowed fire to extend grasslands into humid forests. Grasslands of Imperata cylindrica (kunai) occur in many areas on poor soils such as the iron- and magnesium-rich ultramafics of Sentani. These clearings are probably of considerable antiquity as regeneration may take thousands of years. The pollen of Casuarina becomes much more common across the highlands after ca 1,800 years ago, suggesting that this was a time of widespread silvicultural planting (Haberle et al. 1991). This rapid spread supports a hypothesis that there was effective diffusion of ideas despite linguistic and social barriers. Similarly, the spread of crop plants such as sweet potato also seems to have been almost universal in the highlands, resulting in the new clearance of slopes and higher altitudes. This becomes apparent in the last few centuries, and results in the abandonment of some valley floor swampland field systems. The floor of the Baliem has large areas of apparently abandoned field systems and swamps which may have been abandoned for some time, judging from the Kelela pollen record (Haberle et al. 1993). Away from the major centers there are some sites that show increased exploitation in the later Holocene compared to the present. At higher altitudes forest regeneration and tree fern regrowth is widespread today, suggesting some reduction of human pressures over the past century. While this may be the result of

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changes associated with recent new crops and other social changes, it seems to have started earlier, for example, than the introduction of sweet potato. This is suggested by the charcoal curve from Ijomba site at 3,720 m in the Jaya (Carstensz) Mts where burning has been reduced in the last 1,500 years (Hope and Golson 1995). North of Sentani in the Cyclops Mts, a major period of patchy clearance around Lake Hordorli is evidenced by charcoal and secondary species around 4,000 years bp (Hope and Tulip 1994). The area is little used at present although sago palms can be found in creek lines. This is evidence for more scattered settlement in the mid-Holocene in an area right on the limit for agriculture.

Discussion There is much we still do not know about the environmental history of New Guinea. While we can be sure that the open grasslands and agriculture today are human landscapes, in only very few cases do we know when the landscapes appeared. The more we investigate the more individualistic site histories are uncovered. The midmontane grasslands at Telefomin have been in place for about 4,000 years (Hope 1983) but clearances that lasted some centuries also occurred 11,000 and 7,500 years ago. Other records are incomplete but we are unsure of the causes of a ‘‘gap’’ in sedimentation that occurs in many sites. At Supulah Hill no sediments are known from the period 30,000–2,500 bp, by which time the area was as deforested as it is now. Only a few sites cover the gap in the late Pleistocene. Perhaps at the end of the glacial maximum a phase of dry weather and widespread fires occurred, but catchment clearance may also be implicated. This lack of detail is frustrating because of our inability to relate the modern diversity to its past history. This will remain a major aim as the framework of sites and records is strengthened. The diversity of human culture and language must reflect the need for local adaptation to specific environments that change over distances of a few kilometers. Yet on a time scale of many centuries similar patterns of settlement and technology appear at the same time across the island. Diffusion of cultivars, land management practices, and other techniques such as pottery or silviculture seem to have been relatively rapid despite the isolation of many groups. The most rapid time of climate change and coastal stress, from ca 15,000–8,000 years ago, rewarded adaptive cultures. The burning of some of the high altitude grasslands by 13,000 bp, 4,000 years before the substantial clearance in the montane basins, suggests that trading links were in place across the mountains before the agricultural populations had increased. Similarly, the scattered evidence for clearance and burning by 34,000 bp suggests that some ecological manipulation and selection was already taking place (Fairbairn et al. 2006). Hence the emergence of an agricultural landscape in the highlands of New Guinea can be seen as a result of gradual indigenous development punctuated by external influences such as introduced domestic plants and climate change and variability. Any ‘‘foreign’’ influences would be small

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and possibly isolated to single locations in the Central Cordillera. Our records come from swamps and these may have been the birthplace of an agriculture that depended on aroids and banana. Papua possibly had its ‘‘big game hunters’’ in the Pleistocene, but the evidence is so diffuse that we can only note another unexplained extinction event and wonder if there might be anything in the origin myths of strange animals and birds (Roberts et al. 2001). However, the loss of species of rodents in the New Guinean islands (Spriggs 1997) around 30,000 years ago is clearly coincident with human settlement. Flannery (1992) has suggested that a second phase of effective hunting of arboreal mammals could start only when the dog arrived about 3,500 years ago. With the establishment of large areas of secondary vegetation, a reduction in the significance of hunting has continued to the present day

Acknowledgments I am grateful to Chris Ballard, Michael Bourke, Tim Denham, Tim Flannery, Yance de Fretes, Pawel Gorecki, Simon Haberle, Juliette Pasveer, Ron Petocz, and Pamela Swadling, for their knowledge of New Guinea past and present. I am also indebted to LIPI, the Departments of Kehutanan and Kependudukan dan Lingkungen Hidup, the Universitas Cenderawasih, P.T. Freeport, SIL and MAF, as well as NGOs in Papua such as WWF and CI for their permission and support. The work has been dependent on the farmers and hunters of Papua who shared their unique knowledge with us and provided practical assistance. I thank the Australian Research Grants Scheme and the Australian National University for financial aid.

Literature Cited 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. Denham, T., S. Haberle, and C. Lentfer. 2004. New evidence and revised interpretations of early agriculture in Highland New Guinea. Antiquity 78: 839–857. 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. Fairbairn, A., G. Hope, and G. Summerhayes. 2006. Pleistocene occupation of New Guinea’s highland and subalpine environments. In Archaeology at Altitude. World Archaeology 38: 371–386. Flannery, T.F. 1992. New Pleistocene marsupials (Macropodidae, Diprotodontidae) from subalpine habitats in Irian Jaya. Alcheringa 16: 7–23. Flannery, T.F., M.J. Mountain, and K. Aplin. 1983. Quaternary kangaroos (Macropodidae: Marsupialia) from Nombe rock shelter, Papua New Guinea, with comments

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1096 / geoffrey s. h o p e on the nature of megafaunal extinction in the New Guinea Highlands. Proc. Linnean Soc. New South Wales 107 (2): 75–97. Gorecki, P.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. Groves, R.H., and J. Chappell (eds.). 2000. El Nin˜o—History and Crisis. The White Horse Press, Cambridge. Haberle, S.G. 1998. Late Quaternary vegetation change in the Tari Basin, Papua New Guinea. Palaeogeography, Palaeoclimatology, Palaeoecology 137: 1–24. Haberle, S.G. 2003. The emergence of an agricultural landscape in the highlands of New Guinea. Archaeology in Oceania 38: 149–158. 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. Hanson, L.W., B. Allen, M. Bourke, and T.J. McCarthy. 2001. Papua New Guinea Rural Development Handbook. Land Management Group, Australian National University, Canberra. Hope, G.S. 1983. The vegetation changes of the last 20,000 years at Telefomin, Papua New Guinea. Singapore Journal of Tropical Geography 4: 25–33. 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. CSIRO, Div. Water Resources, Canberra. Hope, G.S. 1996. 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. 1998. Early fire and forest change in the Balim Valley, Irian Jaya, Indonesia. J. Biogeography 25: 453–461. Hope, G.S., T. Flannery, and Boeardi. 1993. A preliminary report of changing Quaternary mammal faunas in subalpine New Guinea. Quaternary Research 40: 117–126. Hope, G.S., and J. Golson. 1995. Late Quaternary change in the mountains of New Guinea. In Allen, F.J., and J.F. O’Connell (eds.) Transitions. Antiquity 69 (265): 818–830. Hope, G.S., and J. Tulip. 1994. A long vegetation history from lowland Irian Jaya, Indonesia. Palaeogeography, Palaeoclimatology, Palaeoecology 109: 385–398. Mountain, M-J. 1993. Bones, hunting and predation in the Pleistocene of northern Sahul. Pp. 123–130 in Smith, M., 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. O’Connor, S., and J. Chappell. 2004. Colonisation, settlement and subsistence in Greater Australia from 60,000 BP. Pp. 15–32 in Sand, C. (ed.) Proceedings of the International Conference for the 50th Anniversary of the first Lapita Excavation Kone-Noumea 2002. Les Cahiers de l’Arche´ologie en Nouvelle Cale´donie 15, Service des Muse´es et du Patrimonie de Nouvelle Cale´donie, Noume´a. Pasveer, J. 2003. The Djieff Hunters. Rijksuniversitat Groningen, Groningen. Roberts, R.G., T.F. Flannery, L.K. Ayliffe, H. Yoshida, J.M. Olley, G.J. Prideaux, G.M.

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The History of Human Impact on New Guinea / 1097 Laslett, A. Baynes, M.A. Smith, R. Jones, and B.L. Smith. 2001. The last Australian megafauna: new ages indicate continent-wide extinction about 46,000 years ago. Science 292: 1888–1892. Spriggs, M. 1997. The Island Melanesians. Blackwell, Oxford. Sullivan, M.E., P. Hughes, and J. Golson. 1987. Prehistoric garden terraces in the Eastern Highlands of Papua New Guinea. Tools and Tillage 260: 199–213. Thunell, R., D. Anderson, D. Gellar, and Q. Miao. 1994. Sea-surface temperature estimates for the tropical western Pacific during the last glaciation and their implications for the Pacific Warm Pool. Quaternary Research 41: 255–264. White, J.P., K.A.W. Crook, and B.P. Ruxton. 1970. Kosipe: a late Pleistocene site in the Papuan highlands. Proceedings of the Prehistoric Society 36: 152–170.

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6.2. A Brief Social and Political History of Papua, 1962–2005 jaap timmer h i s c h a p te r presents an overview and assessment of the social and political developments in West New Guinea since its incorporation in the Republic of Indonesia. The roots of the poor and unequal development in the Province of Papua become evident when quantitative data from 1961 and 1962 produced by the government of Netherlands New Guinea are compared with recent human development indicators and indices developed by the Central Bureau of Statistics (BPS: Badan Pusat Statistik), the National Development Planning Agency (BAPPENAS: Badan Perencanaan Pembangunan Nasional), and the United Nations Development Program Indonesia. The vast variation in human development between urban and rural areas, among regions, and between Papuans and non-Papuans clearly reveals the effects of weak education and health services for rural communities, the preference given to immigrants for manual labor jobs, and the marginalization of Papuans in business ventures at all scales. Thirty-three years of highly centralized New Order governance (1965–1998) has led to widespread feelings of dispossession, a spate of tensions with respect to natural resource extraction projects, a generally low level of educational achievement, limited chances on the job market for most Papuans, and the hampering of delivery of medical services and related efforts aimed at preventing the spread of HIV/AIDS. These developments largely explain the prevalence of expressions of resentment towards the central, provincial, and regional governments currently still voiced by many Papuans.

T

Introduction Any attempt to review the social and political developments in Papua must examine the often unsettling gap between the world of a centralized government and national and international business practices on the one hand and, on the other, the development of living conditions of the people of Papua and local struggles for sovereignty and dignity. Below I identify the characteristics and effects of this gap by reflecting on human development in Papua over the past four decades. The history of Papua, apart from overviews of its ‘‘discovery’’ and foreign expeditions in search of its riches, biodiversity, and peoples, has not attracted much critical study. General but often detailed overviews of the history of West New Guinea include Vlasblom (2004), Moore (2003), Defert (1996), Whittaker et al. (1975), Souter (1963), and Klein (1935–1938); see also Ballard (1999). Most hisMarshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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torical research on Papua largely overlooks the social and economic effects of colonial and post-colonial governments trying to impose a tidier identity on a supposedly unruly people. The classic anthropology of the region’s cultures is relatively rich but generally does not pay much attention to the integration of Papuans into Indonesia and the related changes in local histories and ideals of sovereignty (but see Rutherford 2003; Oosterhout 2002; Glazebrook 2001; Timmer 2000; Giay 1995). In general, the study of contemporary conditions in Papua is flawed with respect to the social, economic, political, and demographic dynamics that explain Papuan responses to the generally large gap between government policies and the needs and aspirations of the people. In addition to realizing that there are pockets of resistance against the Indonesian government throughout the region, it is also important to realize the extent to which Papuans have embraced the opportunities to become Indonesian and to play roles in the informal and business sectors (see Timmer 2005). Towards the end of this chapter these developments are analyzed to indicate that people in Papua know how the Indonesian state works and have learned how to profit from companies wanting to buy their trees, mine their ores, drill their gas, and so on. Over the last few decades, civil society organizations and Papuan intellectuals have also begun to surface in politics and policy roles in Papua in increasingly significant ways. Indications of powerful changes in the social landscape of the region include nationwide appeals and support for peaceful protests, demonstrations, and seminars organized by networks of Papuan students, the drafting of and lobbying for Special Autonomy regulations by academics and bureaucrats in Jayapura, and a growing number of critical writings by Papuans. The high turnout of voters and the generally smooth implementation of the 2004 national parliament, provincial legislation, and presidential elections in Papua demonstrated the growing will to support civilian-led government and the rule of democracy in Indonesia since the fall of New Order’s President Suharto in 1998. This faith in the possibility of having a voice through democratic means is part of a larger process of people in Papua integrating Western and Indonesian reflections on colonial history, Christianity, and New Order nation-building and development. In general, we see that the people of Papua are moving away from colonial shackles and old and new subservient positions. Yet these positive developments occur amid increasingly unsettling disparities in social and economic development in the region and fear of poorly monitored military actions. The study of the social and cultural developments in West New Guinea (now Papua) since its incorporation in the Republic of Indonesia is inevitably hampered because access to the region for researchers and the press, as well as for development organizations, has been severely restricted. Data on demographic changes, economic developments, and social and cultural change are scarce and, if available, generally inaccurate. Nevertheless, the magnitude of changes in Papua since 1962 is obvious, if only through activist accounts, travelogues, news reports, and data collected by the Central Bureau of Statistics (BPS) of Indonesia. More accurate

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analyses are available in the form of a handful of detailed anthropological studies by Indonesian and foreign scholars, a number of studies of economic change, a large variety of detailed NGO reports, and careful assessments of the development potential of the region done by the United Nations Development Program in the 1960s and early 1970s (see below). The general picture that emerges is that, after four decades of Indonesian government in Papua, the imposition of a centralized formal sector on a large variety of local groups with distinct identities and histories of contact with regional neighbors and a larger world, has markedly changed people’s ideas about the state, democracy, and their own future.

Transfer of the Territory For a number of reasons, ranging from Dutch nationalism, geopolitical considerations, and self-righteous moral convictions, to wanting to keep the territory for immigration (in particular for Eurasians from the Netherlands Indies), the Netherlands government resisted including West New Guinea in the negotiations for the independence of Indonesia in the late 1940s (see de Geus 1984; Gase 1984; van Galen 1984; Huydecoper van Nigtevecht 1990; Lijphart 1966; and Penders 2002). Around the same time, the Netherlands government set about economic and infrastructure development as well as, at later stages, political development of the Papuans under paternalistic guardianship. In the course of the 1950s, when tensions between the Netherlands and Indonesia grew over the status of West New Guinea, the Dutch began to guide a limited group of educated Papuans towards independence, culminating in the establishment of the New Guinea Council (Nieuw-Guinea Raad) in 1961. In addition, the Morning Star flag (Bintang Kejora) was designed to be flown beside the Dutch flag, and a national anthem, Hai Tanahku Papua, was adopted to be played and sung during official occasions after the Netherlands national anthem. After a twelve-year dispute that was reaching its peak with the threat of open military conflict, this policy had to be aborted. With the threat of an Indonesian invasion and the unwillingness of the United States and Australia to support the Netherlands military forces, The Hague was forced to accept the Indonesian claim that it was the legitimate successor to the entire territory of the former Netherlands Indies. In December 1961, President Sukarno issued the People’s Threefold Command (Trikora: Tri Komando Rakyat) for the liberation of Irian Barat (West Irian, now Papua). At the heart of this massive mobilization was Operasi Mandala, an Armed Forces of the Republic of Indonesia (ABRI: Angkatan Bersenjata Republik Indonesia) campaign designed to put pressure on the Netherlands government. Following years of persistent but low-scale infiltration into Netherlands New Guinea that met limited support by Indonesia supporters among the Papuans, the Indonesian Air Force and paratrooper units readied themselves at East Indonesian airfields and ports such as Ambon (Maluku) and Makassar (South Sulawesi). The

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Mandala military authority, headed by Major General Suharto, had authority over the entire Eastern Indonesian war zone. Despite the impressive number of troops numbering 42,000, Operasi Mandala was chiefly a political instrument and did not have the capacity or the intention to invade West New Guinea. In fact, most troops of the Mandala force did not come into action. Nevertheless, Dutch defenses intensified over the last few years and were tested in January 1962 when three German-built Jaguar torpedo boats of the Indonesian Navy entered Dutch waters and were fired upon. One ship, the Macan Tutul, was sunk and the commander Yos Sudarso of the Indonesian Navy was killed (Vlasblom 2004: 314–315). To avoid further confrontation and in an international climate of decolonization, and after President Sukarno’s sustained pressing of Indonesia’s claim to the territory, the United States sponsored negotiations between Indonesia and the Netherlands. While Indonesia dropped hundreds of paratroops in Irian Barat (West Irian) and the Dutch Navy carrier Karel Doorman sailed to the region, the two parties sat around the table to negotiate the future of West New Guinea under the auspices of the United Nations. No delegation from Papua was present (Vlasblom 2004: 329–330). The resulting New York Agreement of 15 August 1962 outlined the transfer of Netherlands sovereignty over West New Guinea to an interim United Nations Temporary Executive Authority (UNTEA) from 1 October 1962 to 1 May 1963. This was to be followed by a second phase during which the intervening administration would hand over full administrative responsibility to Indonesia. The agreement formulated the provision that the people of Irian Barat would exercise free choice over their future relationship with Indonesia before the end of 1969. On 30 September 1962, the nationalist Papuan Morning Star flag that had flown beside the Dutch flag on the building of the New Guinea Council in Hollandia (now Jayapura) was lowered. On 1 October 1962, Irian Barat became the first United Nations-administered territory. The United Nations Temporary Executive Authority (UNTEA) lacked the capacity and the authority to be sensitive to anti-Indonesian sentiments among elements of the Papuan elite and a growing fear among some rural communities that perhaps Indonesia would not continue the feverish development efforts of the Dutch over the last decade. As of 1963, the country became occupied by large numbers of Indonesian troops, no longer under the Mandala military force but as part of the Regional Military Command (Kodam: Komando Daerah Militer) XVII/ Irian Barat, which after 1 July 1964 became known as Kodam XVII/Cenderawasih. The victory over what had now become the Province of Irian Barat was a boost to Indonesian nationalism, and was later portrayed as the final chapter of decolonization (see Soekarno 2000). The Indonesians ruling Irian Barat under the banner of the Trikora mobilization were triumphant, while elements of the Papuan elite empowered by the Dutch began to complain about what they saw as a blunt Indonesian takeover. Among urban Papuans, feelings of being marginalized by Indonesian bureaucrats arose as immigrants from other Indonesian islands began

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filling jobs and taking business opportunities. Some of the urban Papuans were arrested or sidelined as ‘‘collaborators’’ with the Dutch, while others continued to play a role in the administration. A plebiscite called Act of Free Choice (Pepera: Penentuan Pendapat Rakyat) was held in July–August 1969, during which 1,020 carefully chosen representatives from eight regions voted overwhelmingly for integration with Indonesia. A total of 1,025 representatives were chosen from regional councils (excluding military members), political parties, civil society and church organizations, and customary leaders. Overall, five people did not show up, of whom one was reportedly executed because he did not stick to the dictated text during a speech repetition (Pepera 1972: 82–83; Vlasblom 2004: 479). While voting for integration, Eduard Hegemur from Fakfak gave a dissonant speech in which he said that he could not betray his friends who fought for independence, and demanded that Indonesia grant the Papuans their legitimate right to self-determination (Pepera 1972: 208; Vlasblom 2004: 474). On top of that, desperate outcries in the form of written notes were handed over to the United Nations observers and demonstrations in Sukarnopura (formerly Hollandia, now Jayapura), Biak, and Manokwari were dispersed swiftly by the Indonesian military. Over the years a growing number of Papuans would dispute the results of the Pepera plebiscite by claiming that the votes for integration were all cast under pressure from the Indonesian military. Elements in Papua became resentful of international ‘‘betrayal’’ and demanded the Netherlands government and the United Nations take responsibility over the allegedly unfair outcome of the Pepera plebiscite. Over the following decades, the belief in Papuan self-determination, coupled with the belief in the undemocratic implementation of the Pepera plebiscite favoring integration into Indonesia, became a key and consistent ingredient in a variety of Papuan nationalist movements. In response to the ‘‘Indonesian occupation’’ of their land, during the course of the 1960s, a liberation organization called the Free Papua Organization (OPM: Organisasi Papua Merdeka) emerged as a local movement in Manokwari and from there spread over the Vogelkop region (see Vlasblom 2004: 469, 513 ff, 632 ff). The Free Papua Organization (OPM) soon became a fragmented network of dispersed groups of guerilla fighters. Its access to weapons was limited and popular support scant. Very few elite Papuans joined the armed struggle. The sense of belonging to a nation that had been invaded by Indonesia was not felt by the vast majority of people living outside the small urban centers. Nevertheless, disillusionment with the Indonesian government became widespread among those who had enjoyed the fruits of the accelerated development effort of the Dutch government since the 1950s. In early February 1969, the military launched Operasi Wibawa (Operation Authority) with the aim of ending the law-and-order disturbances in the Vogelkop Peninsula, in the Paniai region, and near the border with Papua and Papua New Guinea. These were the three main locations where the Free Papua Organization

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(OPM) was pursuing a small-scale armed resistance against the Indonesian government and its armed forces. A more pressing goal of Operasi Wibawa was to ensure a favorable outcome of the Pepera plebiscite on integration, and to consolidate the authority of the Indonesian government as established under a previous operation called Operasi Sadar (Operation Consciousness). During the following few decades, five Free Papua Organization (OPM) command structures kept the ideal of a free West Papua alive but their regional network rapidly dissolved due to internal divisions. The OPM diversified and became largely an instrument in the hands of disgruntled people for establishing their own power bases, now confined to forests mainly in the central highlands. Since the 1970s, most Papuans have grown up with the idea of the Free Papua Organization (OPM) without having a clear sense of where it is, who leads it, and what its strategies are, but cherishing the idea of ‘‘OPM’’ is a state of mind shared by all Papuans.

Years of Reconstruction Until 1935, Netherlands New Guinea was largely unknown to Western and Indonesian outsiders but, starting shortly before World War II, an increasing number of expeditions for mapping local communities and to search for oil and gold began, amid increasing missionary activity. After the war, the Dutch put considerably more effort into the development of infrastructure, largely building on the efforts of missionaries, and into providing health services and schooling to Papuans. Economic development was minimal (see Lagerberg 1962; Schoorl 1996). There were a small number of European coconut, coffee, and kapok plantations and one Japanese venture which grew cotton and collected copal. Throughout the 1950s, economic development was expected to come from further exploration and development of mineral extraction, particularly the development of oil reserves (which proved unsatisfactory towards the end to the 1950s), the vast forest resources (for which there was little interest abroad), a few coconut, cocoa, and nutmeg plantations largely owned by non-Papuans, and some experimental small-scale commercial agriculture projects. The most significant resource extraction development was the oil industry in the Vogelkop region where the Netherlands New Guinea Oil Company (NNGPM: Nederlandse Nieuw-Guinea Petroleum Maatschappij) discovered a major oilfield near Sorong which was brought into production in 1949. However, by 1962 the production at this site had virtually ceased (see Poulgrain 1999). The labor force at the oil fields was mostly Indonesian and Dutch. Rural economic development concentrated on the cultivation of export crops like copra, nutmeg, cocoa, coffee, and rubber. Most of these initiatives were taken by the Agricultural (and Stockbreeding) Department of Economic Affairs, while privately owned plantations were scarce. Economic development during the last phase of Dutch Government in New Guinea ‘‘was geared towards the discovery of

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ways and means to unleash the initiative of the Papuans themselves’’ (Pouwer 1999: 171). In the course of the 1960s, social and economic conditions in Irian Barat (West Irian or Papua) deteriorated. The local administration became a refuge for mainly untrained staff that had to operate with a shortage of funds and equipment. Furthermore, a confrontation campaign that Indonesia launched against Malaysia in 1963 over the future of the former British colonies in North Borneo drained Indonesia’s skills and resources. As a result, Irian Barat was put significantly lower on the Indonesian national agenda, while frustration over the United Nations’ membership of the ‘‘neo-colonial construct of Malaysia’’ led Jakarta to decide to quit its membership of the United Nations. As part of the decolonization agreement, the Dutch promised the Indonesian government that they would contribute with a grant-in-aid of US$ 30 million to a Fund of the United Nations for the Development of West Irian (FUNDWI). As a result of Indonesia’s decision to suspend membership of the United Nations much of the FUNDWI effort at rehabilitation was suspended. Assessment and planning for FUNDWI programs resumed after Jakarta decided to renew its participation in the United Nations in November 1966. In accord with the Netherlands government, the administration of the fund was entrusted by the Secretary General to the Administrator of the United Nations Development Program (UNDP) in February 1967. A FUNDWI Secretariat was established to implement a comprehensive development program in the province. The Indonesian National Development Planning Agency (BAPPENAS: Badan Perencanaan Pembangunan Nasional) incorporated the funds into the first Fiveyear Development Plan (REPELITA I, Rencana Pembangunan Lima Tahun, 1969– 1974). In 1970, US$ 21 million was allocated for FUNDWI-assisted programs largely focusing on infrastructure and rehabilitation (see UNDP/FUNDWI 1975). It was clear that developing Irian Barat would be an arduous task and consequently Irian Barat received about four times more funds than any other part of Indonesia. To this day Papua remains a particularly favored province in terms of resource allocation. However, only a relatively small proportion of these allocated funds are for the development of Papuans. As noted in the final FUNDWI report of 1974, in terms of economic activity at the community level, by far the most potent influence in the private sector was the spontaneous (and organized) influx of immigrants from other Indonesian islands, who took up a large stake in local fisheries, timber business, retail trade, and supply of labor (UNDP/FUNDWI 1975: 10; and see Aditjondro 1986; Arndt 1986; Manning and Rumbiak 1989). This development and the concurrent mounting feelings of displacement among Papuans increased over the years and are to this day a key source of resentment among Papuans toward immigrants and the Jakarta government. After General Suharto assumed powers from President Sukarno in 1965, the government launched the New Order approach to the development and good governance of Indonesia. The effort to stimulate development and economic

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growth continued. Support for human development increased and primary and secondary education services were provided with many subsidies to the most remote communities. Moreover, the government tried to stimulate development by employing a large number of people with impressive technical skills, but failed to prioritize the local Papuan people. It has not always been easy to assign seasoned administrative and technical personnel in this distant province, despite financial and other incentives. On top of that, many of those deployed to the region believed that after a few decades of Dutch colonial politics encouraging antiIndonesian sentiment, they had to make Papuans into Indonesian citizens. This line of thinking also suggested that Papuans were still prone to being influenced by discontented individuals or by outsiders who wanted to destroy the unity of Indonesia. One of the most prominent civilizing efforts of this period was Operasi Koteka (Penis Gourd Operation). This operation was targeted at highland communities, most of which had experienced contact with government no more than one or two decades earlier. One of the aims was to convince them to wear clothing so as to protect them from outsiders who might see their near-nudity as offensive (see Vlablom 2004: 499–503 for an interview with Acub Zainal, then military commander for Irian Barat, who led Operasi Koteka). The operation met with resistance and after a few years the military command began to appreciate the argument of the local Catholic Church that Papuan customs should be respected. In the course of the 1970s, the civilizing effort abated. As for economic development under the New Order, the number of permits for the mining, forestry, and fishing industries increased. After the Indonesian government offered a new arrangement that allowed oil and mining companies to keep a fairly large proportion of their profits, a contract with the United Statesbased Freeport Sulfur was signed. On the basis of exploration results dating back to the Dutch period and an initial period of mining in the mid-1960s (before the decolonization of the territory), copper and gold mining in the western highlands got substantially underway and appeared highly profitable. (See Leith 2003 for an analysis of the nature of power relations between Freeport and President Suharto, the military, the Amungme and Kamoro landowners, and environmental and human rights groups. For details of the exploration of the copper deposit in the 1960s and the setting-up of the mine facilities, see Wilson 1981.) President Suharto celebrated the mining industry and the wealth of the region when he visited the Freeport mine site in 1973. It was on this occasion that he renamed the province Irian Jaya or ‘‘Glorious Irian.’’ In conclusion, throughout the New Order period (1965–1998), Jakarta often stressed that Papuans did not understand that the central government was spending a lot of money to improve the living standards of the people. What was needed, according to policy makers, was to educate these Papuan masses to the point where they became full members of the Indonesian nation. This top-down approach continues today and leads to Papuans and outsiders viewing each other in often incompatible ways, causing disappointment, resentment, and protest. Since

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the fall of President Suharto in 1998, the region has experienced a number of upheavals and conflicts related to Papuans’ increasing aspirations for independence and a variety of inconsistent approaches from the central government.

Four Decades of Human Development This section presents a general picture of the situation in which the people of Papua find themselves after over forty years of efforts of the Indonesian government at development of the region since its integration into the republic. The analysis is largely based on statistical information. For the final years of Dutch administration in West New Guinea, we have the summarizing reports on Netherlands New Guinea produced for the United Nations. In addition, we have the results of a detailed demographic study sponsored by the European Economic Community (EEC) and implemented in 1961 and 1962 in the Cenderawasih Bay, Nimboran, Fakfak, and Muyu regions (Groenwegen and van de Kaa 1964). For 2000 and 2002, we have generally credible statistics and reports on human development produced by the Central Statistical Agency (BPS) Papua (BPS Papua 2002). After an overview and analysis of statistical information on education, health, labor force, and migration, this section concludes with a reflection on the main findings of the recent National Human Development Report (NHDR 2004) that uses the Human Development Index (HDI) to reveal and emphasize the achievements of New Order development. In 1961, the total number of people in West New Guinea was estimated at 700,000 but no exact data were available on ‘‘the exploration resorts’’ or uncontrolled areas in the Central Highlands. The provisional estimates reported by the government suggested that some 250,000 were living in montane West New Guinea (Report on Netherlands New Guinea 1961). More recently and using a large variety of sources, Ploeg (2001) estimates the number of people living in the highlands during the early 1960s at 230,000. In 2002, 668,671 people were living in the highlands, of a total of 2,387,427 people for the whole province (BPS Papua 2002: 80). In 1961–1962, the population density in the regions under administrative control was 2 per km2, while in 2002, now also including the slightly more densely populated highlands, it had increased to 5.6 per km2.

education In 1960, the total number of village schools was around 800, most of which were managed by missionary societies and congregations. In the rural areas, only the very simplest three-year education seemed feasible and few Papuans rose above this level. Despite attempts to increase the level of education during the last decade of Dutch rule in New Guinea, the number of pupils in village and urban elementary schools had risen from 26,417 in 1952 to only 40,615 (of which around 43% were girls) in 1961. Most of the 844 teachers at these schools were Papuans and hardly any of them had teaching qualifications. On average, each school had one teacher for 51 pupils in three classes.

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Reflecting the policy that aimed at the emancipation of Papuans and preparing the population for self-government, the number of Papuans enjoying training at the 25 so-called vervolgscholen (‘‘supplementary schools,’’ most of which had boarding establishments) grew significantly in this period, from 804 to 2,734. This training would allow them to enter vocational and technical courses. Most of these courses were run by government departments, both on behalf of the departments themselves and as part of the effort of rapid Papuanization of the labor force. Towards the end of Dutch rule, there were four junior technical schools, one domestic science school, eight training schools for village teachers, and four teachers’ training colleges. In 1960, the number of Papuan students in teacher training courses grew from 95 in 1950 to 453 in 1961, and in technical schools from 70 to 212. Non-vocational training remained undervalued; the number of Papuans in secondary school only grew from 20 in 1950 to 116 in 1958. In 1960, the number of Papuans attending the MULO (advanced primary school) increased to 430. These quick rises in student numbers are indicative of the last-minute efforts of the Dutch to develop the labor force of Netherlands New Guinea. In the final years of Dutch administration, increasing numbers of people were given the chance to study abroad. In 1960, 29 Papuans were studying in the Netherlands: three at university, two at the tertiary-level Tropical Agriculture Institute, and seven at secondary schools, while the remainder were enrolled in college-level vocational and technical courses. In 1961, 50 Papuans went to the Netherlands to follow advanced training courses. In addition, Papuan students were sent to the Australia-administered Territory of Papua and New Guinea to attend medical college or to undertake a radio engineering course. Two Papuan students attended the Auxiliary Medicine and Dentistry School in Fiji (Penders 2002: 390–391). Illiteracy was reportedly low in those regions where children had been attending schools for two or three generations. The situation appeared most favorable in the Cenderawasih Bay region, the northern regions and in Sorong and Manokwari, where illiteracy rates averaged less than 30%. In other regions such as the southern plains and parts of the highlands that were recently brought under administration, the illiteracy rate ranged from 30% to 70% (Report on Netherlands New Guinea 1960, 1961; Penders 2002: 391). The related regionally disproportionate development would continue over the next few decades. In the course of the New Order period, the administration expanded education in Irian Jaya extensively. While during the Dutch period most rural schools were financed and operated by mission organizations, the number of state schools grew markedly from the 1960s until today. Since the beginning of REPELITA II (1974– 1978), through a Presidential Decree (INPRES: Instruksi Presiden), primary schools have been made available in every village and primary education is now compulsory. In terms of accessibility, this has lead to a major increase in attendance at primary education in most regions of Irian Jaya, but for higher education, access has remained starkly divided between rural regions and provincial and district capitals.

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Because of the high levels of poverty, many parents face difficulties in keeping their children at school because of the money needed for school fees, books, uniforms, and school organizations (Rusman 1998: 376). Moreover, the quality of education in Papua is reportedly low compared to other provinces due to the poor quality of teachers and facilities (Rusman 1998: 377). Not surprisingly, older people in Papua cherish nostalgic ideas about the Dutch education system that to a certain extent provided facilities and support for pupils to attend boarding schools. Many policy makers in the Department of Education, including a number of governors, have regularly expressed the need for reviving subsidized boarding in rural regions of the province. The number of state primary schools grew from 1,337 in 1998 to 1,402 in 2002, with a total of 184,325 pupils in 1998 and 232,941 in 2002. The number of teachers in state primary schools was 10,948 in 2002. Private primary schools grew at a similar rate and numbered 1,005 in 2002 with 7,036 teachers for 140,395 pupils. Taking state and private schools together, there were 7.5 teachers per school for 155 pupils in six classes. The number of state junior high schools increased from 217 to 241 over the period from 1998 to 2002, while over the same period the number of private junior high schools increased from 110 to 115. The number of teachers at state junior high schools was 3,414 for 67,279 pupils, while at private junior high schools there were 722 teachers for 21,011 pupils. In 2002, there were 121 state senior high schools for general education and 37 for vocational training. The numbers for private junior high schools and private senior high schools in 2002 were 115 and 77 respectively. The vocational senior high schools attracted 18,168 pupils in 2002 (BPS Papua 2002: 137–159). During the Dutch period, there was no university in West New Guinea but, after the transfer of the territory, the Indonesian government rapidly established Cenderawasih University (UNCEN: Universitas Cenderawasih). Student number at this university has experienced significant growth over the last decade, from 4,136 to 11,163, of whom a large proportion are enrolled in science and social and political studies (BPS Papua 2002). The Faperta or Agricultural Department of UNCEN based in Manokwari attracted 953 students in 2002 and was formalized as a separate institution, the State University of Papua (UNIPA: Universitas Negeri Papua), in 2000. The aim of the educational development effort was partly to make Papuans into Indonesian citizens, in ways similar to earlier Dutch efforts of making ‘‘selfconscious world citizens ready for independence.’’ The New Order administration also recognized the need to develop human resources to stimulate economic growth and, at least nominally, to prepare the population for jobs in the government and commercial sectors. This effort has not been entirely successful due to the lack of attention paid to increasing access to education for rural communities and the failure to stimulate good vocational training rather than university and college level education for Papuans. At all levels of the educational system, curric-

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ula that are mostly developed in Jakarta are poorly adjusted to local Papuan needs and perspectives, which decreases their effectiveness and motivation. As the Dutch had previously observed, Papuan educational achievement shows marked differentiation by sex and residence. The level of education among women is strikingly low, especially in highland regions such as Jayawijaya. In 1990, the national census showed a 30% illiteracy rate for the whole province, with remarkable differences between the urban regions (Jayapura 3.7%, Sorong 4.0%, Fakfak 3.0%, Merauke 3.0%) and the highlands (Jayawijaya 43.2% and Paniai 36.2%) (Rusman 1998: 369). In the age group 25–29, more than 50% of the women from the rural areas are illiterate, a pattern that is also apparent in urban regions and that reflects social and cultural tendencies and the limited success of female access to educational programs. Regarding dissemination of information on health issues, which is becoming ever more pressing with the rampant spread of HIV/AIDS in the region, it is worth noting that the percentage of women reading a newspaper weekly in Papua is 19.1%, compared to 26.9% for the whole of Indonesia (Lautenbach 1999: 60). The adult literacy rate for Papua was the lowest, at a level of 74.4%. Together with an average of only six years of schooling, this places Papua as the second lowest in the nation, following West Nusa Tenggara, where there are far fewer migrants from west Indonesia to boost provincial figures (National Human Development Report 2004: 97). According to the 1990 population census, there were 465,122 people aged between 15 and 29 in Irian Jaya (Papua), approximately 28% of the total population. 68% of youth lived in rural areas while those in the urban areas comprised mainly of children of migrants (Rusman 1998: 366). This wide gap between educational achievement in rural and urban areas persists until today and leads to resentment among Papuans towards the government elite and migrants who make up the majority of students in high schools, colleges, and universities, and whose chances of getting employment are greater.

health The New Order government also put considerable effort into increasing medical services in Irian Jaya (Papua). At rates comparable to the establishment of schools, the number of public clinics and hospitals grew significantly from the 1970s onwards. Many skilled doctors and medical auxiliaries were made available by Jakarta but they frequently faced shortages of medicines and equipment. The montane interior and the southern plains lagged behind in this development largely because poor infrastructure and harsh working conditions deterred doctors and nurses. In some regions, diseases like malaria, yaws, leprosy, respiratory infections, tuberculosis, and venereal diseases spread where the Dutch had previously been able to curtail them through systematic and well-monitored campaigns (Pouwer 1999: 173). The number of hospitals at the end of 1961 was 22. Of these, seven were general hospitals in the urban centers (with numbers of beds ranging from 27 in Fakfak to 360 in Hollandia, now Jayapura) and 15 smaller hospitals (including one pri-

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vate) in the regions (with numbers of beds ranging from 6 to 32). Of a total of 122 outpatient clinics, 108 served rural communities, while there was only one maternity hospital, one mental hospital, and five leper colonies. The number of medical doctors grew from 63 to 73 between 1957 and 1961, while over the same period the number of nurses with diplomas decreased from 231 to 209. The number of midwives remained basically stable at only eight and the training of midwives was started only in 1960, first at Inanwatan, and later in the districts of Sentani and Demta (near Hollandia, now Jayapura). To combat malaria, there were 19 control spraying teams, while for yaws there was one, for tuberculosis one, and for leprosy two control teams (Report on Netherlands New Guinea 1961: Appendices). In 1984, Irian Jaya had the highest rate of health centers per capita of the country, at 47 per 100,000 people (Lautenbach 1999: 54–56). In the early 1970s, a national scheme called the Applied Family Welfare Program (PKK: Pembinaan Kesejahteraan Keluarga) was implemented in Irian Jaya. The program aimed to enhance the role of married women in the development of the nation. Along with a far-reaching standardization of village government during the same period, the family welfare program was implemented through the wives of government employees and village (desa) heads. In its outline and practical implementation, the program confirmed women’s status as subordinate to their male counterparts. The most successful part of the program appeared to be the establishment of a village health service post (posyandu) that is run by volunteers and the scheme for consultation by traveling nurses from the headquarter’s community health center (puskesmas). However, in most rural regions these services are hampered by regular shortages of medicine, unavailability of trained personnel, and poor planning and coordination. During the New Order period, Indonesia followed closely the World Health Organization’s guidelines for child vaccination. In 1977, the Extended Program on Immunization (EPI) was introduced to combat tuberculosis, diphtheria, pertussis, tetanus, polio, and measles through vaccinations of children under one year. Apart from some remote regions in parts of the highlands and the southern plains, the vaccination programs were successful and still run today. Since the 1970s, infant mortality rates for Irian Jaya as a whole dropped significantly, though less rapidly than for the rest of Indonesia (Lautenbach 1999: 102). In 2002, life expectancy for Papua was 65.2, compared to 72.3 and 72.4 for the Jakarta and Yogyakarta regions, respectively.

labor force In mid-1961, 18,986 Papuans were employed, of whom 68% were in the urban centers of the regions of Hollandia (Jayapura), Cenderawasih Bay, Manokwari, Fakfak, and Merauke. Over 7,100 members of the total labor force in these centers were migrants, mostly Eurasians and Moluccans, the majority of whom were young unmarried men, reflecting the limited availability of trained Papuans and a general favoring of migrants over locals. Over 6,000 Papuans were in paid employ-

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ment in the building trade; 1,200 in factories or workshops; 2,300 in modern-style agriculture, stock-breeding, forestry, timber-felling, hunting and fishing; 1,600 in transport and communication; 4,600 in education, administration, public health, police, fire brigade, the hotel business, and so on; and 3,300 in other businesses. Over 10,500 Papuans were in government service and nearly 8,500 worked for private employers. In 1961, the total number of employed Papuan women was 1,345. The Indonesian Government continued the effort to further ‘‘Papuanize’’ the civil service, though this was done rather hesitantly out of fear that Papuans would gain too much of a voice. In addition, like the Dutch, the new government also experienced limitations in the availability of adequately trained personnel. The sensible practice of training Papuans in courses set up by the different government departments was discontinued and, in the course of time, people from other islands of Indonesia took most of the government positions. It was only in the late 1990s that policies were adopted and sincere efforts were made to Papuanize the government. Currently, around 35% of the labor force in the government is Papuan, which is a poor reflection of the demographic reality in which approximately 60% of the population is Papuan (D. Flassy, pers. comm.). For the whole of Irian Jaya, in 1990, 289,854 (62.3%) of youths were classified as ‘‘economically active’’ and 175,268 (37.7%) who were ‘‘not economically active.’’ The majority of the economically active were employed, while among those who were not economically active, half were attending school and the rest were housekeeping or engaged in other activities (Rusman 1998: 368). New economic activities began to mushroom in urban centers like Jayapura, Sorong, Merauke, and Timika (the last largely related to the Freeport mine). In the future, British Petroleum (BP) and the Indonesian oil company Pertamina will boost economic development in Manokwari and Sorong due to the Tangguh natural gas extraction project (Chapter 6.5). The government’s investment schemes have systematically prioritized the urban centers of the northern districts and the proportion of the local indigenous population who benefit from the growth in the cash economy is relatively small compared to other provinces and the neighboring country of Papua New Guinea (Manning and Rumbiak 1989). As a result, urbanization has increased over the last few decades and immigration of people from other islands largely affects the towns, while Papuans in the interior are discouraged from moving to towns, not the least because of limited employment opportunities. Besides their limited schooling, Papuans must compete with immigrants who come to towns in Irian Jaya through family and friendsof-friends networks, while at the same time many non-Papuan employers tend to consider Papuans lazy and unreliable. Throughout the New Order period and up until today, job opportunities for Papuans have become increasingly limited, further discouraging many Papuan parents from investing in their children’s education. In 1990, the proportion of youth from the highlands who had completed senior high school was two to three times lower than among the youth from the coastal and urban regions (Rusman 1998: 372). Thus, despite the impressive in-

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crease in educational attainment, health services, and a significant decrease in illiteracy since 1962, a major problem yet to be tackled is the unequal distribution of access to education and employment by region, and employment preference for migrants over Papuans.

migration Internal migration and migration from outside the province are perhaps the most significant causes of demographic, social, and cultural change in Papua. Papuan communities have always been very mobile and the demographic picture of precolonial Papua is one of flux. For centuries, people living along the coast looked towards the west and maintained trade and marriage relationship with the Moluccas; Islam spread along certain coastal stretches (Swadling 1996; Kamma 1947/ 1948; Visser 1989; Huizinga 1998). Many of these coastal communities currently play prominent roles in all sectors of development in Papua. In the course of the Dutch period, migration towards urban centers increased because of the availability of jobs and the attraction of manufactured goods. Since the Indonesian government began to stimulate economic development in the region, an older Dutch colonial program of population distribution from highly populated regions such as Java and Bali to Papua and other less populated provinces was continued. President Sukarno and also the following New Order Government supported the emigration program called transmigration (transmigrasi). Transmigration came to be seen as a way to boost the development of Irian Jaya, but the program proved to be largely unsuccessful. The total number of transmigrant families that settled in Irian Jaya remained well below the targets set. In 1984, for example, the transmigration program projected an increase of approximately 138,000 families, or 700,000 persons, for the REPELITA IV (1985–1989) period but the number of people who actually arrived in Irian Jaya was 272,320. The program proved to be very costly because of poor infrastructure in the region and the increasing need for subsidies for farmers who found themselves in a situation where demand for crops was low and transportation to markets was expensive. Over time, the transmigration program became a concerted effort to seek economic, social, and security benefits from, first of all, the more rapid spontaneous migration waves (amounting to an estimated 300,000 to 400,000 people in 1985), secondly, the organized transmigration scheme and, thirdly, the resettlement of Papuan communities. The latter aspect of the program was particularly designed to stimulate the economic integration of Papuans and to ‘‘civilize’’ groups that are considered ‘‘backward’’ or ‘‘isolated’’ (masyarakat tertinggal) (Ndoen 1994). The regional focus of the program has been the areas around Merauke, Manokwari, Paniai, and Timika. A revealing report produced by the Irian Jaya Rural Community Development Foundation (YPMD: Yayasan Pengembangan Masyarakat Desa) indicates that myriad problems resulted from poor implementation of the program and lack of coordination between the three responsible ministries: Interior, Forestry, and Social Affairs (YPMD 1985, summarized by Pouwer 1999: 173–174). On the basis of

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visits to five transmigration projects in various parts of Papua, YPMD concluded that the program fails to appreciate local social and cultural dynamics, that it leads to resentment among Papuans who lose their agricultural land and hunting grounds without compensation, that there are serious delays, that the housing, school, and health facilities for the transmigrants are poor, and that access to markets for the transmigrants has not received enough attention. Overall, despite numerous reports suggesting that the transmigration program was a deliberate attempt by the government to ‘‘Javanize’’ the Papuans, the demographic impact of the program was far more muted than most had anticipated. From 1976 to 1986, the peak period of transmigration, 94,000 people settled in transmigration sites, of whom about 13,000 were local transmigrants. During this period, only one quarter of the estimated total increase of the population of Irian Jaya was accounted for by transmigrants from outside Irian Jaya (Manning and Rumbiak 1989). However, as mentioned above, spontaneous migration and urbanization has significantly affected the composition of the population of the urban regions and the distribution of economic and social developments (see Visser 2001). Currently, the influx of migrants from elsewhere in Indonesia seems to be increasing, which possibly increases tensions between non-Papuans and Papuans.

human development There is a consensus among development agencies that development can be measured in terms of people’s choices. The most critical of these choices include living a long and healthy life, being educated, and having access to the resources needed for a decent standard of living. The National Human Development Report (NHDR) that was produced by the United Nations Development Project (UNDP) in collaboration with Central Statistical Agency (BPS) and National Development Planning Agency (BAPPENAS) brought together the production and distribution of commodities and the expansion as well as use of human capabilities. The report analyzed economic growth, trade, employment, political freedom, and cultural values from the perspective of local people. The measure used is the United Nations Human Development Index (HDI) which is a comparative measure of poverty, literacy, education, life expectancy, and other factors for countries worldwide. It is a standard means of measuring well-being. The emerging picture indicates certain general trends in schooling, health, and economic progress, but does not cover other aspects of human development, which will be discussed in the following section. The 2004 National Human Development Report showed that the Human Development Index (HDI) for Indonesia in 2002 was 66, ranging from 76 in the highly urbanized and industrialized region of East Jakarta to 47 in the district of Jayawijaya in Papua (NHDR 2004: 1). The considerable variation across the country was thought to be related to differences in the availability of profitable natural resources, people’s access to these natural resources, and the distribution of the

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revenues of the extraction of these riches. At the national level, the resource-rich province of Papua ranked 26 places lower in HDI than in per capita gross domestic product (GDP), a clear indication that income from Papua’s mining, oil, forestry, and fishing industries have not been invested sufficiently in services for the people (NHDR 2004: 11). An even greater variation was found within Papua, where district HDIs vary from 47 for Jayawijaya to 71.4 and 73 in Jayapura and Sorong, respectively (NHDR 2004: 109). In the statistics for social welfare produced by the Central Bureau of Statistics (BPS) for Papua in 2002, the districts of Jayawijaya, Paniai, and Puncak Jaya showed the highest number of poor families and the greatest number of so-called ‘‘isolated communities’’ (masyarakat tertinggal) (BPS Papua 2002: 207). This clearly reflects limited access to services and indicates the extent to which the government has failed to increase living standards among rural Papuan communities. Highlanders feel that this is an unfair distribution of resources, particularly when they reflect on their dislocation in the light of the economic growth and wealth enjoyed in the urban centers dominated by non-Papuans and Papuans from coastal regions. In addition to the vast revenue from natural resource exploitation not being shared evenly within the country or among different groups within Papua, many non-Papuans are more likely to find employment in the commercial sector. Despite the fact that employment for Papuans in the bureaucracy has been stimulated over the last few years, there are still great disparities. In general, the statistics, together with observations of the situation on the ground, indicate that overall the government of Indonesia has not been successful in fulfilling some basic needs of the people of Papua despite relatively high expenditures on education and health. For most regions, funding for development programs came through the government, with the exception of private investments in urban centers and the Freeport mine location. The benefit of private expenditure tends to be weighted towards the rich, a large proportion of whom are nonPapuan. This has contributed to significant regional divisions in levels of health service and educational attainment. Also, infant mortality rates for Papuans and poor migrants in both urban and rural regions are much higher than they are for the wealthy in Timika, Sorong, Merauke, and Jayapura. In conclusion, the legacy of 32 years of New Order government in Papua is a number of dramatic changes. Most worrisome, statistics indicate that the region falls below the national average in terms of health, education, and infrastructure, despite containing an overwhelming quantity of natural resources (minerals, timber, oil, and gas). Moreover, regional divisions in development, in particular between northern coastal and central highlands communities, have been reinforced by unequal distribution of resources. What is not included in the statistical assessments of human development is what other (informal) changes people in Papua have experienced. The statistics fail to convey how certain age-old traditions and customs, as well as ethnic and religious identities, have been altered, and how social structures have changed as the results of migrations and the uneven distribution of modern opportunities. The next section discusses these issues.

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Resentment and Resistance During the Suharto-led New Order, an ever-growing but relatively poorly funded military supported a network of alliances for both political control and predatory business. Predatory businesses were organized around the exploitation of natural resources taken from rural communities without proper compensation. As noted above, Papua was one of the regions where mining took on central importance. Forestry and fishery activities were initially limited but grew significantly during the period of reform and decentralization after the fall of the Suharto regime in 1998. Recently, the London-based Environmental Investigation Agency (EIA) and Telapak in Jakarta reported on rampant logging in Papua (EIA and Telapak 2005). The report is an alarming indication of illegal trade threatening the last tract of pristine forest in the Asia-Pacific region. The investigation revealed how one timber species, merbau (kayu besi), a luxurious dark hardwood, is the main target of a billion-dollar trade route from Papua to the booming cities of China’s Yangtze River delta. Numerous local legislators and members of the security force are involved. This is just one effect of the poorly guided national decentralization effort that allows many local leaders to gain a larger share of the profits that were effectively channeled to Jakarta elites during the earlier period. They became ‘‘little Suhartos’’ in popular expression. Nepotism, international finance, military muscle, and the siphoning off of funds and revenues by the military at every level of the government became the key ingredients of decentralization in Papua. Involvement of the military in large-scale resource projects such as the Freeport gold and copper mine includes security deals with the company and the establishment of the military as the main bridge between allegedly rebellious local communities and an industry that feels threatened by attacks from Papuans. In the extreme case of Freeport, a condition of systematic intimidation, manipulation, and terror developed which profited the military, victimized local groups such as the Amungme and Kamoro, and did not improve the reputation of the mine, either locally or internationally (see Ballard 2002). These conditions have made many Papuans feel vulnerable to outside forces, in particular the military and the police. In terms of governance, the region is still among the most poorly developed in Indonesia. Frustration about limited access to services sharpens the fault lines between local people and those who have arrived in Papua through transmigration programs or the larger waves of spontaneous migration. Butonese, Buginese, Makassarese, and Javanese immigrants have filled manual labor and small business opportunities. As a result, economic, ethnic, and religious differences play a significant and sometimes alarming role in land and resource politics. Most of what Papuans have received from the government in terms of support for economic development and infrastructure was designed in Jakarta and appeared largely inappropriate for local circumstances once implemented in Papua.

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In line with the policies of the preceding Dutch administration, the government of Indonesia believes that activities such as exchanging cloth among the people of the western Vogelkop, organizing bridewealth ceremonies in the highlands, and implementing large-scale fertility ritual gatherings in the south-coast plains region are hampering ‘‘the native’s’’ integration into an emerging Indonesian state system and economy. Most administrators working in Papua are convinced that they have to manage a process of cultural incorporation of indigenous cultures within an Indonesian administration and economy. They see themselves as being largely responsible for relating new structures, goals, and values of a centralized administration to the different cultures of the Papuan societies. At the same time, the attitudes of both Papuan and non-Papuan government officials are often marked by a sense of not knowing how to develop the region and, in some cases, outright ignorance and apathy. Lack of knowledge and capacity, and negative ideas about the potential of Papuan communities to participate in modern development, persist today. This constitutes one of the main reasons that Papua remains among the least developed regions of Indonesia. Only when people in Papua were allowed to formulate and plan increased sovereignty within the framework of decentralization were local communities and leaders accorded a meaningful place in the governance of the region. The need for bridging Indonesian institutions and regulations that are founded on nationwide centralized principles of New Order-guided democracy, became a central concern during the drafting of the Special Autonomy Law (see below). The main issues that unite Papuans in a feeling of resentment towards the government of Indonesia are frustrations about the limited successes of development, seemingly systematic marginalization, and ongoing repression. However ‘‘Indonesian’’ many Papuans may behave, talk, and occupy positions in the formal sector, there are clear signs that Indonesian nationalism is very limited, if not nonexistent, among the majority of Papuans. In part, the sentiments of being different and of suffering are expressed in terms of local ‘‘traditional’’ cultural ideas and practices, millennial expectations, and concerns with knowledge. The responses from Papuan communities to a highly centralized administration that fits poorly with the reality on the ground and distributes resources unequally, are varied. In the southwestern Vogelkop region, for example, Imyan people see their autonomy threatened by a variety of demands from the government (e.g., for participation in development programs) and from the church (e.g., demanding that people be good Christians, adhere to the Ten Commandments, attend services, and contribute money and service to the church organization). These demands conflict with traditional duties, such as gardening, hunting, organizing bride wealth payments, and settling disputes (see Timmer 2000). The duties to the government and church are viewed as in conflict with ‘‘traditional’’ duties in terms of demands on time and labor. In contrast, ‘‘tradition’’ or ‘‘custom’’ (adat) is perceived as stable and comprising all contemporary reflections on and practices that people regard as customary. In many of the locations where large-scale resource extraction takes place, such

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as the Freeport copper and gold mine in the Timika region, the recently initiated British Petroleum/Pertamina Tangguh LNG plant in the Bintuni Bay, and logging and fishery businesses, local communities organize themselves against neighboring local groups that also claim the natural resources and demand compensation. One of the effects of this development is emotionally charged revitalizations of customary structures and the establishment of traditional communities (masyarakat adat). Because of the expectations of monetary flows that resource development projects might bring, there are often competing claims over land and resources. This poses a problem not only for local communities that no longer know whom to trust and through whom to raise their voices with outside companies and the government, but also for the government and the companies that find it increasingly difficult to deal effectively with the dispersed Papuan civil society. Besides tradition or custom, Christianity has left a deep imprint on the culture of Papuan communities. Missionization began in 1855 and continues through today in remote regions of the southern plains and the highlands. Two major church organizations have established a firm role in the lives of Papuans: Catholics, mostly in the south and southwest, and Protestants, mainly in the north and northwest (see Kamma 1976; Cornelissen 1988; Boelaars 1992, 1995, 1997; van Nunen 1999), and charismatic evangelicals in the highlands. As a result, local interpretations of Christianity form an important background for the social and political changes taking place in Papua today. Christian symbols and rituals appear in a creative combination with pre-Christian cosmologies, and rituals often become weapons of the weak, functioning as ways to subtly undermine the nation and its ideologies (Timmer 2004). Religious tensions with immigrant communities at times induce Papuans to confirm or stress that they are Christian, in a nation whose majority is Muslim. At the same time, the church is the most prominent and largest well-functioning civil organization, and moreover, it belongs largely to the Papuans. On top of that, church organizations are present in most remote areas where the government and non-governmental organizations generally fail to deliver services.

Special Autonomy For many Papuans, the idea of self-rule and autonomy dates back to the events during the final years of Dutch colonial rule. As mentioned above, in 1955, the Netherlands Government laid the foundations for the institution of a central representative body, the New Guinea Council, which was installed on 5 April 1961. In this representative body, most of whose members (23 out of 28) were Papuans, future policies were worked out in cooperation with representatives of the population, who were elected by majority vote in the regions of Manokwari and Hollandia (Jayapura), while those from other regions were indirectly elected or appointed. The New Guinea Council was granted a co-legislative function, including the right of initiative, the right of amendment, and the right of interpellation,

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and collaboration in drafting the budget. These rights, including political representation for the population, were viewed as practical preparation for the application of the principle of self-determination. Despite the brief time the New Guinea Council had to fully prepare Papuans for governing their own land, it certainly had a catalytic effect on the political awakening of certain elements of Papuan society. However, very few Papuans participated in the New Guinea Council process or emulated the ideas of governance and self-rule advocated by the Dutch. The Indonesian New Order policy minimized autonomy for the Papuans while formally recognizing limited autonomy for the province of Irian Jaya. The New Guinea Council was replaced by a Provincial Council (DPRD: Dewan Perwakilan Rakyat Daerah) but the regulations and practices of governance made the Governor more responsible to the President than to the population of the province. The council was not granted any budgetary rights and the Provincial Development Planning Bureau (BAPPEDA: Badan Perencanaan Pembangunan Daerah) was expected to implement development programs that were designed by the National Development Planning Agency (BAPPENAS: Badan Perencanaan Pembangunan Nasional) in Jakarta. The district and subdistrict heads were directly responsible to their superiors so consultation with local groups and any institutionalization of the right of initiative or control over planning and expenditure became virtually extinct. To counter these unsettling imbalances in governance, a number of people in Papua supported a Special Autonomy package that had been offered by Jakarta in late 2001, in an explicit attempt to curtail the demands for independence advocated by elements of Papua’s society. The new laws were intended to give more autonomy to local communities in terms of equal sharing of the benefits of resource extraction and democratic governance. The draft legislation drew heavily on a draft prepared by the governor of Papua and a team of Papua-based academics that worked for him. But the draft was not really presented to the community and there was no provision for any dialogue in the future leading towards a referendum or a negotiated resolution of the conflict between the government and Papuan communities (see Sumule 2003). Financially, however generous the provisions appeared to be (80% of resource revenues and 70% of oil and gas, for instance), it is becoming increasingly apparent that a lot of that money stays in the hands of a few people. Another serious problem with the Special Autonomy package is that its implementation is difficult because the province is very poorly served in terms of trained human resources; it is not in a position to maximize the benefits from the new revenue. Overall, the promises and the few benefits of Special Autonomy so far only serve to frustrate many Papuans. More and more people are motivated to disagree, sometimes violently, with those who are seen as a menace: ‘‘the Indonesians,’’ ‘‘the immigrants,’’ ‘‘the transmigrants,’’ the big companies, the military, and those Papuans who are seen to have become ‘‘too Indonesian.’’ On top of that, the positive developments that were expected to grow from Special Autonomy were frustrated by a decree on 27 January 2003. The decree

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expedited the implementation of Law No. 45/1999, creating two new provinces (West Irian Jaya and Central Irian Jaya), three new districts (Paniai, Mimika, and Puncak Jaya), and one municipality (Sorong). The decree, which has become labeled pemekaran after the Indonesian word blossoming but referring to ‘‘administrative involution,’’ took most Papuan leaders by surprise. Their surprise was soon followed by distress when they learned that the initiative for the policy had come from quarters within the central government that feared that Special Autonomy would give too much leverage to independence-minded Papuans. Reactions to the accelerated implementation of pemekaran were diverse but generally negative. A spate of popular reactions held that it was yet another attempt by Jakarta to divide and rule the Papuans so as to control the region’s resources. Others, in particular intellectuals, activists, and politicians supportive of Special Autonomy, began to argue that Law No. 45/1999 dividing Papua, is inconsistent with the later Special Autonomy Law and therefore, on first legal principles, must be repealed to the extent of its inconsistency with the Special Autonomy Law (International Crisis Group 2003; Sullivan 2003). In November 2004, the Constitutional Court judge ruled against a lawsuit from the Special Autonomy Defense Team. This Defense Team argued that Law No. 45/ 1999, implementing division of Papua, was orchestrated by the Megawati Sukarnoputri-led Indonesian Democratic Party of Struggle (PDI-P) to weaken the Golkar Party that was dominant in Papua. The Defense Team also argued that the PDI-P had economic interests in Bintuni Bay, where British Petroleum and Pertamina are establishing the Tangguh natural gas plant. The Special Autonomy Defense Team added that both the military and the National Intelligence Board (BIN) would also lose out if Law No. 45/1999 were dropped, because the two institutions have economic interests in maintaining a large military presence in Papua. The Constitutional Court concluded that the establishment of West Irian Jaya remained valid, although Law No. 45/1999 was no longer effective. Eight of the nine judges argued that the Special Autonomy Law took effect after the new province and districts were formed, and that no state institution had been annulled by the Special Autonomy Law. The court ruling was seen as a victory for West Irian Jaya’s acting governor, and left many people in Jayapura, Sorong, and elsewhere in Papua, confused. For West Irian Jaya, the verdict requires a review of the province’s status, specifically the application of Special Autonomy. The pemekaran decree implementing the division of Papua was issued without consultation with local communities, the provincial government, or leaders of religious and other civil society organizations in Papua. The decree started cleavages in the Papuan elite, some of whom supported Special Autonomy while others were keen on establishing separate provinces and claiming their own governorships. The resulting disunity among influential people in the Papuan bureaucratic elite has weakened the leverage of supporters of Special Autonomy who oppose the implementation of the pemekaran decree (McGibbon 2004; Timmer 2005). For those who support division of Papua, the central issues are access to the Spe-

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cial Autonomy Funds and the share of fiscal transfers from the central government that will be disbursed to the new provinces, as well as control over the territory’s riches, and increases in their own personal status.

Conclusion Although the province of Papua is resource rich, its people are among the nation’s poorest. Educational opportunities are limited and those with training face high unemployment rates in an environment that favors employees from other regions of Indonesia. Papuan parents target government jobs if they are seeking a prosperous future for their children because competition in the commercial sector is too stiff. The government sector is also not immune to discrimination against Papuans, perhaps in combination with a fear of people who are inclined to develop a separatist form of nationalism gaining too much influence in the formal sector. In that sense, any attempt at involving more Papuans in decision-making processes, and employing more Papuans in the formal sector, creates a vicious circle. A generally low level of education, with a curriculum focused on making proper Indonesian citizens out of unruly rebellious Papuans, provides Papuans with limited alternative strategies for creating forms of nationalism and connecting to advocacy bodies within Indonesia and abroad. Recent generations of Papuans have, however, seized opportunities for education, especially those from regions where people were less disgruntled by the Indonesian ‘‘takeover’’ and were favored in terms of access to education, such as the Vogelkop region. The acquired skills and knowledge of the current Papuan bureaucrats enables several influential people to establish links with Jakarta for the benefit of the people in Papua, but these talents are mostly used for self-interest in competition among elements of the elite. In contrast, the relatively poorly developed highland and south coastal regions have bureaucracies with a more regional representation, despite a disproportionate number of policy makers from Biak-Numfoor and Yapen (especially the town of Serui) in the Cenderawasih Bay, the Sorong region of the Vogelkop, and Sentani, and the inclusion of a few Javanese and Moluccan representatives. In particular, the recently ascended Sorong and Ayamaru elites are familiar with Indonesian ways of doing politics and, in line with most elements of the elite, have a growing appreciation for engaging in political and economic strategies that primarily benefit themselves. This produces dishonesty and resentment among others in the bureaucracy and the communities that they administer. Highlanders and people from the south coastal regions are often consumed with envy about the power enjoyed by people from the Cenderawasih Bay, the Vogelkop, and Sentani. Many of the current internal differences in Papua and the lingering conflict with Jakarta relate to shifts in power at the center of the state and increased concerns over access to resources as a result of decentralization at the regional level. The gap is widening between Papuan perspectives and a nationalist perspective that advocates an apparently organized state structure but that often disguises

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nepotism, business, and international economic interests. The provincial government, civil society organizations, and local communities have a long way to go to reach a shared commitment to increasing access to services, particularly for rural communities, to initiating social, economic, and political changes that will serve the poor, and to promoting a more equal sharing of resource benefits through Special Autonomy regulations.

Acknowledgments Thanks to Chris Ballard, Mike Cookson, Chris Penders, Anton Ploeg, Leontine Visser, and Dirk Vlasblom for helpful comments on earlier drafts of this chapter.

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1122 / j a a p t i m m e r Glazebrook, D. 2001. Dwelling in exile, perceiving return: West Papuan refugees from Irian Jaya living in East Awin in Western Province, Papua New Guinea. Ph.D. thesis, The Australian National University. Groenewegen, K., and D.J. van de Kaa. 1964. Resultaten van het Demografisch Onderzoek Westelijk Nieuw-Guinea. 4 volumes. Government Printing and Publishing Office, The Hague. Huizinga, F. 1998. Relations between Tidore and the North Coast of New Guinea in the Nineteenth Century. Pp. 385–419 in Miedema, J., C. Ode´, and R.A.C. Dam (eds.) Perspectives on the Bird’s Head of Irian Jaya, Indonesia: Proceedings of the Conference, Leiden, 13–17 October 1997. Editions Rodopi, Amsterdam. Huydecoper van Nigtevecht, J.L.R. 1990. Nieuw-Guinea: Het Einde van een Koloniaal Beleid. SDU, The Hague. International Crisis Group. 2003. Dividing Papua: how not to do it. Indonesia Briefing 9 April 2003. International Crisis Group, Jakarta and Brussels. Kamma, F.C. 1947/1948. De Verhouding tussen Tidore en de Papoesche Eilanden in Legende en Historie. Indonesie¨ I, 361ff.; II, 536 ff. Kamma, F.C. 1976. Dit Wonderlijke Werk: Het Probleem van de Communicatie tussen Oost en West Gebaseerd op de Ervaringen in het Zendingswerk op Nieuw-Guinea (Irian Jaya) 1855–1972, Een Socio-missiologische Benadering. 2 volumes. Raad voor de Zending der Ned. Hervormde Kerk, Oegstgeest. Klein, W.C. (ed.). 1935–1938. Nieuw Guinee. Three volumes. De Bussy and MolukkenInstituut, Amsterdam. Lagerberg, C.S.I.J. 1962. Jaren van Reconstructie: Nieuw-Guinea van 1949 tot 1961. Ph.D. thesis, University of Utrecht. Zuid-Nederlandse Drukkerij, ’s-Hertogenbosch. Lautenbach, H. 1999. Demographic survey research in Irian Jaya: population dynamics in the Teminabuan area of the Bird’s Head Peninsula of Irian Jaya, Indonesia. Ph.D. thesis, University of Groningen. Thela Thesis, Amsterdam. Leith, D. 2003. The Politics of Power: Freeport in Suharto’s Indonesia. Honolulu: University of Hawai’i Press. Lijphart, A. 1966. The Trauma of Decolonization: The Dutch and West New Guinea. Yale University Press, New Haven, Connecticut. Manning, C., and M. Rumbiak. 1989. Irian Jaya: economic change, migrants, and indigenous welfare. Pp. 77–106 in Hill, H. (ed.) Unity and Diversity: Regional Economic Development in Indonesia since 1970. Oxford University Press, Oxford. McGibbon, R. 2004. Secessionist Challenges in Aceh and Papua: Is Special Autonomy the Solution? Policy Studies No. 10. East-West Center, Washington, D.C. Moore, C. 2003. New Guinea: Crossing Boundaries and History. University of Hawai’i Press, Honolulu. National Human Development Report. 2004. Indonesia National Human Development Report 2004: The Economics of Democracy, Financing Human Development in Indonesia. BPS-Statistics Indonesia, BAPPENAS, and UNDP Indonesia, Jakarta. Ndoen, M.L. 1994. Ekonomi politik migrasi di Irian Jaya. Bina Darma 44: 55–65. Penders, C.L.M. 2002. The West New Guinea Debacle: Dutch Decolonisation and Indonesia, 1945–1962. Crawford House, Adelaide. Pepera. 1972. Penentuan Pendapat Rakjat (Pepera) di Irian Barat 1969, pp. 82–83. Pemerintah Daerah Propinsi Irian Barat, Djajapura. Ploeg, A. 2001. The other Western Highlands. Social Anthropology 9 (1): 25–43. Poulgrain, G. 1999. Delaying the ‘discovery’ of oil in West New Guinea. The Journal of Pacific History 34 (2): 205–218.

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A Brief Social and Political History of Papua, 1962–2005 / 1123 Pouwer, J. 1999. The colonisation, decolonisation and recolonisation of West New Guinea. The Journal of Pacific History 34 (2): 157–179. Report on Netherlands New Guinea 1960. Ministry of Home Affairs and Ministry of Foreign Affairs, The Hague. Report on Netherlands New Guinea 1961. Ministry of Home Affairs and Ministry of Foreign Affairs, The Hague. Rusman, R. 1998. Youth, education and employment in Irian Jaya. Pp. 365–381 in Miedema, J., C. Ode´, and R.A.C. Dam (eds.) Perspectives on the Bird’s Head of Irian Jaya, Indonesia: Proceedings of the Conference, Leiden, 13–17 October 1997. Editions Rodopi, Amsterdam. Rutherford, D. 2003. Raiding the Land of the Foreigners: The Limits of the Nation on an Indonesian Frontier. Princeton University Press, Princeton, New Jersey. Schoorl, J.W. (ed.). 1996. Besturen in Nederlands-Nieuw-Guinea 1945–1962: Ontwikkelingswerk in een Periode van Politieke Onrust. KITLV Press, Leiden. Soekarno. 2000. Bebaskan Irian Barat: Kumpulan Pidato Presiden Soekarno tentang Pembebasan Irian Barat, 17 Agustus 1961–17 Agustus 1962. Ragam Media, Yogyakarta. Souter, G. 1963. New Guinea: The Last Unknown. Angus and Robertson, Sydney. Sullivan, L. 2003. Challenges to Special Autonomy in the Province of Papua, Republic of Indonesia. Discussion paper 2003/6. State, Society and Governance in Melanesia Project, Research School of Pacific and Asian Studies, The Australian National University. Sumule, A. 2003. Swimming against the current: the drafting of the Special Autonomy bill for the Province of Papua and its passage through the national parliament of Indonesia. Journal of Pacific History 38 (3): 353–369. Swadling, P. 1996. Plumes from Paradise: Trade Cycles in Outer Southeast Asia and Their Impact on New Guinea and Nearby Islands until 1920. Papua New Guinea National Museum, Boroko. Timmer, J. 2000. Living with intricate futures: order and confusion in Imyan worlds, Irian Jaya, Indonesia. Ph.D. thesis, Centre for Pacific and Asian Studies, University of Nijmegen, Nijmegen. Timmer, J. 2004. Government, church, and millenarian critique in the Imyan tradition of the religious (Papua/Irian Jaya, Indonesia). Pp. 117–136 in Jebens, Holger (ed.) Cargo, Cult and Culture Critique. University of Hawai’i Press, Honolulu. Timmer, J. 2005. Decentralisation and elite politics in Papua. Discussion paper 2005/6. State, Society and Governance in Melanesia Project, Research School of Pacific and Asian Studies, The Australian National University. UNDP/FUNDWI. 1975. Fund of the United Nations for the Development of West-Irian (FUNDWI)–Final Report. UNDP/FUNDWI, Jayapura. van Galen, J.J. 1984. Ons laatste oorlogje–Nieuw-Guinea, de Pax Neerlandica, de Diplomatieke Kruistocht en de Vervlogen Droom van een Papoea-natie. Van Holkema en Warendorf, Weesp. van Nunen, B.O. 1999. Enam Puluh Tahun Penginjilan Fransiskan di Irian Jaya 1937– 1997. Unpublished, Diocese of Jayapura, Jayapura. van Oosterhout, D. 2002. Landscapes of the body: reproduction, fertility and morality in a Papuan society. Ph.D. thesis. Research School CNWS, University of Leiden, Leiden. Visser, L. 1989. The Kamrau Bay area: between Mimika and Maluku, a report of a short visit. Bulletin of Irian Jaya, Irian 17: 65–76. Visser, L 2001. Remaining poor on natural riches? The fallacy of community development in Irian Jaya/Papua. The Asia Pacific Journal of Anthropology 2 (2): 68–88.

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1124 / j a a p t i m m e r Vlasblom, D. 2004. Papoea: Een Geschiedenis. Mets & Schilt, Amsterdam. Whittaker, J.L., N.G. Gash, J.F. Hookey, and R.C. Lacey. 1975. Documents and Readings in New Guinea History, Prehistory to 1889. Jacaranda Press, Milton, Queensland. Wilson, F. 1981. The Conquest of the Copper Mountain. Atheneum, New York. YPMD. 1985. Special issue on resettlement of Kabar dari Kampung, No. 3–4. YPMD, Jayuapura.

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6.3. The Agricultural Systems of Papua manuel boissie` re and yohanes purwanto e o pl e f r om P a p ua , even if they depend on forest products for their livelihoods, mostly rely on cultivated crops for their subsistence. Strictly hunter-gatherer societies do not exist in Papua. Most of Papua’s ca 300 ethnic groups cultivate a variety of crops, and hunt and gather as supplemental activities. Much could be said about the different types of agriculture in the western part of the New Guinea island, as gardening techniques can differ from one valley to another. Many techniques are used, from irrigated taro cultivation in insular systems, to sago gardens in the lowlands, to mixed crop cultivation and taro gardens in the highlands. Our understanding of agricultural practices in the western part of New Guinea, however, is limited, in part because very few scientists are allowed to work there. Studies in the eastern part of New Guinea (i.e., Papua New Guinea) are extensive, including, for example, the works of Bourke et al. (1998) on agricultural systems, and also work by Panoff (1969), Clarke (1971), Sillitoe (1983, 1996, 1998), Waddell (1972), Dwyer (1990), Lemonnier (1982), Juillerat (1982), Lory (1982), Malinowski (1935), Steensberg (1980), and Yen (1993), among others. Some monographs on agriculture in Papua are available, but there is no synthesis of the overall system. Pospisil (1963), Cook (1995), Purwanto (1997), Sunarto and Rumawas (1997), Boissie`re (1999), Ploeg (2000), and others provide in-depth studies of specific kinds of agriculture in Papua, mostly related to a particular ethnic group. Despite the small number of publications, some comparisons can be made between agriculture in Papua and in Papua New Guinea. For the same altitudinal level, techniques and crops used by local people from both countries are very similar; this is not surprising considering the arbitrary drawing of the national borders, and the similarities of the environment and populations on either side of the border. Because the authors cannot give an exhaustive overview of all the agricultural systems in Papua, we shall instead describe the diversity of the systems that have already been studied and that represent the principal ones in Papua. In this chapter, after a historical background of agriculture in Papua, we shall propose a set of definitions for the different terms describing traditional agriculture systems, and then present a classification of the main agricultural systems in Papua. Then we shall turn to describing each type of agriculture, according to the landscape in which it is used. The chapter concludes with a discussion of the recent transformation of agriculture, systems of cash cropping, and new priorities in terms of food security.

P

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

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Historical Background: The Origins of Agriculture in Papua For the most populated places in New Guinea, the highlands and coastal areas, the landscape is heavily impacted by human activities (Chapter 6.1). Forest landscapes are mixed with agricultural landscapes (Kennedy and Clarke 2004), as shifting cultivation coexists with forest (Cook 1995). The question of the arrival of agriculture in New Guinea has aroused a lot of controversy, as has the question of whether New Guinea was a center for plant domestication and the independent development of agriculture (Golson 1989; Yen 1973, 1993; Gillieson et al. 1985; Powell 1970). According to Denham et al. (2003: 189), ‘‘agriculture arose independently in New Guinea by at least 6,950 to 6,440 calibrated years before the present (cal yr bp).’’ Most evidence comes from research at the Kuk site in the Wahgi Valley, Papua New Guinea. For the western part of the island, anthropogenic impacts have been recorded from the Baliem Valley from 7,800 cal yr bp, but the first sign of human activity, according to carbon analysis, was estimated at 28,000 bp (Haberle et al. 1991), probably a result of bush fires. Recent research (Neumann 2003: 181) suggests that one of the centers for agricultural origins is New Guinea, and that ‘‘early forms of cultivation may have started 10,000 years ago, mounds are attested 7,000 to 6,400 years ago, and ditch network was built 4,400 to 4,000 years ago.’’ Mounding introduced a new era of intensification for agricultural practices; ecological characteristics of the mounds buffer the micro-temperature of the soil in which crops are planted, allowing a longer period of garden activities before fallow time, and the possibility of planting crops at higher altitudes (Waddell 1972). Cultivation occurred first in the wetlands around 10,000 bp, and developed independently in the highlands. This period (6,400 bp) was also the time for domestication of Eumusa bananas, an essential crop for New Guinea (Denham et al. 2003: 192). According to Powell (1970), at around 2,600 bp, swamp lands were used intermittently for agriculture, in combination with lands at higher altitude; these swamp gardens presumably involved a complex system of drainage, especially compared to shifting cultivation.

Definitions of Traditional Agricultural Systems Concerning the use of the terms ‘‘horticulture’’ and ‘‘agriculture,’’ which are part of the common vocabulary concerning agro-systems, we follow Kennedy and Clarke (2004: 5), who make slight distinctions between them: ‘‘the term horticulture in Oceania is often used as a synonym of agriculture. We use agriculture as an inclusive cover term for productive activities, of which the other terms describe specific components.’’ Kennedy and Clarke (2004: 7) consider arboriculture—tree cultivation—in New Guinea to be fully part of the agricultural systems. According to them ‘‘in the New Guinea region, the integration of different modes of cultivation of mainly

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perennial plants in complex polycultural production systems justifies the designation of these systems as agricultural.’’ Traditional agricultural systems are characterized by the farmer’s dependence on local resources and technologies. In order to optimize food production in these low-input farming systems, farmers must possess considerable knowledge both of the nature and characteristics of the resources available, and of the methods suitable for sustainable crop production under difficult conditions. Traditional agriculture is thought to support the needs of more than onequarter of the world’s population (Wolf 1986), and it is in a fragile condition, because of the ever increasing degradation of agriculture habitats. Although encompassing a wide range of distinct horticultural systems, all traditional farming methods can be distinguished from industrial or ‘‘green revolution’’ agriculture by a number of characteristic features, among which are their low reliance on the use of agrochemicals and commercial seed, and their total dependence on local climate (Cotton 1996). Traditional farmers, therefore, try to optimize the use of local resources to minimize the risk of crop failure and to ensure the sustainability of their production methods. A wide range of agricultural systems are employed in the traditional farming societies of Papua, ranging from the shifting cultivation of the highlands to permanent wetland agriculture of coastal areas. Many of these farming communities have remained dependent on nutritional supplements from a wide range of wild foods, and display a considerable knowledge of the nature, behavior, and distribution of the useful species available. A variety of agricultural systems are used in combination, in order to spread risk where crop failure is common due to factors such as drought, pests, and disease. The success of these systems often relies on the careful selection of crop varieties well-suited to specific micro-environments, and on the use of cropping patterns which maximize resource use and minimize susceptibility to damage (Richards 1985). Complementing their knowledge of the environment, cultivators generally show considerable knowledge of many domesticated crops. The plant species exploited in traditional systems include true domesticates or ‘‘cultigens’’ (domesticated crops), which depend on human stewardship to complete their life cycle; semi-domesticated plants, which may be propagated and managed but are not obligatorily reliant on human intervention; and protected plants, which even when occurring spontaneously in managed plots are encouraged or protected by practices such as weeding and pruning. Traditional farmers spread risk both by cultivating a number of different crop species and by exploiting several cultivars, or varieties of important staples. For example, the Dani in the Baliem Valley have more than 100 distinct cultivars of sweet potato Ipomoea batatas (Purwanto 1997; Figure 6.3.1). These varieties vary in a range of characteristics such as flavor, size, storage properties, time of maturation, and resistance to pests and disease. These features allow them both to fulfill a range of nutritional and cultural requirements and to ensure some productivity under often unpredictable circumstances. Agricultural practices have a great impact on landscape composition. For exam-

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Figure 6.3.1. Cluster of sweet potato gardens in Tiom area (Baliem Valley). Photo: M. Boissie`re.

THE PL ACE OF PIGS IN PAPUAN AGRICULTURE Pig husbandry is intimately connected to Papuan agricultural systems because a part of the garden’s production is reserved for feeding pigs. Every day, when men and women leave the cultivated plot, they bring home some sweet potato tubers that are too hard for human consumption and some sweet potato leaves for their pigs. When they arrive at the village, women cook them together and give them to the pigs. In some societies (e.g., the Yali), cassava is a food for pigs and humans never eat it, except if there has been a very bad harvest and no other food is available. Pigs play an important role in Papuan societies, even though introduction by Austronesian migrants was relatively recent (about 3,600 years bp, according to Diamond 1997). They are an important part of local cosmologies and are critical for religious rituals, payment of compensation, wedding festivals (bride price), and exchanges between friends and relatives (Brutti and Boissie`re 2002; Lemonnier 1993). Pigs can even be part of agricultural practices themselves: when the gardens are abandoned, pigs are often left free to roam and gather any remaining tubers and greens. In the Chimbu society of Papua New Guinea it has been reported that pigs are used to turn over the soil by foraging in the fallows, to prepare and fertilize the ground for new plantings (Brookfield and Brown 1963).

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ple, the presence of fences transforms a garden’s landscape; in places where gardens are fenced, pigs are left free to wander around the village and in the forest. When no fences protect the gardens, pigs must be confined in fenced areas and cannot move freely around the village. This has implications for the state of the adjacent forest. Forests visited by pigs have little understory, and large pioneer trees (e.g., Albizzia sp. or Caldcluvia sp.) grow well and become dominant. Tools also influence the kind of agriculture that is practiced. In Papua, gardeners mostly use machetes, axes, and digging sticks. Chainsaws are still rarely used. As a consequence, gardening is very laborious and a lot of physical energy must be expended for it to be successful.

agricultural classifications in the literature The classification of agricultural systems differs according to different authors. After describing several alternatives, and we will propose our system, based on general topography, crops, techniques, and our own observations during our research in Papua. Matanubun et al. (1995) define agricultural systems according to the techniques used for gardening. Thus ‘‘agricultural systems in Papua can be divided into two types, shifting and settled cultivation. In the Baliem Valley and Wissel Lakes areas, shifting cultivation is practiced on the hill slopes and foothill areas, whereas settled cultivation is performed on valley floors and river banks. In the Anggi Lakes area, only the shifting cultivation system is practiced, both on mountain slopes and in the flat area’’ (Matanubun et al. 1995: 58). According to Kennedy and Clarke (2004), shifting cultivation can be considered an adaptation to low soil fertility, whereas settled cultivation, with its complex farming techniques, is an adaptation to strong population pressure and can only be practiced in areas of relatively high soil fertility (e.g., in the Baliem Valley). Another general classification of agricultural systems is provided by Haynes (1989), in the framework of a report for the United Nations Development Program (UNDP). Haynes defined two major systems made up of six subsystems, although the practices described occasionally appear very similar between different subsystems. The first major system, lowland agriculture, comprises three subsystems: (1) coastal swamps and rivers: sago cultivation with occasional tuber gardens; (2) coastal plains: coconuts orchards with swidden cultivation (cassava, taro, and yam); and (3) in foothills and small valleys: shifting cultivation (tubers). The second major system, highland agriculture, also has three subsystems: (1) in broad valleys: intensive sweet potato production; (2) in lakeshore plains: sweet potato gardens, coffee cash cropping; and (3) on hill slopes in narrow valleys: sweet potato gardens. Barrau (1962) describes seven main types of subsistence economies in New Guinea distributed in four main types of landscapes. First, in the lowlands are found both a) hunting and/or fishing, associated with gathering and shifting sago cultivation with long periods of fallow; and b) shifting cultivation of tubers, sago, bananas, and Saccharum sp. associated with gathering, hunting, and/or fishing. Second, in foothills and low mountains is found shifting cultivation with long

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fallow periods of tubers, bananas, and Saccharum sp., associated with gathering, hunting, and/or fishing. Third, in the mountains are found both a) semi-sedentary cultivation of tubers and herbs, with fallow periods, composting, and ditch networks; and b) cultivation of tubers and herbs based on long fallows, associated with gathering, hunting, and/or fishing. In the fourth landscape type, coastal mountains, are found the last two subsistence economies. In swamps, gathering of fruits and wild nuts and cultivation with long fallow periods of sago and herbs is associated with fishing and/or hunting. In rainforests, cultivation with long fallow periods of tubers, sago, bananas, and Saccharum sp. is associated with gathering and fishing and/or hunting.

proposed agriculture classification For our purposes, we will keep the main landscape divisions: highland and lowland, and we will add one component to the last type, the insular landscape. Our division of agricultural systems integrates the main ideas of the previous authors, and tries to show, by the description of a small number of case studies, the great diversity present in these three types of agriculture: highland, lowland, and coastal/ insular.

Highland Agriculture Highland agriculture is the best documented type of agriculture in Papua (Gardiner 1987). After a brief discussion of some general features, we will present four examples of highland agricultures from four different societies of the highlands. The highlands are the most densely populated areas in Papua. Places like Enarotali or the Baliem Valley account for a considerable portion of Papua’s inhabitants. According to a 1997 census (available at http://irja.bps.go.id/), about 22% of the Papuan population lives in the Jayawijaya Regency, which includes the Baliem Valley; it is therefore the most populous regency in Papua. Agricultural practices from the highlands represent one of the most important sets of techniques in Papua (Figure 6.3.1). Highlanders cultivate tuber crops, mostly sweet potatoes (Ipomoea batatas), cassava (Manihot esculentus), yams (Dioscorea sp.), and taro (Colocasia esculenta). But many other crops are planted, such as bananas (Musa sp.), red pandanus (Pandanus conoideus), nut pandanus (Pandanus julianettii), and a considerable number of vegetables, which complement the nutritional intake from tubers. Agriculture in the highlands can be roughly divided in two groups: hill agriculture (between 600 m and 1,500 m altitude) and mountain agriculture (above 1,500 m altitude). In both, sweet potato remains the staple crop planted, but different additional crops can be found. Most kinds of agriculture in New Guinea conform to a cycle of events (Figure 6.3.2), called ‘‘garden sequences.’’ These are a succession of stages that lead to the final garden: clearing, cleaning, tree felling, fencing, preparation of the soil, planting, weeding, harvesting, and leaving fallow. For each of these stages, a number of

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Figure 6.3.2. Cycles of cultivation. Source: Boissie`re (2003).

variations can be observed, each of which requires certain specific techniques to implement. The diversity of techniques creates a diversity of garden types. In a single society, different kinds of gardens can be observed, each adapted to specific conditions (topography, soil fertility, availability of land, particular crop, etc.). Most of the steps described in Figure 6.3.2 can be found in most highland gardens; only a few techniques or knowledge differ across garden types. Thus highland gardens can be considered, not only as an object of cultivation, topographically well-delimited, but also as a set of practices and actions on landscape, which can be seen as a dynamic cycle. All of these practices and actions come from the heritage of skills among members of each society, and from their capacity for assimilation of new techniques (Haudricourt 1987), as we will discuss further below. Here we describe examples of agricultural practices that come from four societies of the Papua highlands: the Yali, Dani, Kapauku, and Amungme. These four societies live in the same sort of habitat, in the Central Cordillera of Papua, and have developed many similar behaviors related to agriculture, as well as a great diversity of practices. Located in a vast territory east of the Baliem Gorge, the Yali represent a society of agriculturalists of both the low mountains and highlands (see Boissie`re 1999a,b, 2003; Figure 6.3.3.). The Yali recognize five main types of garden, according to the techniques used prior to planting (Figure 6.3.4). These techniques are listed in Table 6.3.1. Yali gardens end after two years of cultivation, even in the case of mound gardens. But as seen in Figure 6.3.2, the active phase of the garden is only

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Figure 6.3.3. Sweet potato gardens in Holuwon. Photo: M. Boissie`re.

part of a long 15–20-year cycle, with human intervention at each step of the cycle. During the fallow period of a garden, the Yali often go to that location to hunt, to look for non-timber forest products (NTFPs), or simply to collect remaining crops, such as Pandanus. Yali agriculturalists use an incredible variety of techniques and also incorporate techniques from neighbors (e.g., from the Dani, who inhabit the Baliem Valley). The traditional agricultural system practiced by the West Dani and Dani-Baliem society is shifting cultivation of sweet potatoes (Purwanto 1997, 2003; Soenarto and Rumawas 1997). Rotation to a new location depends on the condition of the land, but generally occurs after every two or three harvests. The selection of the location of a new field by shifting cultivators is based on the condition of the vegetation that grows on it. When the vegetation regrowth looks vigorous, the cultivator assumes that soil fertility has recovered and that the land can be cultivated again. The stages of establishing a sweet potato garden are: discussions regarding site selection, slashing, felling, burning, fence construction, tilling, planting, protection (including weeding), and harvesting. The establishment of a sweet potato garden is accompanied by traditional ceremonies. Sweet potato cultivation techniques of the West Dani and Dani-Baliem in the central mountains of Jayawijaya are adapted to prevailing environmental conditions, especially with respect to the land, topography, slope, and other aspects of the environment, such as microclimate. For each environmental condition Dani

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Figure 6.3.4. Woman planting sweet potatoes in Holuwon. Photo: M. Boissie`re.

society has different techniques for sweet potato cultivation. Basically there are three different techniques, depending on whether sweet potatoes are grown in swamps, in well-drained soil on flat land, or on slopes. For sweet potato gardens grown in a swamp or other inundated site (Figure 6.3.5), a deep furrow (1.5–2 m deep and 1.5–3 m long) is dug to make the mound, reduce ground water level, and to capture nutrients from the decomposition of various grasses (weeds), which are thrown into the furrow. The Dani use the mud formed in the furrow as fertilizer. Analysis of nutrient composition shows that the mud contains more nitrogen and potassium than the soil in the garden bed. This technique also offers the advantage of stabilizing soil temperature and soil moisture; sometimes the furrow may even be used as a fishpond. For sweet potato gardens grown in well-drained soils on flat land, the constructed furrow is not deep, but a mound is made as a place to plant each individual sweet potato cutting. Mound-building regulates soil temperature and soil moisture for optimal growth of sweet potatoes. For sweet potato gardens grown on slopes, the garden bed is made by a deep furrow. A main drainage channel is made in the middle and on the sides of the garden. This drainage channel is

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Table 6.3.1. The different types of Yali gardens Garden name

Practice

Location, topography

Esap yabuk or Wen yabuk

Herb gardens or mound gardens. The soil is turned over to build mounds, and fertilizer is used. Sweet potato is the main crop.

Gentle slopes

Kwenang yabuk

Garden where the soil is turned over, without use of fertilizer or mounds. Ground nuts and sweet potatoes are planted here.

Gentle slopes

Wealangge

Gardens directly planted after clearing. The soil is not turned over to avoid landslide.

Steep slopes

Soli yabuk

Garden where the soil is turned over onto fertilizer, without mounding, onto large beds. Rare.

Gentle slopes and flat lands

Busuk yabuk

Garden partly prepared as an esap yabuk, and partly as a wealangge. Rare.

Places that are partly steep and partly flat; adaptated to the topography.

designed to protect the garden from excessive water, which could carry away the soil. According to the Dani, if there is good drainage, sweet potato tubers are sweeter in this kind of garden. Generally, in sweet potato gardens, vegetables and other crops are planted too, such as Setaria palmifolia, Colocasia esculenta, Psophocarpus tetragonolobus, Brassica oleracea var. botrytis, Brassica oleracea var. capitata, Phaseolus vulgaris, Zea mays, Dioscorea alata, Dioscorea esculenta, Saccharum officinarum, Musa spp., Pandanus conoideus, P. julianettii, and Nicotiana tabaccum. Similarities can be found between the Kapauku agricultural system and that of the Dani. The Kapauku, or Etoro, according to Pospisil (1963), have two types of agriculture: one adapted to steep slopes and one to valley bottoms. Pospisil (1963) considers shifting cultivation on mountain slopes as an extensive cultivation method. The grower selects a site, according to land tenure (land is owned individually) and to the length of time for fallow (between 7 and 12 years), and clears the vegetation by felling the trees and burning the remaining vegetation. Fences are built on these gardens to protect them from pigs (domesticated or wild). Very few crops can be planted: only sweet potatoes and some greens (e.g., Amaranthaceae). On the slopes, no fertilizer is used, and after a weeding process the garden is only harvested once before the long fallow period. Cultivation on the valley floor is considered intensive by Pospisil (1963). On

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Figure 6.3.5. Sweet potato gardens along the Baliem River. Photo: Y. Purwanto/doc LIPI.

the valley bottom, the black soil is deep enough to allow the use of drainage ditches, composting, and crop rotation. Aside from sweet potatoes, several crops can be successively planted in this sort of garden: manioc, taro, sugar cane, bananas, bottle gourds, cucumbers, and so on. Although the soil is rich, this land can be subject to flooding when there are heavy rains. The size of gardens here is smaller than on mountain slopes. The valley bottom technique has two features: cultivation of a small surface, with crop rotation; and no soil preparation other than the use of fire for clearing. This garden is very similar to that on the mountain

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slopes, and only a single crop is cultivated there (e.g., sugar cane, sweet potatoes, or taro). If ditches are built, then similarities may be found with the Dani, such as in the use of large drainage ditches in valley swamps. Lastly, the Kapauku differentiate another category of cultivation on the valley bottom, called Bedamai, an intensive complex cultivation, where ‘‘the soil itself is cared for by an intensive drainage system, the use of compost fertilizer, and spading in order to form raised beds for the planted crops’’ (Pospisil 1963: 122). Soil is turned over onto a layer of fertilizer to form numerous beds. The Amungme practice shifting cultivation with the use of fire heaps, and each garden is re-opened after a fallow of at least five years (Cook 1995). Shrubs are cleared and then trees cut down; some trees are left standing. Cleared debris is left for a month to dry, then debris is burned in small cribs, and the rest of the garden’s surface is also burned. Once this is completed, mounds 20 cm high are made with digging sticks. Most crops are planted together in these mounds at the same time. Five categories of gardens can be identified: house gardens, sweet potato gardens, riverside gardens, high altitude taro gardens, and Pandanus orchards. House gardens are small gardens planted in each neighborhood with traditional and nontraditional plants; these gardens are cultivated more intensively than the other garden types. House gardens are a source of cuttings, food, and a place to experiment with new crops. Sweet potato gardens represent the main cultivated area in Amungme territory, and are planted year-round. Sweet potatoes are planted in mounds. These gardens are mixed-crop cultivation, and dozens of different plants can be found there (sugar cane, hibiscus, ferns, Amaranthus, spinach, pumpkin, cucumber, bananas, etc.). Riverside gardens contain crops adapted to humid areas, such as Xanthosoma, but also gourds (Lagenaria siceraria) may be grown there. High altitude taro gardens contain taro (Colocasia esculenta), locally known as ‘‘Mo,’’ a favorite source of carbohydrate for the Amungme. These gardens are reserved for men. Less attention is given to these types of gardens, where fewer crops other than taro are grown (compared to sweet potato gardens): only some maize, beans, sugar cane, and legumes are typically found there. However, many cultivars of taro can be grown in the same garden. Finally, Amungme cultivate Pandanus orchards. Pandanus spp. are generally planted when a garden is opened, but remain in small orchards long after other cultures have been abandoned. Fallow land is used for a long time, for harvesting some remaining food crops and for non-timber forest products.

Lowland Agriculture The lowlands can be divided in three categories, according to landscape and season: swamps, foothills and low mountains, and savanna. The first two categories are largely represented in the Mamberamo Watershed, one of the biggest watersheds in Papua; the third is found only in the vicinity of Merauke, in the southeast part of Papua. Very few studies have been conducted on lowland agricultural systems (Tucker

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1987). However, two main agricultural practices in the lowland can be identified: one is linked to sago cultivation (sago is an important crop for the lowlanders), and the second is swidden cultivation of tuber crops (e.g., sweet potato, manioc, yam), wherein banana is a key element. In the Mamberamo Watershed, sago is the main food, and as a consequence, land tenure seems to be stricter for sago orchards than for other kinds of gardens (CIFOR 2004). Swidden gardens are considered a secondary source for food, and gardeners can open these anywhere, but sago gardens are subject to a formal system of land use and tenure. The following anecdote illustrates lowlanders’ indifference to swidden agriculture: in Kwerba, some villagers opened a garden, cleaned the land, prepared the soil, and once it was planted, they just abandoned it, because, according to them, they didn’t want to spend more energy to take care of it. Each sago garden belongs to a villager and, for example, in Kwerba, a spiny cultivar of sago is used to delimit personal orchards. Sago is planted in swampy places or at the bottom of small valleys (Schuiling and Jong 1996). This crop can be planted in orchards, but is also found in the forest, growing wild or escaped from former orchards. Sago is mostly cultivated from the suckers of year-old plants (Schuiling and Jong 1996). All leaves except the youngest ones are removed. The suckers are planted in shady places, with the apical meristem above water level. After three months, the shading plants can be removed and the sago must be weeded regularly. Usually, sago is worked by women alone or communally (and the harvest is divided among all participants). This is the case in Mamberamo, but also in many other places (Townsend 1992). The organization of labor is divided among the women: while some scrape the stems, others filter the starch, and others pack it in bags made from sago leaves (Figure 6.3.6). In the Mamberamo Watershed, tuber cultivation is normally secondary, and sago is the primary crop, often mixed with banana cultivation. Yam, cassava, and sweet potato are not staples and are cultivated only to add variety to villagers’ diet. Bananas are usually planted in mixed-crop gardens and, when the garden is harvested and left for a fallow period, only the bananas remain to form orchards where the villagers regularly come to harvest the fruits. Usually coconuts can be found in the lowlands, in the vicinity of the villages, and these are mainly used for their fruits, for household consumption. According to Haynes (1989), this kind of agriculture must also be supplemented by hunting, fishing, and gathering.

Coastal and Insular Agriculture In the coastal areas and islands (e.g., the Vogelkop region; see Haynes 1989), shifting cultivation takes place with taro and yam as staple foods supplemented by gardens of mixed crops (e.g., cassava, banana, sweet potato). Recently, plantations of oil palm and cocoa have been developed in the coastal areas and lowlands; this kind of commercial cultivation will be discussed in the following section. Taro can be found growing in a wide range of altitudes, from sea level to 2,700 m

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Figure 6.3.6. Woman processing sago starch in Mamberamo. Photo: M. Boissie`re.

in New Guinea, where it has developed resistance to cold. Taro in insular (island) systems is cultivated with special techniques, such as irrigated gardens. Two examples of people with coastal and island agriculture systems are the Ormu of the Cyclops Mountains (near Sentani) and the Kimam of Kolepom Island (near Merauke). The Ormu, who live in coastal and insular areas, can be divided into two groups: land (keret darat) and sea groups (keret laut). The coastal and insular societies have different subsistence activities: land-based activities include gardening, gathering, and hunting, and sea-based activities include fishing. Although most families practice agricultural activities, differences between the land groups and sea groups of Ormu lie in the intensity and the predominance of one activity over others. For the sea Ormu (keret laut), catching fish is the principal daily activity; agriculture is a side activity. By contrast, for the land group (keret darat), fishing is secondary and agriculture the main activity. As in other traditional societies in Papua, coastal agricultural activities here follow a fallow system (Purwanto 2004). The main cultivated crops are: (1) tubercules, including taro (Colocasia esculenta, Ipomoea batatas, Manihot esculenta, and Dioscorea spp.), sago (Metroxylon sago and M. rumphii), and other crops like corn (Zea mays), rice (Oryza sativa), and some leguminous crops (Arachis hypogaea,

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Vigna sinensis, Glycine max, etc.); (2) vegetables (Psophocarpus tetragonolobus, Vigna sinensis, Amaranthus spp., Capsicum annum and C. frutescens, Lycopersicon esculentum, Solanum melongena, etc.), and (3) fruits (Mangifera indica, Pometia pinnata, Annona muricata and A. squamosa, Averhoa bilimbi and Av. carambola, Anacardium occcidentale, Cucurbita moschata, Lansium domesticum, etc.). Crops for daily consumption are cultivated in gardens around the village and in house gardens. Some of the important economic species (mostly for earning cash) are cloves (Eugenia aromatica), Psidium guajava, Anacardium occidentale, Averrhoa carambola, Av. bilimbi, Citrus spp., Mangifera indica, Carica papaya, Nephelium lappaceum, Artocarpus heterophyllus, Artocarpus champedem, Artocarpus integra, Lansium domesticum, Cocos nucifera, Areca catechu, Durio zibethinus, and Persea americana. Gathering sago is an important activity for coastal societies. Two different varieties of sago trees are distinguished by the Ormu, based on the presence or absence of thorns. Originally these two taxa were considered to be separate species (Metroxylon sagu and M. rumphii), but most botanists now consider them to be two varieties of the species M. sagu. Sago extraction is conducted on 8- to 15-year-old, 10–15 m tall (diameter above 50 cm dbh) trees. Besides by stature, sago trees ready to be harvested can be recognized by new frond leaves being shorter than previous ones, and fronds beginning to expand and conical inflorescence buds starting to appear distally on the tree. When the inflorescence is starting to form branches (looking like deer horns), then the sago tree must be cut immediately. Also, the thorns of mature sago trees are shorter than those of younger trees. An interesting example of specific strategies of coastal and insular agricultural systems is the Kimam ethnic group that lives on Kolepom Island (formerly Frederik-Hendrik Island), close to the Papuan coast. A characteristic of Kimam agriculture is that it is not intensive, as opposed to, for example, that the agricultural system found in the Baliem Valley. Serpenti (1965) explains that root crops are the staple, and sago is the most important supplementary food in Kolepom Island. Crops are cultivated in swamp areas, on artificial garden-islands. Serpenti (1965) reports that ‘‘sago and coconut areas [are] surrounded by a circle of islands especially devoted to the cultivation of root-crops.’’ The first step in the opening of a root crop garden is choice of suitable soil; if the soil is not sufficiently rich, then sometimes compost is added. The main crops in these mixed gardens are yams, taro, cassava, and sweet potatoes. The Kimam build new islands for their gardens during the dry season when the water level is low. Places with a high concentration of reeds are selected as locations for the island gardens, and the reeds are cut at the water level, in patches two to three meters wide. A ditch is made on both sides of the island in the clay, and the island constructed from the clay in the surrounding water. The island is consolidated with a thick layer of drift grass, and then further with clay, to raise the level of the island. Construction of these islands can take several dry seasons. Usually only a strip of less than one meter square is planted the year that the island is built. Old islands need to be

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regularly rebuilt as the clay dissolves in the water. This laborious work is done communally, with the help of friends and relatives. Unlike most other societies of Papua, sago starch collection is mainly the work of Kimam men. Sago is planted on low beds, close to each other. Only the pith necessary for immediate use is collected; the rest of the trunk is stored in water. Women then prepare the sago flour. Coconut is planted on artificial islands, as are other crops; these islands are often not far from the habitations. Last but not least, bananas are planted between other crops, and play an important role in the Kimam diet.

Recent Transformation Transformations in the techniques used for cultivation have taken place, including the use of chain saws and circulation among ethnic groups of techniques such as mound building or irrigation systems (see Sumule 1994, for the Arfaks). Yet the greatest change comes from the diversification of crops cultivated. The appearance of new crops, such as ground nuts (peanuts), lettuce, and cabbage, introduces more variety into agricultural systems, but also makes gardeners more dependant on the local market economy, and affects gardeners’ choices of which crops to grow and which land to use. The shift from an economy based on household subsistence to an economy based on market demand can be observed. This is already common in Papua New Guinea (Boissie`re 2003), and is becoming increasingly widespread in Papua. Local markets and markets at the district and regional levels are supplied by the yields from villagers’ gardens. Migrants from Java or Sulawesi compete with indigenous producers. This is obviously the case in Timika, for example, where agricultural products come primarily from transmigrants’ gardens. The market of Timika is mostly ‘‘occupied’’ by transmigrants, and cultivators from Amungme or Kamoro ethnic groups are relegated to the edges of the marketplace. The cohabitation of local indigenous people in Papua with transmigrants and spontaneous migrants has considerably altered traditional agricultural systems, and has forced traditional groups to adapt to new realities (cultural, but also economic) and to new techniques. Confusion and hesitation about their future can sometimes be observed among many Papuan societies, where cultural pressures, increasing population, creation of economic needs, settlement of migrants, and new technologies oblige them to make choices concerning their way of life, including their agricultural practices. Many of these societies succeed in accommodating their traditional agriculture to new circumstances, and others try to adopt the new models completely, although not always with success. Factors affecting Papuan societies include systems of land tenure, social transformations, and cash cropping. As regards land tenure, the value of the land has changed in regional, national, and international contexts, because of increasing demand and land speculation, but also with respect to local perceptions. Land value had been based on social

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organization, religion, and land quality for agriculture, but has changed to value based on economic and geographical criteria. This difference in perception is frequently the cause of seemingly endless conflicts of interest between the government, private companies, and customary rights of local societies. It has also become more difficult to balance natural resources with local needs. The increasing settlement of migrants (Fearnside 1997) and the introduction of new technologies that allow cultivation of new crops, cause an increasing pressure on the environment (Sumule 1994). The increasing exploitation of natural resources by companies (e.g., PT Freeport Indonesia has extended its mining activities deep into the Amungme territory, causing damage to forests and rivers; see Ballard 2001) often leads to irreversible damage to the environment, and a decrease in available resources and agricultural land. Pollution by mining companies even causes populations to be displaced to new areas where it is often difficult for them to continue their traditional agricultural practices. Many social changes have also occurred. Currently, half of the population of Papua is not indigenous, which has led to many changes in agricultural practices (cultivation of new crops, such as rice), but also to changes in the fabric of social organization. Traditionally, Papuan societies live in small groups based on relations of blood or marriage, and strong relationships are thus built among groups and villages. Town living, with its more heterogeneous population, has changed indigenous societies, especially with respect to the practices and relationships of their members. Acculturation is occurring and it is affecting all aspects of indigenous societies, including their livelihoods (McGibbon 2004). Cash cropping is a major change in the Papuan economy, with wide-ranging effects. As mentioned earlier, changes in Papua societies have led to changes in agricultural systems, the techniques used, the crops planted, and the destination of produce. From a subsistence type of agriculture, a more cash crop-oriented agriculture has emerged on the island. Cultivators willing to grow newly introduced crops for cash have had to adopt new techniques to cultivate them in order to obtain yields that will provide adequate incomes. Some experiments have been made with coffee plantations (Coffea arabica and Coffea robusta), cacao (Theobroma cacao), cloves (Eugenia aromatica), coconuts (Cocos nucifera) and walnuts (Aleurites moluccana). In some cases, agroforestry techniques have even been used for more efficient cultivation. Even so, production remains very low and transportation costs limit most products to local or regional markets. The Dani of the Baliem Valley provide a case in point. Coffee was introduced into this area about thirty years ago but its cultivation was only really developed about fifteen years ago, under the auspices of an Indonesian Institute of Sciences (LIPI: Lembaga Ilmu Pengetahuan Indonesia) initiative and the Agriculture Division of the Jayawijaya Regency. Local people do not profit from the coffee plantations because the coffee market in the Baliem Valley is controlled by the migrants. Moreover, migrants who have enough capital are starting to purchase the coffee plantations belonging to the Dani. Dani society has also developed paddy field cultivation in the vicinity of Wa-

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mena. The aim of the rice field development program was not to change Dani traditional agriculture, but to decrease rice importation into the Baliem Valley and to exploit swamp areas that were not suited to sweet potato cultivation (Purwanto et al. 1990; Purwanto 1997). Yield for the Toraja rice cultivar in the Baliem Valley is about three tons per hectare. Even if rice cultivation has been a success, local people haven’t adopted it as a staple food. The Dani eat only a small portion of the yield and sell the rest. Other cash crops cultivated by the Dani are fruits and vegetables. Traditionally, the Dani cultivate sweet potatoes and local vegetables. New crops, such as broccoli, mustard, potatoes, and carrots, have only recently been integrated into this system. Now vegetable production in the Baliem Valley represents almost half of all the vegetable production in Papua. In general, produce is transported by plane to Jayapura, Timika, and Biak. According to statistics of Papua Province (Badan Perencanaan dan Pengendalian Pembangunan Daerah 2002), vegetable production from Jayawijaya (Baliem Valley) was 12,327 tons per year of cabbage, 3,735 each of shallot and garlic, 6,623 of potatoes, and 3,523 of tomatoes. For societies living in the lowlands, such as the Ormu, Moi, Sentani, and others in the Jayapura Regency, the main crops cultivated for cash are coffee (Coffea spp.), nutmeg (Myristica fragrans), cloves (Eugenia aromatica), coconut (Cocos nucifera), cashew nuts (Anacardium occidentale), sago (Metroxylon sago), and cacao (Theobroma cacao). Jayapura Regency is the biggest producer of cacao and coconut for all of Papua. Other commercial crops are fruits such as rambutan (Nephelium lappaceum), durian (Durio zibethinus), mangos (Mangifera indica), oranges (Citrus sp.), and pineapples (Ananas comosus). The high diversity of fruits cultivated for cash is the direct result of their introduction by migrants. The same situation is observable in Merauke, Fakfak, and Manokwari, where crop plantations have an important role for household cash earning; about 40% of the yield is used for household consumption. In coastal areas, coconut production is also important for cash generation (for copra production, see BPS Statistics of Papua Province 2002). Even so, local people rarely produce copra, or grow oil palm, sugar palm, or vine palm; plantations of these crops are generally owned by companies. Sago, however, is likely to be processed by local people and sold to local entrepreneurs and middlemen, or exported within Indonesia. Among new crop plantations being developed by the government, oil palm (Elaeis guineensis) is one of the most important. In 1997, only 19,250 ha were allocated to oil palm production, with a yield of about 51,242 tons of oil (Directorate General Plantation Estates 1998, in Casson 1999). These plantations are managed by companies or by the local government, and indigenous people rarely benefit from them. In Papua, many companies apply for plantation concessions essentially to gain access to the valuable timber growing on the concession lands, which they cut and sell, and oil palm is never planted (Casson 1999). The introduction of new species has had repercussions socially, culturally, and economically. The high yields from rice fields at Tulem (Baliem Valley) changed

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the status of the villagers; this ‘‘success story’’ boosted villagers’ position both socially and economically, as they became important rice producers in the valley. Effects can also be observed at the cultural level: traditionally in Dani society, men cannot plant seeds, weed gardens, or harvest. These tasks fall to women, except for soil preparation and construction of fences. With the introduction of new crops and techniques, this division of labor has changed, and there is now greater participation of men in these heretofore restricted activities. Even sweet potato cultivation techniques have changed with the appearance of cash crops. Previously, sweet potatoes were cultivated in monoculture (with only various cultivars of other sweet potatoes and some vegetables). Now, sweet potato cultivation is done in a mixed cropping system, with a higher diversity of plants cultivated, among them cash crops. The appearance of these new cash crops grown has influenced the way the local population faces economic pressure and market forces. In theory, the economy of these local societies should improve with the introduction of new crops, but this is not the case because migrants control all tools for cultivation, seeds, and the market. The migrants’ monopoly of the means and market does not favor indigenous people’s bargaining power for the selling of their products. Moreover, local populations are vulnerable to economic crises. Much needs to be done in terms of support to local communities and in providing a ‘‘level’’ economic playing field for local producers.

Food Security and the New Priorities After the El Nin˜o Southern Oscillation (ENSO) event of 1997–1998 and its effects on Papuan society (Ballard 2000; Boissie`re 2002), food security has become an important issue in the region for governmental and nongovernmental agencies. At that time, droughts and fires affected most of the Asia-Pacific region, and isolated groups living in New Guinea received some attention. What was a rare event in the Asia-Pacific region as a whole is, in fact, a fairly regular event in Papua, and indigenous people for generations have adopted complex strategies that reduce variability in food supply. As previously mentioned, traditionally, highland societies such as the Dani, Yali, Nduga, Hupla, Kenyam, Amungme, and others rely on sweet potatoes for their livelihood. In contrast, societies living in coastal and lowland areas depend on sago extraction, gardening, and gathering. Food security is therefore perceived differently by highlanders and lowlanders. Typically, highlanders try to keep gardens continually active to assure food throughout the year. They generally have at least two or three gardens, planted at different times, so that when the production of the first garden decreases, they can shift to another. For that reason, food shortage is rare in the highlands, except during natural disasters (e.g., drought, pest invasion). Diversification of garden crops is another strategy to reduce risks of food shortage or attack by parasites (e.g., fungi, bacteria). Planting a diverse array of cultivars strengthens the capacity of sweet potatoes as a crop to resist disease.

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Polyculture is more likely to provide food security than monoculture (Morren and Hyndman 1987; Paiki 1996), even if in many cases, monoculture still exists in New Guinea. In a Dani garden of four to five hectares in the Baliem Valley, more than 60 local cultivars of sweet potato, 5–12 local cultivars of taro, two or three local cultivars of cassava, and possibly two cultivars of maize may be found (Purwanto 1997). Polyculture also helps mitigate damage caused by poor harvests, and improves the variety and quality of food. This strategy is also used by lowlanders, who cultivate gardens with various crops such as dry rice, maize, peanut, cassava, taro, yam, and a variety of vegetables and fruits. Harvesting gardens in the highlands requires techniques that also play a role in the strategy to insure food security. The tubers of sweet potatoes are not all harvested at the same time within a garden; only the largest tubers are harvested and the small ones are left until they become bigger. This harvesting method ensures a continual stock of tubers. Hence, gardens can be considered as places of food storage. Another strategy for increasing food security is the domestication of wild plants for food. These plants are collected when people go into the forest, and then are planted in the gardens, small orchards, or close to habitations. Several pandans (e.g., Pandanus conoideus, P. brosimus, and P. julianettii) are treated this way. Another strategy to ensure food security is to adopt and develop new agricultural systems adapted to Papuan environmental conditions. Local gardeners are always eager to add new crops to their gardens; gardeners are willing to experiment with new cultivars, even if they keep the more traditional ones (e.g., sweet potatoes) as a staple. This curiosity of gardeners often leads to the learning of new technical skills, but sometimes they simply grow new crops using their usual set of practices (see Boissie`re 1999b for a Yali example). Extension of cultivated land is one response to population growth, even if it also causes many problems with traditional land tenure, land availability, and, possibly most importantly, changes in the local landscape as a result of conversion of forest to agricultural land. In some other places (e.g., the Baliem Valley) one technique is to open gardens on slopes, terracing the land to grow sweet potatoes.

Conclusion Although little research has been conducted on the agricultural systems of Papua, we have tried in this chapter to give an overview of the principal studies using a number of examples taken from existing monographs and the authors’ previous studies. A wide range of agricultural practices are used in Papua. This diversity of practices can be observed in several procedures, including: the many steps necessary to open and cultivate a garden; the diversity of crops (species, varieties, cultivars) planted; the techniques used, that are adapted to each crop, to the topography of the cultivated area, and to the social organization of the gardeners; the way the cultivators manage the cycles of cultivation (e.g., active and fallow phases); and the situation of each society, either isolated (‘‘stand alone’’) or reliant on markets (cash earning) for its economy. Confronted by economic and social

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change, agriculture in Papua can survive precisely because of its diversity and the flexibility of the gardeners, who are able to adopt new techniques and crops, or integrate them into more traditional practices. The main danger for the sustainability of these ancient practices comes from the disruption of the demographic balance between indigenous communities and migrants from other Indonesian islands. The new settlers bring with them new crops and techniques, and are able to appropriate existing markets, leaving little space for local traditional agriculture. Access to land is a key issue for the development of agriculture; the status of land is unclear and too often the rights to land are in the hands of the local government, private companies, or migrants. Even if decentralization has begun to give land rights to local communities, only isolated areas have been completely secured for traditional agriculture. The greatest threat is to the forest: population growth means that more land must be converted to agriculture, and forests are giving way not only to gardens, but also to plantations (e.g., sago, oil palm). The dynamics of agriculture in Papua’s evolving landscape need to be recognized, as well as the rights of indigenous people to their land and how traditional activities can form the basis for longterm economic stability.

Acknowledgments: We are grateful to Chris Ballard for valuable comments and to Dawn Frame for editing the English version.

Literature Cited Badan Perencanaan dan Pengendalian Pembangunan Daerah, Badan Pusat Statistik, 2002. Statistics of Papua Province, 2002. Papua in Figures, Jayapura. Ballard, C. 2000. Condemned to repeat history? ENSO-related drought and famines in Irian Jaya, Indonesia, 1997–1998. Pp. 123–148 in Grove, R. H., and J. Chappell (eds.) El Nin˜o—History and Crisis: Studies from the Asia-Pacific Region. The White Horse Press, Cambridge. Ballard C. 2001. Human rights and the mining sector in Indonesia: a baseline study. IIED, WBCSD, Mining, Minerals and Sustainable Development, 18. Barrau, J. 1962. Les plantes alimentaires de l’Oce´anie: origines, distribution et usages. Annales du Muse´e Colonial de Marseille, 7th series, volumes 3–9 (1955–1961). Faculte´ des Sciences de Marseille, Marseille. [Publication of his thesis, completed in 1962 in the Faculte´ des Sciences, Marseille] Boissie`re, M. 1999a. Ethnobiologie et rapports a` l’environnement des Yali d’Irian Jaya (Indone´sie). Ph.D. diss., Universite´ de Montpellier II. Boissie`re, M. 1999b. La patate douce et l’arachide. Transformations d’une agriculture Yali (Irian Jaya, Indone´sie). JATBA 41 (1): 131–156. Boissie`re, M. 2002. The impact of drought and humanitarian aid on a Yali village in West Papua. Asia Pacific Viewpoint, 43 (3): 293–309. Boissie`re, M. 2003. La me´moire des jardins: pratiques agricoles et transformations sociales en Nouvelle-Guine´e. Annales de la Fondation Fyssen 18: 111–128

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1146 / manuel boissie` re and yohanes purwanto Bourke, R.M., B.J. Allen, P. Hobsbawn, and J. Conway. 1998. Agricultural systems of Papua New Guinea. Working paper No.1, Papua New Guinea, text summaries, vol. 1 and 2. Australian National University, Papua New Guinea Department of Agriculture and Livestock, and University of Papua New Guinea. Brookfield, H.C., and P. Brown. 1963. Struggle for Land. Cambridge University Press, Cambridge. Brutti, L., and M. Boissie`re. 2002. Le donneur, le receveur et la sage femme. Echanges de porcs a` Oksapmin (PNG). Journal de la Socie´te´ des Oce´anistes 114/115: 141–157. Casson, A. 1999. The hesitant boom: Indonesia’s oil palm sub-sector in an era of economic crisis and political change. CIFOR, Bogor. CIFOR. 2004. Building capacity for multidisciplinary landscape assessment in Papua. Three phases of training and pilot assessments in the Mamberamo Basin. CIFOR, Bogor. Clarke, W.C. 1971. Place and People: An Ecology of a New Guinea Community. University of California Press, Berkeley, California, and Australian National University Press, Canberra. Cook, C.D.T. 1995. The Amung way: the subsistence strategies, the knowledge and the dilemma of the Tsinga Valley people in Irian Jaya, Indonesia. Ph.D. diss., Southern Illinois University at Carbondale, Illinois. Cotton, C.M. 1996. Ethnobotany: Principles and Applications. John Wiley & Sons, New York. 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 (11): 189–193. Diamond, J. 1997. Guns, Germs and Steel. W.W. Norton & Company, New York. Dwyer, P.D. 1990. The Pigs That Ate the Garden: A Human Ecology from Papua New Guinea. The University of Michigan Press, Ann Arbor, Michigan. Fearnside, P.M. 1997. Transmigration in Indonesia: lessons from its environmental and social impacts. Environmental Management 21 (4): 553–570. Gardiner, S. 1987. Highland Horticultural Development, Irian Jaya. Jayapura: BPPDUNDP/IBRD Project. Library of Congress microfiche. Gillieson, D., P. Gorecki, and G. Hope. 1985. Prehistoric agricultural systems in a lowland swamp, Papua New Guinea. Archaeology in Oceania 20 (1): 32–37. Golson, J. 1977. 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. Golson, J. 1989. The origins and development of New Guinea agriculture. Pp. 678–687 in Harris, D.R., and G.C. Hillman (eds.) Foraging and Farming: The Evolution of Plant Exploitation. Unwin Hyman, London. 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 (1): 25–40. Haudricourt, A.-G. 1987 (1965). L’origine des techniques. Pp. 287–298 in La technologie science humaine. Recherches d’histoire et d’ethnologie des techniques. Ed. of the Maison des Sciences de l’Homme, Paris. Haynes, P. 1989. Agriculture, soil and climate in Irian Jaya. Irian 17: 88–105. Juillerat, B. 1982. Note sur les rapports de production dans l’horticulture-arboriculture Yafar (Nouvelle-Guine´e). JATBA 29 (3/4): 285–293. Kennedy, J., and W. Clarke, 2004. Cultivated landscapes of the Southwest Pacific. RMAP Working Papers, No. 50, Canberra.

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The Agricultural Systems of Papua / 1147 Lemonnier, P., 1982. Les jardins Anga (Nouvelle-Guine´e). JATBA 29 (3/4): 227–245. Lemonnier, P. 1993. Le porc comme substitut de vie: formes de compensation et e´changes en Nouvelle-Guine´e. Social Anthropology 1: 33–55. Lory, J.-L. 1982. Les jardins Baruya. JATBA 29 (3/4): 247–274. Malinowski, B. 1935. Coral Gardens and Their Magic: A Study of the Methods of Tilling the Soil and of Agricultural Rites in the Trobriand Islands. 2 vol. George Allen & Unwin, London. Matanubun, H., A. Rochani, and A. Sumule. 1995. Some aspects of the indigenous knowledge of selected sweet potato farming systems in Irian Jaya. Pp. 57–62 in Schneider, Ju¨rg (ed.) Indigenous Knowledge in Conservation of Crop Genetic Resources, Proceedings of an International Workshop Held in Cisarua, Bogor, Indonesia January 30–February 3, 1995. CIP-ESEAP/CRIFC, Bogor. McGibbon, R. 2004. Plural society in peril: migration, economic change, and the Papua conflict. Policy Studies 13, East-West Center, Washington, D.C. Morren, G.E.B., Jr, and D.C. Hyndman. 1987. The taro monoculture of central New Guinea. Human Ecology 15 (3): 301–315. Neumann, K. 2003. New Guinea: a cradle of agriculture. Science 301 (11): 180–181. Paiki, F.A. 1996. Symptoms of taro leaf blight disease (Phytophtora colocasiae) and relationship with yield components in Biak, Irian Jaya. Science in New Guinea 21 (3): 153–157. Panoff, F. 1969. Some facets of Maenge horticulture. Oceania 40 (1): 20–31. Ploeg, A. 2000. Dr. P.J. Eyma’s writings on agriculture in the Paniai area, Central Highlands, Western New Guinea. Journal of the Polynesian Society 109 (4): 401–420. Pospisil, L. 1963. Kapauku Papuan Economy. Yale University Publications in Anthropology No. 67. Department of Anthropology, Yale University, New Haven, Connecticut. Powell, J. M. 1970. The history of agriculture in the New Guinea highlands. Search 1 (5): 199–200. Purwanto, Y. 1997. Gestion de la biodiversite´: relations aux plantes et dynamiques ve´ge´tales chez les Dani de la valle´e de la Baliem en Irian Jaya, Indone´sie. 2 vols. Ph.D. diss., Universite´ Paris VI. Purwanto, Y. 2003. Ethnoecological study of the Dani-Baliem society and the environment changes in Baliem Valley, Jayawijaya, Irian Jaya. Berita Biologi 6 (5), Edisi Khusus Kebun Biologi Wamena dan Biodiversitas Papua. Purwanto, Y. 2004. Aspek sosial budaya dan etnobiologi masyarakat di sekitar kawasan cagar alam cycloops. Report for Indonesian Institute of Sciences (LIPI), Bogor. Purwanto, Y., R.D.A. Darmayana, and E.B. Waluyo. 1990. Penembangan pesawahan di Lembah Baliem, Wamena. Pp. 267–281 in Proceeding Lokakarya Pengembangan Wilayah Pedesaan Wamena. T.T.G. LIPI, Wamena. Richard, P. 1985. Indigenous Agriculture Revolution. Unwin Hyman, London. Schuiling, D.L., and F.S. Jong. 1996. Metroxylon sago. Pp. 121–126 in PROSEA, Plant yielding non-seed carbohydrates. Serpenti, L.M. 1965. Cultivators in the swamps: social structure and horticulture in a New Guinea society (Frederik Hendrik Island, West New Guinea). Ph.D. diss., University of Amsterdam. Van Gorcum, Assen. Sillitoe, Paul. 1983. Roots of the Earth: Crops in the Highlands of Papua New Guinea. New South Wales University Press, Kensington. Sillitoe, P. 1996. A Place Against Time: Land and Environment in the Papua New Guinea Highlands. Harwood Academic Publishers, Amsterdam.

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1148 / manuel boissie` re and yohanes purwanto Sillitoe, P. 1998. It’s all in the mound: fertility management under stationary shifting cultivation in the Papua New Guinea Highlands. Mountain Research and Development 18 (2): 123. Soenarto, F. Rumawas. 1997. An agro-ecological analysis of wen-tinak, a sustainable sweet potato wetland production system in the Baliem Valley, Irian Jaya, Indonesia. Science in New Guinea 23 (2): 55–66. Steensberg, A. 1980. New Guinea Gardens: A Study of Husbandry with Parallels in Prehistoric Europe. Academic Press, London. Sumule, A. 1994. The technology adoption behaviour of the indigenous people of Irian Jaya: a case study of the Arfak tribals. Ph.D. diss., Department of Agriculture, University of Queensland. Townsend, P.K. 1992. Social constraints to the subsistence production of sago. Pp. 161–167 in Levett, M.P, J. Earland, and P. Heywood (eds.) Proceedings of the First Papua New Guinea Food and Nutrition Conference. Tucker, A. F. 1987. Ekosistem-Ekosistem Tani di Irian Jaya dan Arah Pembangunannya. 2 vols. STAKIN, Sentani. Waddell, E. 1972. The Mound Builders: Agricultural Practices, Environment, and Society in the Central Highlands of New Guinea. American Ethnological Society Monograph 53. University of Washington Press, Seattle, Washington. Wolf, E.C. 1986. Beyond the green revolution: new approaches for third world agriculture. World Watch Paper 73), Worldwatch Institute, Washington, D.C. Yen, D.E. 1973. The origins of Oceanic agriculture. Archaeology and Physical Anthropology in Oceania 8 (1): 65–85. Yen, D.E. 1993. The origins of subsistence agriculture in Oceania and the potentials for future tropical food crops. Economic Botany 47 (1): 3–14.

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6.4. Patterns of Commercial and Industrial Resource Use in Papua dessy anggraeni Overview of Socioeconomic Conditions in Papua a p ua i s t he l a r ge s t Indonesian province with the smallest population, the majority of whom are poor. More than two million people live in an area of 41.48 million ha, 72.50% (30.07 million ha) of which is forested, according to the calculation of forest cover map by Forest Watch Indonesia, the Indonesian Forestry Planning Agency (BAPLAN: Badan Planologi Kehutanan), and Conservation International in 2002 (Forest Watch Indonesia 2002). The rate of population growth declined during 1991–2000 to 2.50% per year in 2000, from 3.86% per year in 1990. According to the Central Bureau of Statistics (BPS: Badan Pusat Statistik), in 2000 the population of Papua reached 2,219,500 people, more than 70% of whom lived in rural areas (BPS 2000). Within Papua, there are more than 250 tribes with their own languages and traditional systems (Taime 2002; Chapter 1.3).

P

regional gross domestic product The data show that regional gross domestic product (GDP) increased between 1993 and 2002. Here, GDP is defined as the total added value (goods and services) produced by economic sectors as results of the activities by production units (BP3D: Agency for Planning and Coordination of Regional Development, Badan Perencanaan dan Pengendalian Pembangunan Daerah, and BPS of Papua 2002). In 2002, the regional GDP reached almost nine trillion rupiah at constant prices, and more than 23 trillion rupiah at current prices. The data also show that the mining, gas and oil, and quarrying sectors contributed the largest proportion of regional GDP during these years. Therefore, these sectors contributed significantly to the growth of GDP in Papua. Without mining, total GDP in Papua in 2002 was only Rp 10.80 trillion, at current prices. In 2002, the total GDP decreased slightly from about 24 trillion rupiah to 23 trillion rupiah. This is because the total value added from the mining sector decreased from 14.68 trillion rupiah in 2001 to 11.39 trillion rupiah in 2002 due to a fall in the price of copper concentrates. It is also worth noting that when the economic crisis hit Indonesia, the GDP of Papua jumped from less than 10 trillion rupiah in 1997 to more than 19 trillion rupiah in 1998. This is because the total value from mining increased by 38% in that year and increased economic growth in Papua by 13% (BP3D and BPS of Papua Province 2002). The average per capita income (which is obtained by dividing total GDP at current price by number of people at midyear) at 2002 prices was about 9.67 Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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million rupiah (when mining is included), which is the fourth highest per capita income in Indonesia, following East Kalimantan, Jakarta, and Riau provinces. Yet in real terms, local people in Papua have not enjoyed this income because the majority of this income belongs to one of the mining companies, namely PT Freeport Indonesia. Without mining, the average per capita income at 2002 prices was about 4.53 million rupiah, which is much lower than the per capita income when mining is included (BP3D and BPS of Papua Province 2002). Despite the high mean per capita income, the average conceals the significant disparity among the regencies in Papua. For example, in 2000, the mining regency of Mimika with 45,750 people had an average per capita income of about Rp 76.8 million, whereas the hilly, isolated, and populous agricultural district of Jayawijaya with 472,800 people had only a little over Rp 500 thousand (Mollet 2001).

total exports from papua Total value of exports from Papua Province in 2002 reached about US$ 1.7 billion, down slightly from the total value in 1998 of US$ 1.9 billion. Most of Papua’s market for export is in Asia, which consumes more than 60% of exports (⬎40% to Japan, ⬎20% to South Korea, ⬎10% to the Philippines). Remaining exports go to European countries (Spain, Finland, and Germany), United States, and Australia (BPS Papua 1999). Mining products (e.g., copper concentrates) comprised the majority of export value (US$ 1.439 billion or more than 84% of total exports), followed by plywood (US$ 70 million or 4% of total export value), frozen fish (US$ 67 million), gas and oil (US$ 54.9 million), and frozen prawns (US$ 52 million) in 2002 (BPS Papua 2002).

total domestic and foreign investment in papua Data from the Regional Promotion and Investment Board (BPID: Badan Promosi Investasi Daerah) of Papua Province (2002) showed that both domestic and foreign investment has increased significantly during the period between 1991 and 2002 (total domestic investment and foreign investment increased by Rp 18.43 trillion and US$ 43.83 billion, respectively, during this period). Domestic investment has increased from Rp 477 billion in 1991 to Rp 3.729 trillion in 2004, while foreign investment has increased from US$ 955,769 in 1991 to US$ 6.57 billion in 2002. In 1999, more than 86% of foreign investment was allocated to the mining sector, while the rest was invested in services (7.5%) and timber industry development (3%). These proportions changed in 2002, when foreign investment in the mining sector decreased to 70% (though the absolute amount is still relatively high, at US$ 4.6 billion) and the investment in services increased to 21% (to US$ 1.4 billion from only US$ 389 million in 1999). Services include construction and infrastructure development, marketing services, telecommunications, logistics, and other services related to the mining sector. In 1999, domestic investment was allocated as follows: timber industry (26%), plantations (25%), forestry development (21%), services (8.5%), fisheries (8.5%), and hotel and tourism development

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(7.5%). These proportions also changed in 2002, when the domestic investment for timber-based industries decreased to 12% while the investment for fishery increased to 26%.

employment by sector in papua Although mining contributed the biggest portion of total GDP in Papua, most people in Papua are part of the agricultural sector, which includes food crop farming, commercial crop plantations, animal husbandry, forestry, and fishery. The number of people working in the agriculture sector was estimated to be 768,774, contributing about 77.3% of all employment in Papua.

infrastructure Currently, there are two main road systems in Papua, one across the Central Cordillera (Jayapura-Wamena-Enarotali-Nabire) and one across the western sector (Sorong-Manokwari). Road density (ratio of length of road to size of the area) in Papua is one of the lowest in Indonesia (0.04 km/km2), far below the average of road density at the national level (0.17 km/km2). High traffic volume is found only in the cities, especially in Jayapura; it is very low in other regional road sectors. In terms of sea transportation, Papua has 11 national seaports (Biak, Bintuni, Fakfak, Kaimana, Manokwari, Merauke, Nabire, Pomako, Sarmi, Teminabuan, and Wasior) and two international seaports (Jayapura and Sorong). Air transport also constitutes a significant component of the regional transportation system and offers access to isolated areas. In Papua the main airports are in Jayapura and Biak, supplemented by many small airports across the area (BP3D of Papua Province 2004).

Patterns of Current Commercial and Industrial Resource Use in Papua

mining Papua is rich in minerals, oil, and natural gas. Because it is located along the ‘‘Ring of Fire’’ where the Indo-Australian and Pacific plates collide, it is in one of the highest mineralization zones in the world (Dwiyana 2001). There are two types of extractive mining activities in Papua: mining by local people and mining by large companies.

Mining by Local People Gold mining activity by local people commenced in the early 1990s and is carried out in four areas by local people with licenses from the Ministry of Mining and Energy. Those four locations are Subdistrict Web, Subdistrict Uwapa, Topo District, and Subdistrict Senggi (Mining and Energy Services of Papua Province 2000). Subdistrict Web, Jayapura District, covers rivers in three villages with total area of 385 ha. Subdistrict Uwapa, Nabire District, is considered to be a big gold mining area, employing about 12,000 people, covering the rivers Buaya (2,870 ha), Sowasowa (6,175 ha), Adai (4,365 ha), and Matao (4,620 ha). The third and fourth

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gold mining areas are Topo District, Nabire, covering some small rivers diverted from the rivers listed above; and Subdistrict Senggi, East and West Sentani, Jayapura District covering the rivers Pas, Pis, Maru, and Ukopulo.

Mining by Companies under Work Contracts (KK) Under Mining Regulation Law No. 11/1967, the Government of Indonesia opened foreign investment in mining activities under Work Contracts (KK: Kontrak Kerja). PT Freeport Indonesia is the only mining company in Papua that is producing under this law. Other mining companies are still in the process of exploration, including PT Nusamba Duta, PT Siriwo Mining, and PT Iriani Mutiara Idenburg (exploring for gold); PT Cyprus Amax Iriani (exploring for gold, copper, and metal); PT Gag Nikel and PT Iriani Mutiara Mining (exploring for nickel); PT Ingold Antares (exploring for gold and metal); PT Iriani Sentani (exploring for gold and nickel); PT Persada Pertama Mulia (exploring for coal); PT Karunia Pola Daya Bumi and PT Kumamba Mining (exploring for mineral sands); and PT Mineralindo Mas Salawati and PT Nabire Bakti Mining (exploring for gold and copper). Based on the data from the Mining Advocacy Network (JATAM: Jaringan Advokasi Tambang) (2000), the total area approved for small-scale mining and contracted for commercial mining investigation, exploration, and exploitation add up to about 11 million ha, mostly in the northern parts of Papua, accounting for about 25% of the total provincial territory. Figure 6.4.1 shows the distribution of mining concessions in 2000. As mentioned above, PT Freeport Indonesia (PTFI) is the only mining company that is successfully mining in Papua. After depletion of the very rich Ertsberg ore body, the company moved on to the nearby Grasberg deposit in 1988, and has invested US$ 4 billion from investing partners in 23 countries (Marsh 1997, cited in Dwiyana 2001). PTFI directly employs ca 7,800 people, with another 1,600 contract workers. Of this total of 9,400 employees, approximately 2,500, or 27%, are Papuans (FCX 2003). The original Contract of Work (KK) between PTFI and the Government of Indonesia was entered into in 1967 and was replaced with a new Contract of Work in 1991, encompassing an exploration area of 26,150 km2. In 1997, the Government of Indonesia altered the permission granted to PTFI and the area designated for exploration was reduced to 12,997 km2 (Mining and Energy Services of Papua Province 2000). The initial term of the current Contract of Work expires in 2021, but can be extended for two 10-year periods subject to approval by the Indonesian government. In 1996, PT Freeport Indonesia established joint ventures with Rio Tinto PLC, an international mining company with headquarters in London, England. The joint venture covers mining operations in Block A and gives Rio Tinto, through 2021, a 40% interest in production from Block A (FCX 2003). The Grasberg complex contains the world’s largest gold deposit and the second largest copper deposit, all in one ore body. In terms of mineral reserves, at the end of 2003, the Grasberg complex still contained aggregate proven and probable re-

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Figure 6.4.1. Distribution of mining concessions in Papua prior to 2000. serves of 54.4 billion pounds of copper and 60.4 million ounces of gold. According to PT Freeport Indonesia (2003), reserves established as of 2003 will give them the ability to mine for decades to come. PTFI’s reserves reflect estimates of the reserves that can be recovered before the end of 2041 (the end of the two potential 10-year extensions). Mill throughput averaged 203,000 metric tons of ore per day in 2003, which is lower than previous years (about 230,000 MT/day). The lower mill throughput during 2003 reflected the impact of the open pit slippage and debris flow events and the subsequent clean up efforts. However, it is also worth noting that the production from PT Freeport Indonesia’s Deep Ore Zone (DOZ) underground mine nearly doubled in 2003 from the previous year, with an average of 40,500 metric tons of ore per day. DOZ operations continue to perform above design capacity of 35,000 metric tons of ore per day, consisting of 1.08% of copper per metric ton ore, 0.98 grams of gold per metric ton ore, and 3.72 grams of silver per metric ton ore. PT Freeport Indonesia is completing studies to increase the capacity of the Deep Ore Zone mine to 50,000 metric tons of ore per day, which would make it one of the world’s largest underground operations (FCX 2003). Approximately one-half of PTFI’s concentrate production is sold to its affiliated smelters, Atlantic Copper (PTFI’s wholly owned smelting and refining facility in Huelva, Spain) and PT Smelting (PTFI’s 25% owned smelter facility in Gresik, East Java,

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Indonesia); the remainder is sold to other customers (FCX, 2001). Table 6.4.1 shows the production of mill throughput per day during the period of 2000 and 2003. Since discovery of the Grasberg mine in 1988, the complex has already yielded an aggregate 16.6 billion pounds of copper and 25.8 million ounces of gold. Production of gold, copper, and silver from PTFI fluctuated between 1996 and 2003. Copper production has increased from 36 billion pounds to 40 billion pounds, while gold has decreased from 47.4 million ozs to 46.6 million ozs. Silver production increased from 100 million ozs to 117 million ozs per year. Though it employs less than 1% of the labor force, the mining sector contributes the largest portion of total GDP in Papua, now 60%—an increase from 50% in earlier years. This changed slightly in 2002, where the contribution of mining sector to total GDP decreased to 54%, almost the same as before economic crisis hit Indonesia in 1998, when the average contribution of mining sector was about 50%. During the period of 1995 to 2000, the direct benefits of PT Freeport Indonesia to the Government of Indonesia have totaled US$ 1.28 billion, consisting of dividend, royalty, taxes, and retribution, while the indirect benefits have totaled US$ 4.34 billion, consisting of salary/wages, local procurement, local area development, and charitable contribution and reinvestment. The benefit of PTFI to Indonesia declined during the period of 1995 to 2000. In 1995, the benefit of PTFI to Indonesia was US$ 1.27 billion and in 2000 it dropped to US$ 643 million (FCX 2000). The main reason was the drop of gold and copper prices in Indonesia, from US$ 322/oz in 1995 to US$ 276/oz in 1999 for gold, and from US$ 2,459/ton in 1995 to US$ 1,336/ton in 1999 for copper, though in 1997 the price of copper rose to US$ 2,101/ton from US$ 1,971/ton in 1996 (World Bank 2001).

Gas and Oil Crude oil was Papua’s first main extractive product, tapped in wells surrounding Sorong, Salawati, and Bintuni Bay. The first productive gas and oil company in Papua was Pertamina Operation EP Sorong that covered area of 150 km2 in Klamono, Salawati Block 01X, and Wiriagar since 1964. Serious exploration to find

Table 6.4.1. Production of mill throughput per day, 2000–2003 (metric tons of ore per day) Source of product

2000

2001

2002

2003

Grasberg open pit

201,150

211,400

194,500

155,700

3,000

5,500

21,800

40,500

19,350

20,900

19,300

6,800

223,500

237,800

235,600

203,000

Deep ore zone underground mine Intermediate ore zone underground mine Total mill throughput

Source: Freeport-McMoRan Copper and Gold, Inc., 2003.

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new reserves was continued. In 1998 new reserves of natural gas were found in Wiriagar-Bintuni. This new natural gas field was called ‘‘Tangguh.’’ Exploration of the Tangguh LNG Project in Bintuni Bay (a project between the world’s third largest oil group, Britain’s BP PLC, and Indonesia’s state-owned oil and gas company Pertamina) has shown that the supply of liquified gas in the bay could last 15 to 20 years, with an annual supply of up to 3 million tons. Plant construction by BP is to be completed by 2006. This project would cpntribute tremendous additional income to the central Indonesian government, adding to the US$ 3 billion sent to Jakarta annually from the provinces (Dourueng 2002). The gas and oil sector has been contributing significant revenues to the national and provincial government. Total GDP from the gas and oil sector was steady at around Rp 300 billion until 1997; it jumped dramatically in 1998, reaching Rp 824 billion. It dropped significantly in 1999 to Rp 577 billion due to the drop of gas and oil production to 1,834,085 brl in 1999 compared to previous years, which averaged 7,792,832 brl (Dwiyana 2001). Gas and oil has contributed 3.37% to 5.54% of total GDP in recent years. Gas and oil contributed 3.12% of total export value with almost US$ 55 million in 2002 (from 1.35 million brl of natural oil and 341 thousands brl of residual oil). Figure 6.4.2 shows distribution of gas and oil concessions in Papua until 2000.

Figure 6.4.2. Distribution of gas and oil concessions in Papua prior to 2000.

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agriculture: food crops, cash crops, and animal husbandry The contribution of the agricultural sector to total GDP in Papua has increased over the years and in 2002 its contribution to total GDP reached 8.8%, or about 2.1 trillion rupiah at current prices. Among these agriculture sectors, food crops have contributed the biggest portion (72%). Agricultural commodities in Papua include rice, corn, cassava, potato, soybean, peanuts, vegetables, and fruits. Simply processed foods are also included in this sector, such as sago and sticky rice. Commodities for husbandry include poultry, pigs, and cows. Commodities for cash crops agriculture in Papua include coconuts, nutmeg, cacao, oil palm, cloves, coffee, rubber, and cashews. Plantation cash crops are divided into small-scale agriculture (smallholders) and private/state-owned agriculture (large-scale agriculture). In theory, large-scale agriculture should support small-scale agriculture (plantations owned by local communities) and act as an ‘‘agent of technology’’ for them (Plantation Services of Papua Province 1999a). In 1998, the land area under plantation development was 128,183 ha, including small-scale (97,159 ha), state-owned (31,024 ha), and privately owned (18,270 ha). Land licensed for plantation totaled ca 1.2 million ha. According to Plantation Services (1999a), potential land for plantation development in Papua is about 6,115,443 ha, which means that plantation development as of 1998 comprised 2% of its potential. Plantation Services of Papua Province (1999b) attributed the slow rate of plantation development in Papua to: difficult geographical conditions, including hilly and isolated areas; lack of transportation and other supporting infrastructure; unskilled human resources (farmers); lack of training for agricultural development; lack of accurate data to be used for developing plantation planning; and lack of facilities for pest control. The two types of large-scale commercial agriculture are state-owned and private. The one state-owned company in Papua, PTP Nusantara II, operates in two districts, Jayapura and Manokwari. By contrast, at least six purely private companies have been operating in Papua: PT Cockran, PT Nusa Irian Jaya Indah, and PT Varita Maju Utama in Manokwari; PT Adi Jaya Mulia in Fakfak; CV Jaya Abadi in Merauke; and PT Sinar Mas Group in Jayapura/Lereh. Large-scale commercial agriculture in Papua grew quite rapidly between 1993–1998, especially between 1995 and 1996, when growth reached more than 165% (from 12,668 ha to 33,600 ha). During that six-year period (1993–1998), the average growth of private plantations was about 8,200 ha/year. Normally, land for plantation comes from land conversion under the Timber Extraction License (IPK: Ijin Pemungutan Kayu) system. Until 1998, the area of large-scale commercial plantations was about 49,286 ha, with oil palm accounting for a total of 41,284 ha, or almost 84% of all privately owned plantation area in Papua. Oil palm production contributed 82% of all commercial crop products in Papua, followed by coconut with 11%, and cacao with 6% (BPS 1997). Things are different for small scale plantations, where coco-

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nut plantations are the largest part of small-scale plantations with 33% (covering 32,181 ha), followed by cacao with 23% (22,995 ha), oil palm with 19% (17,533 ha), cashews 7% (7,016 ha), nutmeg 6% (5,430 ha), coffee with 5% (5,065 ha), and rubber with 4% or 3,750 ha (Plantation Services 1999). Merauke District has the greatest potential for plantation development, covering 44% of total potential area for plantation development in Papua, followed by Manokwari with 16%, Jayapura with 13%, Sorong with 11%, and Fakfak with 3% (Plantation Services of Papua Province (1999b). In terms of employment, plantation sectors employed 150,686 people in Papua or almost 20% of the labor force. The government of Papua plans to increase the production of commodities from plantations in order to increase the plantations’ contribution to the regional economy (at the incremental rate of 4.2% per year). The government of Papua has been trying to increase the production of cacao by 7% per year, coconut by 5% per year, and oil palm by 15% per year. Figure 6.4.3 shows the distribution large-scale commercial plantations that have been operating in Papua.

forestry According to a decree of the Ministry of Forestry (2001), roughly 42 million ha of Papua is forested. Of this, more than 52% has been designated as production

Figure 6.4.3. Distribution of large-scale commercial plantations in Papua prior to 2000.

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forest. This includes 2 million ha of limited production forest, 10 million ha of permanent production forest, and 10 million ha of conversion forest (Forestry Services of Papua Province 2001). The commercial trees in Papua’s forests produce about 25–40 m3/ha, with diameter between 20 and 49 cm. The number of harvestable trees varies between 16 and 42 trees/ha (UNCEN 1999). Commercial trees in Papua are dominated by merbau (Intsia spp.), matoa (Pometia spp.), and other mixed commercial trees (Forestry Services of Papua Province 2001). In 1984, Licensed Logging Concessions (HPH: Hak Pengusahaan Hutan) were developed in Papua with PT You Lim Sari given the first Logging Concession License (HPH) with area of 367,000 ha, in Jayapura. In March 2003, the number of Licensed Logging Concessions (HPHs) rose to 56 (with total area of more than 12 million ha) under Ministry Decrees and as many as 11 Industrial Timber Licenses (IUPHHK: Izin Usaha Pemanfaatan Hasil Hutan Kayu) from the Governor or the Head of Regency (Bupati), comprising a total area of 944,666 ha. In March 2003 many Licensed Logging Concessions (HPH) and Industrial Timber Estates (HTI: Hutan Tanaman Industri) had not yet begun operation, including 35 Licensed Logging Consessions (HPHs) with a total size of 4,413,726 ha (Kayoi 2004). Data from Forestry Services of Papua Province (2001) show that the size of each concession varies from 51,600 ha (PT Hanurata I) to 691,700 ha (PT Mamberamo Alas Mandiri). The growth and distribution of logging concessions can be seen on Figure 6.4.4. The districts of Merauke, Jayapura, and Manokwari have the largest areas of Licensed Logging Concessions (HPHs) in Papua covering 30%, 13%, and 12%, respectively. There are several possible reasons for this, such as the advanced status of these districts, the availability of flat forest land, and the availability of rivers for log transport (Wurarah 2001). Data from the Ministry of Forestry and Crops Estate (1999) and Forestry Services of Papua Province (1999) showed that 13 private companies were interested in investing capital in the establishment of Industrial Timber Estates (HTIs) with the purpose of supplying timber for the pulp and paper industries. The designated timber estate area in 1998 was about three million ha, located in Merauke, YapenWaropen, Fakfak, and Sorong. Most of the Industrial Timber Estate (HTI) land can be found in Merauke (2.7 million ha), which contains more than 90% of all proposed HTI areas in Papua. Potential timber volume in the Industrial Timber Estate (HTI) system is about 150 m3/ha, with a life cycle of ten years. Table 6.4.2. shows a list of designated Industrial Timber Estates (HTI) in Papua in 1999. Logging operations contributed more than 7% of total GDP in 1994, but decreased to 4% in 2002 (BP3D and BPS of Papua Province 2002). The forestry sector employed ca 1.2% (or about 21,835 people) of the labor force. However, in several districts, including Merauke, Mimika, Manokwari, and Yapen-Waropen, logging accounted for over 20% of the total district GDP in 1999, and presumably a much higher percentage of the labor force was employed in the forestry sector in those districts. The importance of this sector is also reflected in its contribution to provincial budgets.

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Figure 6.4.4. Distribution and growth of logging concessions in Papua, 1984–1997. In Papua, production forest constitutes about 22 million ha, about half of which has been designated as logging concessions. Comparison of land cover from RePPProt (1985) and satellite imagery in 1991 and 1997 indicates significant changes in land cover during that 12-year period. In 1984, forest cover totaled 34,958,300 ha (or about 84.30% of total area), whereas in 1997 it totaled 33,548,021 ha (or about 82.30% of total area). This means that the deforestation during those period was about 1,410,279 ha; each year Papua lost more than 100,000 ha of its forest (Kapisa 2004). Average log production in Papua between 1995 and 2000 totaled 1,701,543 m3 per year, just 38% of the government’s target, so in 2003 the government reduced its timber production target to 2.7 million m3 per year (Kayoi 2004). Data from 1995 to 2000 show the decline of log production from Licensed Logging Concessions (HPHs) but show the increase from areas with Timber Extraction Licenses (IPKs: Ijin Pemungutan Kayu). Several reasons account for the drop in log production in Papua, one of which is a decrease in the number of active HPHs (in 2001, 12 of 54 HPHs were inactive). However, it is worth noting that even though official log production has been decreasing, illegal log production has been increasing. Data from the Forestry Information Center, Ministry of Forestry (2003), show that about 600,000 m3 per

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Table 6.4.2. List of designated Industrial Timber Estates (HTI) in Papua in 1999 Company

Location

Area (ha)

PT Maharani Rayon Jaya

Merauke

206,800

Decree of Minister of Forestry (SK. Menhut No.5/Kpts-II/1998)

PT Okaba Rimba Makmur

Merauke

300,000

Finished FS and EIA

PT Eucalyptus Tanaman Lestari

Merauke

298,900

Finished FS and EIA

PT Permata Warna Timur Lestari

Merauke

300,000

Survey FS and EIA

PT Dafonsoro Digul Daya Usaha

Merauke

127,000

No follow up

PT Dammore Bayu Permai

Merauke

173,000

Boundary preparation

PT Merauke Hutan Lestari

Merauke

372,500

Processing

PT Mukti Artha Yoga

Merauke

158,930

Converting from HPH to HTI

PT Irma Sulindo

Yapen, Waropen, and Fakfak

199,628

PT Kamundan Irjan Sakti

Merauke

216,000

PT Wana Kerta Eka Lestari

Merauke

300,000

PT Bangun Kayu Irian

Sorong

PT Mitra Jaya Group

Merauke

Total

Step of Activity

Conversion

96,125

Survey FS and EIA

303,000

Conversion

3,051,883

Note: FS  Feasibility Study; EIA  Environmental Impact Assessment; HPH  Licensed Logging Concession; HTI  Industrial Timber Estate. Source: Forestry Services of Papua Province (1999).

month of logs have been cut and smuggled illegally in Papua, causing total loss of Rp 600,000 billion per month or Rp 7.2 trillion per year. In December 2001, there were 47 timber-processing operations in Papua with a total processing capacity of up to 1,611,220 m3 per year and a total investment of Rp 903 billion (US$ 130 million). Because of the economic crisis in 1997, legislation was implemented permitting log export (SK. DG. Production Forest 135/ Kpts/IV–PPPHH/98). Logs and processed timber from Papua have been exported to India, China, Hong Kong, Korea, Japan, Philippines, Malaysia, Europe, and the United States. The central government’s recommended volume of log exports from Papua during 1997–2000 was 204,552 m3, but the actual total was much lower, 68,577 m3. However, in 2001, the central government prohibited log export from Indonesia. Aside from Licensed Logging Concessions (HPHs), the existence of community cooperatives (kopermas: Koperasi Peranserta Masyarakat) has posed a threat to forest conservation in Papua. It is hard to find precise data on how big community cooperatives (kopermas) actually are and where they are located in Papua because

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of the ambiguity of regulations. Currently, 120 community cooperatives (kopermas) have logging licenses granted by the Papuan government (Papua Post, 18 November 2003). One community cooperative (kopermas) can only manage a maximum of 1,000 ha, with a production target of 15,000 to 25,000 m3 per community cooperative. However, in 2004, the Papua Government stopped granting logging licenses under the community cooperative (kopermas) scheme because of problems in implementation, including disorganization and lack capacity of the community cooperative, and the potential for corruption (Cenderawasih Pos, 14 February 2004). Non-timber forest products, such as rattan, sago, gaharu (eaglewood), masoi bark, lawang (clove flowers) oil, resins, and sandalwood, have been of commercial interest over varying time periods. Currently, the total production of these nontimber forest products (NTFPs) reaches 22,420 tons per year, being managed by 55 companies that have Licenses to Collect Non-timber Forest Products (IHHBK: Ijin Pemungutan Hasil Hutan Bukan Kayu) (Kapisa 2004). In addition to non-timber forest products, the government has also legalized the trade of some unprotected animals in Papua, including birds, reptiles, amphibians, insects, and some mammals. Data from the Provincial Forestry Office (1998) show considerable trade of wildlife from Papua’s forests. Bird and reptile trading has recently increased sharply, with offtake rates far exceeding quotas. Similarly, the amphibian and mammal trades have also grown. Deer and wallaby are hunted extensively near Merauke, and venison is exported by air and by ship to other areas in Papua and to other provinces. Birds such as parrots, cockatoos, and bird of paradise are trapped and exported to markets in Java and overseas. The crocodile trade has also contributed to local revenue in Papua. Five large companies control the crocodile trade in Papua. In 1998–1999, they sold 11,593 skins of Crocodylus novaeguineae and 9,043 skins of Crocodylus porosus, and caught 2,691 Crocodylus novaeguineae juveniles and 42,599 Crocodylus porosus juveniles (Forestry Office of Papua Province 1998). Table 6.4.3 shows data from Provincial Forestry Office (1998) indicating a considerable trade of wildlife from the forest area.

fisheries Papua’s commercial fisheries have grown steadily, with an annual growth rate of about 8% over the period between 1993 and 1997. The number of fishermen increased between 1993 and 1999 at a rate of 16% per year. In 1997, 73,013 fishermen in Papua operated 25,977 vessels across Papua’s oceans. The growth rate of fishing ships is about 6.2% per year (Fishery Services 1998). In 1997, 62 commercial fishing companies operated in Papua, primarily in Sorong, Biak, Merauke, Manokwari, Serui, and Jayapura. Seven of these companies belong to foreign investors, mostly operating in Sorong; these are PT West Inan Fish Industries, PT Irian Marine Product Development, PT Alfa Kurnia Fish Enterprise, PT Dwi Bina Utama, PT Artha Samudera, PT Kodeco Fishery, and PT Cendana Indo Pearls (BPID Papua 2001).

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875

376

10,243

600

27,007

14,787

Quota Transported

Source: Forestry Office of Papua Province (1998).

Mammals

900 33,855

63,400

Amphibians

Reptiles

55,890

Birds

Insects

Caught

Group

1994/95

2,253

390

4,835

Realization Transported

104

2,305

400

4,794

1995/96 Realization Transported

4,346

405

5,233

1996/97 Realization Transported

Table 6.4.3. Wildlife trade in unprotected animals in Papua, 1994–1999

1,375

450

1,975

1,330

450

1,775

1997/98 Realization Caught Transported

675

33,925

900

61,800

66,734

210

11,682

600

24,152

16,065

1998/99 Realization Caught Transported

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The contribution of the fishery sector to GDP rose steadily between 1994 and 2002. In 2002, the fishery sector contributed ca 1.4 trillion rupiah (6.2 %) to GDP in Papua (BPS 2003). Thus Papua’s fisheries are more important economically than the forestry sector. Most fishery products from Papua are exported; in 1997, 73,285 tons were exported while only 6,097 tons (8% of all fishes harvested) reached the domestic market (Fishery Services of Papua Province 1998). The value of exported fishery products was estimated to be more than US$ 82 million in 1997, whereas the domestic market was only valued about Rp 25 million in the same year. The export of fishery products fluctuated between 1977 and 1999, but has significantly increased, reaching US$ 138 million in 1999. Mixed large fishes (kakap) were dominant in terms of the volume, whereas frozen prawns contributed the largest portion of total export value. The largest jump in total export value from the fishery sector happened in 1990 when the total value grew by 43%. This is in line with the increase in total export volume during that year, when the total volume of fishery exports jumped by 45%. The total export volume continued to increase and in 1999 total export reached the highest point with more than 95 thousand tons. Frozen mixed fish and frozen prawns comprise almost 90% of total export value from fisheries. Figure 6.4.5 shows the contribution of each fishery product to total fishery export value in 2002. In terms of export volume, frozen mixed fishes contribute the largest portion of total export volume (more than 70% between 1998 and 2002; Fishery Services of Papua Province 2002). Merauke Regency contributed 35% of the total produc-

Figure 6.4.5. Contribution of various fishery products to total export value from the fishery sector, 2002.

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tion from the Papuan fishery sector, followed by Sorong Regency with more than 27%, as can be seen on Figure 6.4.6. A study by The Ministry of Marine Affairs and Fisheries (MMAF) and the Indonesian Institute of Sciences (LIPI) (2001) in Arafura Sea showed that maximum sustainable yield (MSY) in Arafura Sea in 2001 was about 771 thousand tons/year for all fishery products (pelagic fishes, demersal fishes, reef fishes, shrimps, lobsters, and squids) making the total allowable catch (TAC) an estimated 617 thousands ton/year (80% of the MSY). The total fishery production in that year (2001) was only 263 thousand tons, or about 34% of total MSY and 43% of TAC. However, even though total fishery production in Arafura Sea was still far below the MSY, some products (demersal fish, reef fish, shrimp, lobster, and squids) exceeded their MSYs. In addition to fishing for food, freshwater ornamental fishes are harvested. Data from Fishery Services (1998) shows that ornamental fish trade increased between 1993 and 1997. Arowana (Scleropages jardinii) is Papua’s most important freshwater ornamental, and export trade is growing in Papua. Other commercial ornamental fishes from Papua are rainbow fish, banded archerfish, glassfish, goby fish, catfish, and grunters (Chapter 5.5). The total number of ornamental fishes traded during the period 1993–1997 was 1.1 million fishes with a total value of Rp 8.93 billion. Arowana contributed the largest portion of ornamental fish trade in Papua with more than 63%.

Conclusion Papua contains abundant natural resources, such as minerals, oil and gas, forests, and marine resources. Mining and oil provide the largest segment of Papua’s GDP,

Figure 6.4.6. Percentage of total fisheries production by regency in Papua, 2002.

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while timber, agriculture, and fisheries contribute minor but important amounts. Papua’s economy continues to grow, and all of these sectors are expected to grow for the foreseeable future. The well-being of Papua’s rural people will depend upon the government investing the wealth generated by the various industries in the rural communities, to promote health, education, and locally-driven sustainable development. Environmental well-being and conservation of Papua’s remarkable biodiversity and ecosystem services will depend on large-scale planning by the various government agencies in ways that take into account the importance of these resources to the long-term well-being of Papua’s many rural societies.

Literature Cited BP3D (Agency for Planning and Coordination of Regional Development, Badan Perencanaan dan Pengendalian Pembangunan Daerah) and BPS (Central Bureau of Statistics, Badan Pusat Statistik) Papua. 2002. Gross Regional Domestic Products of Papua Province. Jayapura. BP3D (Agency for Planning and Coordination of Regional Development, Badan Perencanaan dan Pengendalian Pembangunan Daerah) of Papua Province. 2004. Draft of Presidential Decree on Spatial Planning in Papua Island. Technical Team of Spatial Planning and Infrastructure Development in Papua, Jayapura. BPID (Regional Promotion and Investment Board, Badan Promosi Investasi Daerah) of Papua Province. 2001. Directory of Domestic and Foreign Investment in Papua Province. Jayapura. BPID (Regional Promotion and Investment Board, Badan Promosi Investasi Daerah) of Papua Province. 2002. Domestic and Foreign Investment in Papua. Jayapura. BPS (Central Bureau of Statistics, Badan Pusat Statistik). 1999. Irian Jaya in Figures. BPS, Jayapura. BPS (Central Bureau of Statistics, Badan Pusat Statistik). 2000. Irian Jaya in Figures. BPS, Jayapura. BPS (Central Bureau of Statistics, Badan Pusat Statistik). 2002. Irian Jaya in Figures. BPS, Jayapura BPS (Central Bureau of Statistics, Badan Pusat Statistik). 2003. Irian Jaya in Figures. BPS, Jayapura. Daorueng, P. 2002. Indonesia: Gas Project Promises Income West Papuans Not Excited. Global Policy Forum Website. Inter Press Service News Agency. April 30, 2002. Dwiyana, A. 2001. Oil, gas and mining development and decision making process in Papua. Mining Services of Papua Province. Issue paper prepared for Rapid Assessment of Conservation and Economy (RACE), Jayapura. Fishery Services of Papua Province. 1998. Fishery in Papua in number 1993–1997. Fishery Services Papua Province, Jayapura. Fishery Services of Papua Province. 2002. Statistics of Fishery in 2002. Jayapura. Forestry Information Center (PIKA), Ministry of Forestry. 2003. Ministry of Forestry coordinates with Mabes TNI in combating illegal logging. Press release no. 51/II/PIK1/2003. Ministry of Forestry, Jakarta. Forestry Services of Papua Province. 1998. Statistic of forestry and plantation in Papua Province. Jayapura. Forestry Services of Papua Province. 1999. Statistic of forestry and plantation in Papua Province 1998/1999. Jayapura.

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1166 / d e s s y a n g g r a e n i Forestry Services of Papua Province. 2000. Statistic of forestry and plantation in Papua Province. Jayapura. Forestry Services of Papua Province. 2001. The forestry condition in Papua Province. Presented at Forestry Workshop of Papua Province. Jayapura. Forest Watch Indonesia (FWI). 2002. Calculation from forest cover map. Preliminary result of imagery interpretation. FWI–BAPLAN–CI. Freeport-McMoRan Copper and Gold, Inc. 2000. Working to produce value. Annual report 2000. Freeport-McMoRan Copper and Gold, Inc. 2001. Annual report 2001. Freeport-McMoRan Copper and Gold, Inc. 2003. The strength of our metals. Annual report 2003. JATAM (Mining Advocacy Network). 2001. Compilation of mining companies in Papua (update 2000). Jakarta. Kapisa, N. 2004. Policy in forest resource management under Special Autonomy in Papua. Jayapura. Kayoi, M. 2004. Direction and policy of Papua forestry development in Special Autonomy era. Forestry Services of Papua Province, Jayapura. Mining and Energy Services of Papua Province. 2000. Walking with potential mineral and energy in Papua toward Millennium III of human development. Jayapura. Mining and Energy Services of Papua Province. 2001. Annual report from Mining and Energy Services (1995/1996–1999/2000). Jayapura. Ministry of Forestry and Crops Estate. 1999. Directory of forestry enterprises and plantation. Jakarta. MMAF and the Indonesian Institute of Sciences (LIPI). 2001. Pengkajian Stok Ikan di Perairan Indonesia [Stock Assessment in Indonesian Marine Territorial Waters]. Center for Capture Fisheries Research MMAF and Center for Oceanic Research and Development, LIPI, Jakarta. Mollet, J.A. 2001. District budget collection under decentralization and the impacts to the natural resource in Papua. Issue paper prepared for Rapid Assessment for Conservation and Economy (RACE). Cenderawasih University, Jayapura. Plantation Services of Papua Province. 1999a. Statistics of plantation in Papua. Jayapura. Plantation Services of Papua Province. 1999b. Five years plantation development plan in Papua Year VII (REPELITA VII) 1998/1999. Jayapura. Taime, T. 2002. Community Livelihood in Papua. Issue Paper prepared for Rapid Assessment for Conservation and Economy (RACE) in Papua. Health Services of Papua Province. University of Cenderawasih (UNCEN). 1999. Study of Optimal Size of Logging Concession in Forestry Management in Papua. Tim Faculty of Agriculture. University of Cenderawasih, Manokwari. World Bank. 2001. Indonesia—Environment and Natural Resource Management in a Time of Transition. The World Bank, Washington, DC. Wurarah, R. 2001. An economic analysis of incentives to government, local communities and logging companies to engage in logging in Papua. Issue paper prepared for Rapid Assessment for Conservation and Economy (RACE), Papua. University of Papua, Manokwari.

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6.5. Natural Resource Economics of Papua james b. cannon a p ua i s o ne of few places remaining in the world where traditional cultures and tropical forests remain relatively intact. Yet significant areas of forest have already been lost, and Papua’s ‘‘wilderness’’ status is in jeopardy. The standard development argument for logging and converting these forests is that doing so generates government revenues, creates jobs, and increases incomes and opportunities for the people. Still more forests will be lost to development, but significant mining and gas resources mean Papua can afford not to log and convert more forest, and still have the revenues to grow economically and reduce poverty. Protecting Papua’s remaining forests also costs little in terms of lower economic growth or the ability to reduce poverty. Many of Papua’s conservation areas and remaining intact forests have small human populations and are relatively inaccessible. While much can and should be done to meet the needs of the poor and reduce poverty in these areas, major development investments in conservation areas contribute less to economic growth, employment, and poverty reduction in the aggregate than development investments focused in accessible areas with many people. Papua therefore has the opportunity to use revenues from gas and mining extractive industries to pursue development options that generate greater benefits than more environmentally destructive alternatives. This chapter looks at the macro level and asks how much of Papua’s remaining forests are at risk, where are the most threatened areas, where are the lowest-cost opportunities for conservation, and what are the real trade-offs with development. The answer to the last question in particular is largely determined by the future use of mining and gas resources in Papua, both in terms of the rate and extent of extraction and how the revenues are reinvested. This discussion focuses mainly on technical analyses, recognizing that while carrying out such analyses is necessary for sustainable development, they are but the first step in a long process of promoting transparency, resolving conflict, improving governance, and building government and other capacity. The particular challenges facing Papua are described in the final section. The extraction of natural resources dominates Papua’s economy and landscape. Natural resource extraction currently generates about 75% of Papua’s GDP, with about 60% of GDP coming from the Grasberg mine operated by FreeportMcMoRan Copper and Gold, Inc. (Chapter 6.4). Freeport’s mining operations are projected to continue to 2041 (Freeport-McMoRan Copper and Gold, Inc. 2004) and as more natural resources are extracted the sector will dominate even more in future. Of particular note is a huge new natural gas project, BP Tangguh, with reserves of 14.4 trillion cubic feet. Gas production is expected to start in 2008. Although negotiations continue over the amount of production capacity to be

P

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

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installed (Ministry of Industry 2005), initial plans were to produce seven million tons of liquified natural gas (LNG) per year (U.S. Embassy, Jakarta 2005). If these exports were taking place now they would be worth about two billion dollars, based on the revenues Indonesia earned from total LNG exports in 2004 (U.S. Embassy, Jakarta 2005a). In comparison, total other exports combined from Papua in 2002 were worth 1.7 billion dollars (Chapter 6.4). Although gas production will start soon, the structure of the financing and existing agreements about the order in which investors get paid back and how revenues are shared means Papua may not see direct revenue flows for a decade. However, Papua may not need to wait for some of the benefits from the Tangguh development because there is the possibility of borrowing against future gas revenues to make public investments now (British Petroleum 2005). The extraction of natural resources from Papua has created an economic windfall. Papuan GDP has grown at an average of nearly 10% since 1991, around double the national rate. Average GDP per capita in 2002 was around US$ 1,000, making Papua the fourth richest province in Indonesia on paper. However, this high average figure hides enormous disparities. While Mimika, the Regency with the Grasberg mine, had a GDP per person of around US$ 9,000, those in the populous Regency of Jayawijaya had very much less at around US$ 60 (Mollet 2001). The mining boom has encouraged urban development. Papua’s urban areas are among the top few wealthiest regions in Indonesia, while Papua’s rural areas are the poorest in all of Indonesia (World Bank 2001a). More equitable sharing of benefits from the natural resource industries could have helped reduce poverty significantly, depending on where and how the revenues were used. In total, over 40% of the people of Papua live on less than a dollar a day (Mollet, 2001). This poverty can be hard to understand given the scale of extraction of natural resources from Papua. One explanation is that the value of these natural resource industries to the local areas is not as great as first appears. GDP counts all economic activity, yet the majority of the wealth generated by mining and forestry in Papua leaves Papua as returns to external investors and workers and revenues to other parts of Indonesia. One measure of the revenues that remain with local people—the Gross National-regional Product (GNrP)—is estimated at only 40% of GDP (Sheng 2004). In contrast, the ratio of GNP to GDP for Indonesia as a whole is approximately 95% (developed countries are typically close to 100%, with some even higher). Fortunately, in the future, Papua is set to receive a greater share of the revenues from extractive industries, and have a greater say in how the industries are run and how revenues are reinvested. In 1999 Indonesia adopted Law No. 22/1999 and Law No. 25/1999, which decentralized certain powers and responsibilities to the provinces and local level governments. Local governments got greater control over their finances, including managing a greater share of government spending and making the budget and setting local taxes (Gesellschaft fuer Technische Zusammenarbeit (GTZ) 2003). Elected regional councils were given greater legislative powers and oversight of the regional administrations (Gesellschaft fuer

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Technische Zusammenarbeit (GTZ) 2003). In 2001, Law No. 21/2001 gave Special Autonomy to Papua, which provided additional powers and a greater share of revenues than under the earlier general decentralization laws. The arrangements for revenue sharing gave the then-Papuan government 80% of the total government revenues from forestry, fishing, and general mining, and 70% of those for oil and gas. The Papuan government also received funds from the General Allocation Fund (DAU: Dana Alokasi Umum) to cover routine expenditures, and funds from the Special Allocation Fund (DAK: Dana Alokasi Khusus) to cover development investments, with a special provision for health and education (Anggraeni 2005). As Figure 6.5.1 shows, the total revenues of the Papuan government increased dramatically following decentralization, reaching nearly two trillion rupiah (about US$ 235 million) in 2002, though a significant part of the additional funding was to cover Papuan government salaries formerly paid by the central government. Unfortunately, the Special Autonomy Law has only been partially implemented. Papua is now also in the process of being divided into three provinces, and the number of regencies has nearly tripled since 2000. The partial implementation and subsequent structural changes in numbers and boundaries of regencies and provinces makes it difficult to understand how the revenues of the various new local governments will develop. Nonetheless, assuming continued revenue-sharing arrangements similar to those envisaged in the Special Autonomy Law, then Papua

Figure 6.5.1. Papuan government revenues climbed after the decentralization of the Indonesian government. Source: Anggraeni (2005).

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as a whole should benefit greatly given the anticipated additional revenues from the Tangguh gas project. Can development objectives in Papua be achieved without sacrificing Papua’s forests? The experience of other energy- and mineral-rich regions of the world suggests that it is possible, though many factors come into play, including avoiding investing revenues in large-scale infrastructure to open up remaining forest areas (Wunder 2003). The impacts of natural resources and their economics are large, interlinked, and complex, and shape the entire economy, particularly in a resource-rich setting like Papua. One way that the resource sectors are interlinked is through government revenues and expenditures. Government revenues from natural resources are fungible; they can be spent on different things and it does not matter which sector contributed the funds. A boom in one sector therefore reduces the need for revenues to be raised from another. Sectors are also linked through prices. A boom in one sector increases demand for construction and services, driving up the costs of labor and other inputs across the economy. These higher costs reduce the profitability of other sectors. In national settings, exports from booming oil, gas, and mining sectors increase the inflow of foreign exchange, strengthening the currency and decreasing the competitiveness of other export sectors. Forests are protected by the resulting decline in logging for export or conversion to grow cash crops for export. Other factors can complicate the picture, but oil wealth in other countries, particularly those with otherwise small economies and low populations, has been shown to reduce forest loss (Wunder 2003). Hence the wealth from Papua’s mining and gas resources could reduce the pressure for logging and forest conversion. However, Indonesian national interests will encourage a greater rate of natural resource extraction, because the economic need at the national level is much greater. Nonetheless, with an increasing share of revenues, and a greater say over development decisions, going to Papua, mineral wealth should reduce the need for further logging and deforestation. Greater revenues, together with large expanses of perceived ‘‘underdeveloped’’ areas, can inspire dreams of ‘‘mega-development’’ projects, such as the Mamberamo dam project championed by former President Habibie. Such ‘‘mega-development’’ projects in remote areas run the risk of swallowing enormous resources without generating equivalent returns. Avoiding bad investments is a good start towards improved development, but is not enough. The key is making good investments, in terms of sectors, but also in terms of location. New findings in development economics point towards the importance of geography in determining development success. Analyses of development patterns globally have shown that urban and coastal areas with high population densities are generally the areas where development progresses most quickly (Gallup et al. 1999). Similar studies done within single countries produced similar results. For example, in India, physical geography explained 66% of urbanization, which in turn explained 82% of variation in growth among states since 1990

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(Sachs et al. 2002). The work on economics and geography does differ in explaining some of the causes for variation (e.g., Easterly and Levine 2002), but all agree that, historically, some places have been much better suited for development than others by virtue of their location. Applying these findings from economic geography to Papua suggests development efforts might yield the greatest results (in terms of the greatest good in aggregate for the greatest number of people) if they are focused in locations with the most promise for economic development and poverty reduction. Such locations include existing areas of high intensity agriculture, urban centers, and coastal areas. If this proves to be the case, then maximizing development benefits may also maximize the conservation of remaining forest areas, delivering a macroscale ‘‘win-win’’ for the economy and the environment. While major development efforts may be better directed away from remoter areas, investments in health, education, publicly subsidized transport, and other government services must be directed towards more remote areas to help meet the needs of the poorest of the poor in remote rural locations. This brief overview of Papua’s natural resources and economy suggests Papua has the opportunity to avoid the fate of other parts of Indonesia, where only a few percent at most of original lowland forest cover remains (Jepson et al. 2001). Papua has ample revenues from mining and gas operations to fund development objectives. Provided those funds are invested wisely, Papua should be able to grow economically and reduce poverty without further reducing forest cover. Exploring whether this macro-scale ‘‘win-win’’ can be achieved or not requires an improved understanding of the spatial distribution of forest use to identify the areas already impacted and those most at risk in future. Macroeconomic modeling is then required to assess how alternative development options perform against various economic, social, and environmental criteria. Finally, even if an economic and environmental ‘‘win-win’’ is theoretically possible, various obstacles may prevent it being achieved. Critical obstacles include political and social conflict, corruption, lack of government capacity in various guises, and imperfect democratic processes. These obstacles are described in the discussion section.

Identification of Economic and Environmental Issues First, though, is a description of the data and methods that were used to develop the spatial and macroeconomic analyses, followed by the results.

data and methods Between 2001 and 2002 Conservation International (CI), with the support of USAID, carried out a project to integrate conservation information into development planning and improve broader technical capacity and understanding of environment and development issues at the provincial level. Indonesia was then undergoing a process of decentralization, and the project was designed as a partici-

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patory process, both to improve quality and to strengthen local government capacity and decision-making processes. Papua was a particular priority because of its biodiversity importance and the extra decentralization envisaged under Special Autonomy. As a first step, Conservation International undertook a Rapid Assessment of Conservation and the Economy (RACE) to improve understanding of development trends, issues, and interactions with the environment. In this RACE, a CI expert first prepared an overview of the Papuan economy and key issues (Anggraeni 2001) for discussion with local experts from Papuan academia, NGOs, and local government. Key issues were identified for further analysis, and Papuan experts were invited to write analyses of important development and environment issues facing Papua. These issues were: mining, oil, and gas (Dwiyana 2001); forestry economics (Wurarah 2001); forestry policy (Nugroho 2001); district budget revenue collection (Mollet 2001); provincial decision-making processes (Sugiono 2001); institutional capacity (Wiratno 2002); and community livelihoods. Each report was reviewed and critiqued by other experts in Papua in technical focus group meetings, to improve the quality of the analyses, broaden understanding, and develop a consensus. Over 100 local experts were involved in the development and discussion of these reports. All of the authors and experts then met to discuss and cross-reference their various reports and develop an integrated assessment of development trends and impacts and opportunities for conservation and development. As inputs to the meeting, a Conservation International expert developed an initial integrated economic analysis (Sheng 2002) and a Center for International Forestry Research (CIFOR) expert developed an integrated spatial analysis (Mertens 2002a). Participants at the meeting identified the major issues facing Papua as poverty and the underdevelopment of health and education services, a heavy reliance on extractive industries, the threats to the environment posed by these extractive industries, and too little institutional capacity in development planning and natural resource management. Significant progress had been made in improving understanding of development trends and their impacts on and linkages with the environment. However, participants in the integrated workshop recognized the need for further work. They prioritized developing objectives and indicators for sustainable development, comparing different large-scale development options, exploring the potential for diversifying into non-extractive economic activities, and addressing land use conflicts and natural resource management issues in conservation areas. Participants also recognized the need for a formal mechanism to help ensure these additional activities were carried out, and to facilitate continued dialogue among experts in different fields and from different institutions. They supported the proposal for a permanent forum on conservation and development (Sheng 2002), later named the Forum for Conservation and Development in Papua (FKPTP: Forum untuk Konservasi dan Pembangunan di Tanah Papua). The participants assigned a small team to develop the forum concept and sug-

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gest statutes, an organizational structure, membership numbers, and balance among different parties. The proposal of the small team was supported with a formal letter from the Governor of Papua in 2004, and the first meeting of the Forum for Conservation and Development in Papua (FKPTP) was held in March 2005 (Conservation International 2005c). In parallel to the launching of the FKPTP, Conservation International funded experts of the Millennium Institute (MI) in Washington, D.C., to develop a macroeconomic model of Papua and carry out initial analyses of alternative development options (Pedercini 2004). This model compared how key economical, social, and environmental indicators would be expected to vary under different patterns of public and private investments. This chapter draws heavily on the spatial analysis carried out by the Center for International Forestry Research (CIFOR) (Mertens 2002a,b), which provided all of the maps of Papua showing vegetation types, land classifications, infrastructure impacts, and future threats posed by logging and other activities. This chapter also draws heavily on the macroeconomic modeling carried out by the Millennium Institute (MI) (Pedercini 2004), which provided all the graphs of economic, social, and environmental indicators. Both the CIFOR and MI studies relied extensively on the data and analyses of earlier Rapid Assessment of Conservation and Economy (RACE) work by CI and many partners in Papua. Together these two studies help give a picture of what current development patterns, or ‘‘business as usual,’’ will mean for Papua’s economy, people, and forests, and how possible alternatives compare.

spatial analysis of natural resources and infrastructure For the purposes of these analyses, five different types of forests were identified in Papua: mountain forest, lowland forest, swamp forest, mangrove, and ‘‘other.’’ The mountain forests are mainly found along the main mountain spine, while lowland forests cover most of the rest of the island, with the exception of small coastal areas of mangrove and the large swamp forests of the south coast (Figure 6.5.2). Land uses were divided into three broad classifications: production forest, conservation forest, and ‘‘other.’’ Production forest included various sub-classifications, including types of logging operations intended to be long-term, and also forests to be logged and converted to other uses. Conservation forest included various types of reserves and parks, and also watershed protection forest. The ‘‘other’’ broad classification included agroforestry, plantations, transmigration sites, and other land uses. These land uses are shown in Figure 6.5.3. The boundaries of the three broad classifications are generally consistent across different spatial planning maps, but the boundaries and status of some of the subclassifications can change and are sometimes unclear. Over 60% of Papua was classified as lowland rainforest, while the large southern swamp forests account for over 10% (Table 6.5.1). Half of Papua’s lowland forest

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Figure 6.5.2. Vegetation classification in Papua. Source: Mertens (2002a).

was zoned as production forest, and 37% as conservation forest. Over 90% of the mountain forests of Papua were considered watershed protection forest because of their altitude and slope. 56% of mangroves, but only 19% of the southern swamp forests, were zoned for conservation. Production forest made up 44% of the total forest area (Table 6.5.2). Almost 70% of the production forest estate was lowland forest, with swamp forest making up most of the rest of the estate. Conservation forests accounted for 46% of the total forest area (Table 6.5.2). Almost 50% of the total conservation forest area was lowland forest, while mountain forests made up 30%. Swamp and mangrove forests each made up 6% of the total conservation area. Comparing Table 6.5.1 and Table 6.5.2 reveals that lowland and swamp forests constitute a disproportionately large share of the production forest estate. Lowland forests make up 60% of the total forest area but 70% of production forests, while swamp forests make up only 10% of the total forest area but 20% of the production forest area. In contrast, mountain forest makes up only 14% of the total forest area, but 30% of the conservation area. The mountain forests contribute a disproportionately large share of conservation forest because almost all are protected as watershed protection forests. Other steep-sloped forest areas are also classified as watershed protection forests, and protected forests in total make up over 60% of the conservation area (Table 6.5.3).

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Figure 6.5.3. Land use classification in Papua. Source: Mertens (2002a).

Table 6.5.1. Areas of vegetation classifications, and percentages in production and conservation forest land use classifications

Area (km2)

Percent of total area

Percent in production forest

Percent in conservation forest

Lowland

252,100

61

50

37

Mountain

57,900

14

4

94

Swamp

52,600

13

71

19

Mangrove

19,800

5

31

56

Other

28,500

7

36

53

Forest cover

Source: Author’s analysis of information from Mertens (2002a).

The rest of the conservation area is made up of a variety of parks and reserves, which together account for 17% of the total land area of Papua. Two-thirds of the production forests have been handed out as Licensed Logging Concessions (HPHs: Hak Pengusahaan Hutan); Table 6.5.3), almost all in lowland forest. Forest concessions began to be handed out in Papua in the early 1980s, as timber resources were exhausted elsewhere in Indonesia. By the early 1990s over

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Table 6.5.2. Percentage of total area and different vegetation classifications covered by production and conservation forest land use classifications Area (km )

Production forest

Conservation forest

179,900

187,000

Percent of total area

44

46

Percent lowland forest

69

49

Percent montane forest

1

30

2

Percent swamp forest

20

6

Percent montane forest

3

6

Percent other forest

6

8

Source: Author’s analysis of information from Mertens (2002a).

Table 6.5.3. Forest sub-class areas and percentages of forest class and total area Forest class

Forest sub-class

Area (km2)

Percent of forest class

Percent of total area

Production

Logging concessions Other

119,200 60,700

66 34

29 15

Conservation

Watershed protection Parks and reserves

115,200 71,800

62 38

28 17

Other

Other

43,900

100

11

Source: Author’s analysis of information from Mertens (2002a) and Wurarah (2001).

six million hectares had been awarded, and twelve million hectares by the time the system stalled in 1998 (Figure 6.5.4). The location and current status of Papua’s logging concessions are shown in Figure 6.5.5. The Licensed Logging Concession (HPH) system stalled because of decentralization and changes in the way forests were managed and by whom. The central government’s Ministry of Forestry, which had administered the Licensed Logging Concession (HPH) scheme, no longer had the authority to give out concessions in production forests. Local governments were given greater powers to issue Timber Extraction Licenses (IPK: Ijin Pemungutan Kayu) for small community cooperatives (kopermas: Koperasi Peranserta Masyarakat). Many of these Timber Extraction Licenses (IPKs) were awarded, though area and production data remains incomplete and fragmented. Nonetheless, from the production figures registered with the government, it appears that production from community cooperatives (kopermas) made up for some of the decline in official production from the Licensed Logging Concessions (HPHs). Following further legal changes, Timber Extraction Licenses (IPKs) became technically illegal. Although official figures were no longer reported, logging—now technically illegal—continued under both Timber Extraction Licenses (IPKs) and

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Figure 6.5.4. The rise of Logging Concession Licenses (HPHs), Timber Extraction Licenses (IPKs), and the rise and fall of legal logging. Source: Author’s analysis of Provincial government statistics, reported in Wurarah (2001).

the old Logging Concession Licenses (HPHs). Recent official estimates for total illegal logging in Papua range from as much as 600,000 m3 per month (Ministry of Forestry 2003a) down to 600,000 m3 per year (pers. comm., Head of the Papuan Provincial Forestry Service, who noted that the same volume of timber was logged illegally as was logged legally). Improved analysis is urgently required to assess the true extent of illegal logging. Exploratory mining leases cover a significant area of Papua, particularly in central and northern coastal mountain ranges (Chapter 6.4). Although several million hectares of forest are zoned for conversion to oil palm or timber plantations, the actual area of these activities remains quite limited (Chapter 6.4). Roads have been identified as a critical determinant of most forest loss around the world. Roads increase access to greater areas of forest, and lower the transport costs for loggers, farmers, and estate crop companies. The result is increased wildlife hunting, increased logging, and increased forest conversion. The effects can extend a considerable distance from the main roads themselves, as logging roads and local feeder roads splinter out into the surrounding forest. Analyses of road impacts typically consider a range of buffer sizes, from 10 km to 50 km or more (Mertens 2002a). Current and planned regional roads are shown in Figure 6.5.6. The buffer zones associated with these new and existing roads are show in Figure 6.5.7. The existing road network in Papua has already had great impact in the southeast, the Vogel-

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Figure 6.5.5. Locations and status of Papuan Licensed Logging Concessions (HPHs). Source: Mertens (2002a).

kop, and to the south of Jayapura. The length of existing national and provincial roads is about 2,700 km. The planned roads will increase this total by more than 1,250 km. Some new roads are needed in Papua, particularly to link existing population centers. Most of Papua is very remote; only 14% of the land is within 10 km of an existing road. The proposed new roads would increase that figure to 21%, though over half the land would be within 50 km of a road (Mertens 2002a). Figure 6.5.8 shows the portion of different land uses currently within 10 km of existing roads and explains how that portion increases if the new roads are built. Using a 10 km buffer, access to transmigration sites, limited and standard production forest, conversion forest, and timber and oil palm plantations all increase. Some of the increases are quite marked; the area of production forest and plantations within 10 km of a road goes up by 50% and 90%, respectively. The increase in plantation area within 10 km of proposed roads is at first glance surprising, because most oil palm plantations were established close to existing roads. In fact, the bulk of the increase comes from the improved access to large plantation areas in the far southeast provided by one proposed new road, the Okaba-Bade segment in Merauke. The area of watershed protection forest within 10 km of proposed roads also increases, though largely because of the proposed Wamena to Enarotali road cutting across the central mountain range. This road connection would link

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Figure 6.5.6. Existing roads and planned roads with the names of towns at either end. Source: Mertens (2002a).

the major population centers of Wamena and Jayapura to those on the southern coast, and may therefore have a strong social and economic case in favor of construction. Reassuringly, in terms of impact on areas protected for their biodiversity, the area of strict and standard nature reserves and national parks within 10 km of a proposed road would remain relatively unchanged. Roads, however, are only one mode of access, and using fixed buffers in an analysis does not capture the fact that a road’s influence may extend well beyond 10 km in easily traversed land, and less far in particularly rugged landscapes. Coastal roads may also have little impact on accessibility, because accessibility is already high due to easy access from the sea. More sophisticated methods can be used to analyze the history of land uses and draw on the resulting information to help predict what may happen in the future. One such analysis considered how proximity to rivers, population centers, and ports, topography, soil type, value of standing timber, and other factors had influenced logging in Papua to date (Mertens 2002b). This analysis needs to be refined and to include other factors (proximity to coastline in particular). Nonetheless the results demonstrate the value of the approach and highlight some of the areas

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Figure 6.5.7. Areas at different distances from existing and planned roads. Source: Mertens (2002a).

most at risk from future logging. The cost of extracting and transporting timber is shown in Figure 6.5.9. Note the incorrect identification of coastal areas such as offshore islands as ‘‘high cost,’’ an anomaly that results from not including proximity to the coast as a determinant of cost. This cost information is then combined with estimates of potential revenues to generate profitability estimates. A map showing an initial estimate is shown in Figure 6.5.10. The spatial modeling reported here highlights that although 70% of the inland swamp forests of the south coast are allocated to production forestry, few concessions were licensed in the area. This area is also relatively unaffected by existing and proposed roads. A large part of the western half of the Mamberamo River basin is similarly unaffected by the existing and proposed roads. Although much of the southern edge of the basin is within 50 km of the Wamena road, the topography is so rugged that much of it may be at low risk of logging. Further work is required to understand the spatial threat posed by logging. Nonetheless, it appears that large parts of both the southern swamp forests and the Mamberamo basin could be conserved with little economic loss. Even if logging profits are revised upwards, the lost income may have little impact because the Papuan government already has a large income from mining, and soon will from gas. Government investments to open up these areas to logging would be

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Figure 6.5.8. Percentage of land, by land use classification, that lies within 10 km of existing and proposed roads. Source: Author’s analysis of information from Mertens (2002a).

misplaced because other, more promising options exist, as the next section describes.

macroeconomic modeling of development options Spatial modeling gives a picture of how things are likely to look, but does not provide insights into what these developments will be worth in terms of GDP, job creation, and other common economic measures. For this more sophisticated computer-based macroeconomic models are required. One such model, Threshold 21 (T21), developed by the Millennium Institute, was modified for use in Papua (Qu et al. 2003). T21 has been used to explore vital development questions in many countries (Qu et al. 2003), and is particularly well suited to analyze the type of development questions facing Papua (Pedercini 2005). The T21 model uses standard economic production functions for different sec-

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Figure 6.5.9. Costs of extracting and transporting timber. Source: Hendi Sumantri (Conservation International) and Mertens (2002b).

tors and interlinks these with each other and models for other processes such as population growth. The models of each economic sector are drawn from established models used around the world by governments and development banks. The software platform utilized by T21 enables users to view the full structure of the model as a flow diagram (Figure 6.5.11). Parameter estimates are generally based on statistical analyses of historical data from Papua. Where assumptions are necessary, they are clearly identified and can be easily modified by local stakeholders. Four development options were identified for analysis (Sheng 2004). The first was a ‘‘business-as-usual’’ scenario. Under this scenario existing patterns of development were extrapolated, with some road and hydropower development, and slight increases in health and education spending. The second scenario—the ‘‘roads’’ scenario—included a significant expansion in the trans-Papua road network to be funded by increased logging. The third scenario explored the economic performance of the Mamberamo dam mega-development project championed by former President Habibie. This development scenario envisaged large investments in dam construction and hydropower in the Mamberamo river watershed. The power generated was to be used in aluminum smelters and other energy hungry industries that would be attracted to the region by low-cost power (Sheng 2004).

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Figure 6.5.10. Preliminary logging profitability estimates. Source: Hendi Sumantri (Conservation International) and Mertens (2002b).

Figure 6.5.11. Structure of the T21 model. Source: Millennium Institute, reported in Sheng (2004).

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These three scenarios were all being actively discussed in Papua in the late 1990s to early 2000s. Conservation International and the Millennium Institute proposed a fourth, ‘‘population centers’’ scenario that would focus development investments in improving government services and infrastructure in existing population centers, and improving access to credit for local entrepreneurs. These four scenarios describe realistic choices facing Papua, but they are not all-encompassing or mutually exclusive; Papua could follow one or more, or yet another strategy. The intentions in analyzing these four scenarios were threefold. The first aim was to demonstrate the value of such modeling for supporting development decision-makers. The second aim was to determine how current possible development options performed in isolation and thus determine whether massive infrastructure and mega-development projects performed better or worse than other options. The third aim was to verify whether or not there were plausible alternative development paths that could meet development objectives competitively while protecting forests. The model results should be interpreted as a ranking of various non-exclusive development options, and not an attempt to predict Papua’s economic future in absolute terms. The T21 model can generate many different indicators and measures of performance. Ideally the indicators and criteria chosen to evaluate development options would be chosen through a transparent and participatory process among local communities and stakeholders (Sheng 2004). This has yet to be done in Papua. In lieu of such agreed indicators, standards such as GDP and government debt were used to measure economic performance. Employment and locally retained income were used to estimate benefits to local people more directly, and pollution and forest land were used as indicators of environmental performance (Sheng 2004). T21 models an economy a number of years into the future. A comparison of the value of key indicators in 2020 is shown in Table 6.5.4. The results contain several key findings. First, the area of forest remaining in Papua in 2020 varies dramatically among the different scenarios. Under the ‘‘roads’’ scenario, forest cover is reduced the most, to about 8 million hectares. In contrast, the ‘‘population centers’’ scenario maintains over twice as much forest cover, about 17 million hectares. Forests would be cut quite quickly during the construction of the roads to 2015, after which time the rate of loss (hectares per year) slows (Figure 6.5.12). Second, although forest cover varies a great deal between these two options, GDP does not. Only the Mamberamo dam scenario generates a significantly greater GDP than other options (Figure 6.5.13). However, this comes at the cost of a significantly higher debt burden. Third, although the Mamberamo dam scenario generates significantly higher GDP, much of the revenue would leave Papua as returns to borrowed foreign capital and wages for highly qualified expatriates. In fact, the ‘‘population centers’’ scenario develops significantly more locally retained income or gross nationalregional product (GNrP) than the other scenarios (Figure 6.5.14).

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Rp billions, 1993 prices Rp billions, 1993 prices

GNrP

Government debt

Source: Millennium Institute, reported in Sheng (2004).

Rp billions, 1993 prices

Papua GDP

Economic

Composite, dimensionless

Millions of people

Employment

Pollution Index

Millions of hectares

Forests

Environment, Social

0

3,351

8,478

1.1

0.9

23.5

Size in 2000

Table 6.5.4. Comparison of development performance

0

7,020

19,076

1.7

1.1

15.9

Business as usual

11,006

7,120

23,068

2

1.2

13.4

2,933

7,084

19,621

1.7

1.1

8.3

Projected size in 2020 Mamberamo Dam Roads

0

7,799

20,315

1.8

1.3

17.1

Population centers

Natural Resource Economics of Papua / 1185

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Fourth, although the Mamberamo dam scenario generates more employment to 2015, investments in population centers should ultimately generate higher employment (Figure 6.5.15). In other words, not only can the development choices made today can conserve up to 9 million hectares relative to other options, they can also generate greater benefits for local people in terms of employment and locally retained income.

Discussion The information presented above suggests Papua can develop sustainably, and maintain most of its forest cover, without compromising economic growth, job creation, or poverty alleviation objectives. Just because this may be theoretically possible does not mean it will be easy to achieve. Success will require the right policies and macroeconomic conditions in Indonesia. More rapid growth can be expected if the Indonesian government successfully builds confidence nationally in public institutions, including security and enforcement agencies, maintains macroeconomic stability, and strengthens sectoral and investment priorities to invigorate the private sector (Baird 2002). Success will also require clear progress towards resolving the political and social conflicts over Papua’s status relative to Indonesia. The past treatment of Papua’s

Figure 6.5.12. Projected forest loss for each scenario. Source: Millennium Institute, reported in Sheng (2004).

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Figure 6.5.13. Projected GDP for each scenario. Source: Millennium Institute, reported in Sheng (2004).

Figure 6.5.14. Projected GNrP for each scenario. Source: Millennium Institute, reported in Sheng (2004).

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Figure 6.5.15. Projected employment for each scenario. Source: Millennium Institute, reported in Sheng (2004).

peoples has been shameful, including human rights abuses, misappropriation of land, and inequitable sharing of natural resource revenues. The longstanding struggle for independence has pitted the Indonesian National Army (TNI: Tentara Nasional Indonesia) against an array of pro-independence individuals and groups. Oppressive crackdowns, group punishment tactics, and persecution of suspected leaders are among a long litany and history of serious human rights abuses (e.g., Monbiot 1989; Blair and Phillips 2003). There is also a huge wealth gap between urban and rural people, and Papua as a whole received a small share of revenues from extractive industries. A greater share of and control over resources is needed to ensure revenues are shared more equitably and spent to alleviate poverty. The recent decisions to divide Papua into three provinces and many new districts may temporarily reduce the pressure for greater autonomy by distracting local leaders into discussions over how to share power under the new governance structures (International Crisis Group 2003). However, these maneuverings are unlikely to dissipate provincial-level demands for greater autonomy in the long term, unless as a result Papuans experience real benefits themselves in terms of improved government services, a greater say in their own governance, and greater economic opportunities. Unfortunately, the increased uncertainty brought about by political boundary and governance changes continues the confusion over which part and level of government has what powers and is responsible for what decisions and tasks. It is

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critical that the provision of government services is improved quickly, and that corruption and strong-arm tactics are dealt with rapidly. Unless these problems are addressed quickly local people will not see significant benefits, which will increase the likelihood that conflicts over Papua’s future will continue. Resolving Papua’s long-term status is a top priority, but opposing objectives are strongly held and finding a peaceable solution looks daunting. One analysis notes that independence supporters will only support a Special Autonomy agreement as long as they believe that it will not end efforts to achieve full independence. Yet on the other hand, those who oppose independence will not support any Special Autonomy arrangements that might further the independence movement (Blair and Phillips 2003). It is far from clear how these positions can be reconciled, but it is clear that significant political progress is unlikely until tension between the two opposing positions is diffused. One of the priorities for reducing tension is to change the role and behavior of the Indonesian military in Papua. The Indonesian military is chronically underfunded and—in line with official policy—makes up budget shortfalls by running its own businesses and receiving private payments for security services. They also have the dual role of maintaining internal security as well as defense from external threats. The changes needed are part of a broader effort to normalize the military according to modern practices, meaning refocusing it solely on defense against external threats (e.g., Blair and Phillips 2003). Two immediate priorities are being discussed for action in Papua. First is to end the role of the military in providing security for large extractive industry operations, and end the millions of dollars in payments to the military by the companies involved (e.g., Global Witness 2005). Second is to reduce the incidents of human rights abuses by placing tight operational limits on the worst performing elements of the military, strengthen internal policing of abuses, and improve overall training and conditions for military personnel (Blair and Phillips 2003). Progress on resolving political and social conflicts in Papua is necessary but not enough to ensure successful sustainable development, or conservation of Papua’s forests. Successful sustainable development will also require better economic and social policy and budget choices in Papua. Although many decisions are beyond the sole control of Papuan local governments under the current decentralization situation, many government roles and responsibilities have shifted to the local level and local governments can significantly affect Papua’s future. The quality of governance in local governments is therefore critical, and depends on capacity building and rooting out bad practices and corruption. The policies and budgets adopted at local levels will also have a great impact, not only in terms of the general direction of development discussed at length above, but also in how they help or hinder the private sector and the delivery of public services to society as a whole. The massive mining and gas revenues are themselves a double-edged sword. If they are well spent they can help Papua develop sustainably and become prosperous. If spent badly then the opportunity to improve conditions in Papua may be

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missed. In a worst-case scenario, poor government policies and misspending of government revenues may also exacerbate environmental damage. Good options do exist, as the analyses presented here show. However, successfully delivering these choices means ensuring that (1) development options are evaluated properly and transparently, (2) government decisions are made to pursue these options, and (3) the capacity exists to implement these decisions effectively. Programs to provide technical training and support participatory dialogue can help evaluate development choices. The Forum for Conservation and Development in Papua (FKPTP: Forum untuk Konservasi dan Pembangunan di Tanah Papua) described earlier is one such example, with a focus on carrying out more technical analyses. Several complementary programs are underway, including a joint program by USAID, the University of Papua, and British Petroleum called the Bird’s Head Alliance to strengthen government capacity and increase participation in decision making (British Petroleum 2004). The impact of these complementary programs can be increased through greater support and coordination. Ensuring government decisions are consistent with the better choices is a far greater challenge. The major challenges facing Papuan government authorities trying to make good choices include corruption, lack of transparency in decision making, and self-serving political leaders unconstrained by weak democratic oversight. These challenges are not unusual around the world, and a range of proven responses exists. These include actions such as support for civil society in a watchdog role, support for a free press, creation of publicly appointed investigative bodies (e.g., ombudsmen), careful auditing of donor-funded government projects and services, funding and other support to create pro-reform alliances, and training in democratic values. Rooting out corruption in local government is arguably the most urgent priority for Papua. This may ultimately require changes in the Indonesian government and criminal justice system, but in the meantime alternative governance and oversight structures may reduce corruption. According to the World Bank, ‘‘communitybased projects, such as the Kecamatan Development Program, have shown how local decision making and transparency can help control the misuse of public funds’’ (Baird 2002: 1). The large scale of revenues up for grabs from the natural resources sector can also exacerbate corruption. Various safeguards can be put in place to reduce the scope for corruption. These include installing independent committees with oversight over spending the revenues, or simply earmarking percentages of the revenues for spending in certain sectors such as health, education, and basic infrastructure. A recent example of such responses comes from Chad, where the World Bank has taken several measures to try and ensure the government spends oil revenues efficiently and transparently (World Bank 2003b). Even if governments make the right decisions, they must still implement them effectively in order to reap the full benefits. This highlights the need for investments in government capacity in basic budget management, delivering health and educa-

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tion and other services, and running public institutions. Strengthening civil society helps citizen groups to participate, demand, and assist in providing these services. These systematic system-wide improvements will take time to have effect. Yet rapid successes are needed to rebuild faith in government by satisfying the most desperate poverty reduction needs and rectifying the worst injustices that underpin and feed social unrest. The need for rapid economic progress points towards prioritizing investments in the sectors and locations where the need is greatest and the marginal returns are highest. Environmental benefits will also arise from investing in the most deprived communities, because such investments will reduce local community and government pressure to seek revenues from other sources, primarily logging. Prioritizing investments towards those most in need will also contribute to reducing concerns over how the wealth from natural resources will be distributed. Natural resource wealth is not evenly distributed across Papua, and communities far from major mines, towns, or gas fields are greatly concerned about getting their share of the wealth. Wealth is unlikely to be distributed through employment, because the numbers employed are limited relative to the available work force and few Papuans have technical training needed for more highly paid jobs. Therefore another priority need in Papua is the development of mechanisms for the fair distribution of wealth from natural resources. Until these distribution mechanisms are in place, local governments are likely to accelerate the use of the natural resources they do have, generally by logging their forests. Thus the absence of a fair method for sharing Papua’s wealth gives mineral- and gas-poor areas the incentive to liquidate their other resources more quickly. Other short-term priorities may include reversing some aspects of transmigration and supporting small- and medium-size business enterprises. The large numbers of rural poor in Papua arise in part because of the large number of poor settlers, particularly those who arrived as part of Indonesia’s transmigration scheme and were often relocated according to official resettlement schemes in extremely remote and inhospitable areas. The authorities should consider closing down transmigration sites that are involved in conflicts with local Papuans, or where economic conditions are particularly awful. Small- and medium-size businesses are also vital to the future success of Papua. They will flourish as government performance improves, but specific measures can also help in the short term, such as adopting more business-friendly policies and establishing lines of credit for local firms. This discussion has so far focused on what government can do to improve circumstances in Papua. The responsibility does lie mainly with the Papuan government. However, the private sector must also play a part, particularly Freeport and British Petroleum (BP). The two major international corporations active in Papua have a critical role to play in helping Papua deliver these short-term priorities and make progress longer-term goals. Both are taking steps to do so. Freeport has made a number of environmental and social commitments (Freeport-

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McMoRan Copper and Gold, Inc. 2004), as has BP (British Petroleum 2004), and they have both funded a range of social development programs in and around their field operations. Both Freeport and BP correctly emphasize they are not governments, and should not take on roles of government such as providing education and health services. Both companies emphasize that they are in Papua as commercial partners to the Government of Indonesia, and argue this limits the extent of engagement with government (e.g., British Petroleum 2005). However, both companies will be in Papua for decades to come and, as major parts of the fabric of society, they can and do influence government. Hence they can legitimately support programs that aim to improve government performance and governance, and apply pressure to government to secure these improvements. Indeed BP has undertaken efforts to improve government capacity through the Bird’s Head Alliance, mentioned above. Efforts to improve the quality of government should be extended across all of Papua, not only because this is the appropriate thing for a responsible company to do, but also because it will help improve stability and contribute directly to corporate performance.

Conclusions The future of Papua’s forests depends on the development path Papua decides to pursue. If Papua chooses extensive road development then there is a risk that forest cover may be reduced by over 60% from current levels. Viable alternative development options can conserve large forest areas while meeting development goals. The revenues from Papua’s mining and gas industries should mean Papua has the funds to pursue these alternative development options. Spatial modeling techniques can help identify areas at varying degrees of risk. Areas at low risk of development or which have already been selectively logged can be protected without trading off progress towards development objectives. Conservation investments can then focus on areas at high risk that have value for development but are essential to conserve in order to protect as much of the full suite of Papua’s biodiversity as possible. The spatial and macroeconomic modeling results presented here make a useful contribution towards identifying sustainable development options and priorities for conservation. Significant investments have already been made to develop these powerful tools, but they require further development and fact checking to fulfill their true potential. Both modeling approaches must be strengthened and extended through participatory work with Papuan academics and experts from other sectors. Local stakeholders may wish to add other criteria for evaluating the performance of different development choices, such as the contribution to promoting stability and security, which may outweigh growth, poverty reduction, or other typical economic measures of performance. Putting accurate information on development options into the public domain and promoting participation and transparency will not guarantee decisions in

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favor of conservation. However, broader understanding and participation will help ensure a broader spread of development benefits, help ensure consideration of the needs of underrepresented groups, including poor and remote communities, and help avoid development projects that have negative environmental impacts while benefiting only a few well-connected stakeholders. Further work is needed to identify different stakeholders and determine how their interests will be affected by alternative development choices. Development choices will need to be refined and additional activities added to mitigate negative impacts on different stakeholders and thus to overcome their opposition. Efforts must be made to build ownership, to encourage government to make the right decisions, and to build government capacity so that these decisions are implemented effectively. The Government of Indonesia and local governments in Papua must take the lead, with the international community providing support through major programs of capacity building of government and civil society, investments in priority poverty reduction efforts, and promotion of good governance. If these development goals can be achieved in Papua, then so too can conservation goals.

Literature Cited Anggraeni, D. 2001. Interim report of resource economic assessment in Irian Jaya Province. Conservation International Indonesia Program, Jakarta. Anggraeni, D. 2005. Rapid assessment for conservation and economy (RACE) in Papua: a summary. Conservation International Indonesia Program, Jakarta. Baird, M. 2002. Farewell remarks by Mark Baird, World Bank Country Director for Indonesia. Jakarta Foreign Correspondents Club, August 27, 2002. Blair, D.C., and D.L. Phillips. 2003. Indonesia commission: peace and progress in Papua. Report of an independent commission sponsored by the Council on Foreign Relations Center for Preventative Action. Council on Foreign Relations, New York. British Petroleum. 2004. Towards a brighter future: Tangguh integrated social strategy review 2004. BP Berau Ltd., Jakarta. British Petroleum. 2005. BP Response to the TIAP report of February 2005. BP, London. Available at http://www.bp.com. Conservation International. 2005. Papua Provincial Government supports establishment of the Forum for Conservation and Development in the Land of Papua (FKPTP). Press release no. 029/CI-PP/IV/05, 30 March 2005. Conservation International Indonesia Program, Jakarta. Dwiyana, A. 2001. Oil, gas and mining development and decision-making process in Irian Jaya. Conservation International Indonesia Program, Jakarta. Easterly, W., and R. Levine. 2002. Tropics, germs and crops: how endowments influence economic development. Working paper no. 15. Center for Global Development, Washington, D.C. Freeport-McMoRan Copper and Gold, Inc. 2004. Making the commitment: 2004 working towards sustainable development report. Freeport-McMoRan Copper and Gold, Inc., New Orleans. Gallup, J.L., J.D. Sachs, and A. Mellinger. 1999. Geography and economic development. Working paper no. 1. Center for International Development, Harvard University, Cambridge, Massachusetts.

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1194 / james b. cannon Gautam, M., U. Lele, H. Kartodihadrjo, A. Khan, Ir. Erwinsyah, and S. Rana. 2000. Indonesia: the challenges of World Bank involvement in forests. Evaluation Country Case Study Series. Operations Evaluation Department, World Bank, Washington, D.C. Gesellschaft fu¨r Technische Zusammenarbeit (GTZ). 2003. Decentralization in Indonesia since 1999—an overview. Deutsche Gesellschaft fu¨r Technische Zusammenarbeit (GTZ) GmbH. Available at http://www.gtzsfdm.or.id Global Witness. 2005. Paying for Protection: The Freeport Mine and the Indonesian Security Forces. Global Witness Publishing, Washington, D.C. Indonesia Ministry of Industry. 2005. Indonesia Review, Department of Trade. Edition 6, Dec 2004. Ministry of Industry, Jakarta. International Crisis Group. 2003. Dividing Papua: how not to do it. Indonesia briefing, April 2003. International Crisis Group, Jakarta/Brussels. Jepson, P., J.K. Jarvie, K. MacKinnon, and K.A. Monk. 2001. The end for Indonesia’s lowland forests? Science 292: 859–861. Mertens, B. 2002a. Spatial analysis for the Rapid Assessment of Conservation and Economy (RACE) in Irian Jaya: Report 1. Center for International Forestry Research report for Conservation International, Washington, D.C. Mertens, B. 2002b. Spatial analysis for the Rapid Assessment of Conservation and Economy (RACE) in Papua: Report 2. Center for International Forestry Research report for Conservation International, Washington, D.C. Ministry of Forestry. 2003. Press Release No. 51. Indonesia Ministry of Forestry, Jakarta. Mollet, J.A. 2001. District Budget Collection under Decentralization and the Impacts to the Natural Resource in Irian Jaya. Cenderawasih University, Jayapura, Indonesia. Monbiot, G. 1989. Poisoned Arrows: An Investigative Journey through Indonesia. Penguin Group, London. Nugroho, B. 2001. A Review of Forest Policies in Papua. Conservation International Indonesia Program, Jakarta. Pedercini, M. 2004. Evaluation of alternative development strategies for Papua, Indonesia. Master’s thesis, Information Science Department, University of Bergen, Norway. Pedercini, M. 2005. Potential Contributions of Existing Computer-Based Models to Comparative Assessment of Development Options. Conservation International, Washington, D.C. Qu, W., G. Barney, and M. Pedercini. 2003. Documentation for the Threshold 21 (T21) Integrated Development Model, 8th Edition with Additions from Mozambique and Papua. Millennium Institute, Arlington, Virginia. Sachs, J.D., N. Bajpai, and A. Ramiah 2002. Understanding regional economic growth in India. Working paper no. 88. Center for International Development, Harvard University, Cambridge, Massachusetts. Sheng, F. 2002. Biodiversity conservation and economic development in the Indonesian Province of Papua under the fiscal decentralization and the Special Autonomy: an integrated issue papers report. Conservation International, Washington, D.C. Sheng, F. 2004. Comparative Assessment of Development Options. Conservation International, Washington, D.C. Sugiono, B. 2002. Analysis of Decision-Making Process in the Development Planning and Implementation in Papua Province. Conservation International Indonesia Program, Jakarta. U.S. Embassy, Jakarta. 2005. LNG report: troubles in Indonesia’s LNG industry. Energy News, June 6, 2005. Embassy of the United States, Jakarta. Wiratno. 2002. Institutional Capacity Assessment. Conservation International Indonesia Program, Jakarta.

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Natural Resource Economics of Papua / 1195 World Bank. 2001. Poverty reduction in Indonesia: constructing a new strategy. Report no. 23028-IND. World Bank, Washington, D.C. World Bank. 2003a. Decentralizing Indonesia: a regional public expenditure review overview report. Report no. 26191-IND. World Bank, Washington, D.C. World Bank. 2003b. The Republic of Chad: country assistance strategy. Report no. 26938CD. World Bank, Washington, D.C. Wunder, S. 2003. Oil Wealth and the Fate of the Forest. Routledge, London. Wurarah, R. 2001. Issue paper on forestry sector. University of Papua, Manokwari.

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section seven 

Conservation of Papuan Natural Resources

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7.1. Threats to Biodiversity scott frazier n d on e s i a’ s e a ste r n mo s t p ro v i n ce of Papua (formerly known as Irian Jaya) comprises the western half of the world’s second-largest and highest tropical island, New Guinea. Any discussion of the geophysical and ecological evolutionary history of New Guinea (and its satellites) is applicable to Papua and to its neighbor on the eastern half of the island, the nation of Papua New Guinea (PNG). Extending over approximately 421,981 km2, Papua encompasses the broadest spectrum of landscape forms, ranging from coral reef and coastal habitats, vast inland freshwater swamps and lowland forests to montane forest and even snow-capped peaks. One protected area in Papua, Lorentz National Park, a World Heritage Site (see UNEP-WCMC 2001) captures this range of landscapes: ‘‘[Lorentz] is the only protected area in the world to incorporate a continuous, intact transect from snowcap to tropical marine environment, including extensive lowland wetlands’’ (UNESCO 2005). To explain this spectacular topographical range, it is necessary to touch upon plate tectonics briefly. New Guinea resides at the northern margin of the relatively stationary Australian continental plate, which has been subjected to 40 million years of collisions with a series of southward-migrating island arcs along the boundary with the Pacific Plate. Each of these arcs has had a separate tectonic history and carried a correspondingly different biota (Polhemus and Polhemus 1998; Chapter 2.1). This long continuous tectonic sequence of events has resulted in the orogenesis that so characterizes the New Guinea island, yielding an extreme topography of abrupt terrain transitions—mountains, escarpments, valleys, and gorges—giving way to vast swaths of lowland forests and swamps, crossed by myriad rivers. These landscape features have acted as barriers to migration and movement of various species (Supriatna 1999), and ensured that there were a plethora of geobiological isolates in which independent evolution could run its course. Because of this geologic history, mainland Papua can be roughly divided into four geographical zones, each with a distinctive biota (Muller, in prep.). The first, a Central Cordillera, divides the entire island, north and south. Vast lowland areas occur on either side of this range (the second and third zones). This includes New Guinea’s largest and least-disturbed tropical humid forest catchment—the Mamberamo Basin—which is essentially, the entire northern watershed of western New Guinea. The Mamberamo Basin encompasses nearly 8 million hectares, and is more than 93 percent forested (Forest Watch Indonesia 2004), making it a vast, intact storehouse of globally significant biodiversity. South of the Central Cordillera is found one of the world’s largest freshwater wetlands and one of the world’s most extensive mangrove systems. The fourth zone, the Vogelkop (or Bird’s

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

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Head), is connected to the rest of New Guinea by a rugged narrow and curving isthmus. There are a number of isolated mountain ranges punctuating the lowlands (e.g., Papua’s Cyclops, Van Rees, and Foja mountains). Outlying islands form separate and distinct ecological units (e.g., the Raja Ampat Islands, Biak, Yapen, and Aru) in Papua, as they do on the eastern Papua New Guinea side of the island. The Raja Ampat Islands, at the western end of the Vogelkop, provide Papua’s most striking example of rich and diverse coastal and marine ecosystems in terms of size, pristine condition, and heterogeneity. Raja Ampat resides at the heart of the Coral Triangle, the earth’s center of coral reef biodiversity (Chapter 5.2). Thus the tectonic and volcanic forces that have shaped New Guinea’s landscape diversity, along with its size and equatorial position within the elongate chain of archipelagoes stretching from Asia into the southeast Pacific, have given rise to a great range and mosaic of natural ecosystems populated by this extraordinary terrestrial and marine biodiversity (see Chapters 4.1–4.10, 5.1–5.13). New Guinea is important to global biodiversity because it supports the third largest contiguous tract of tropical ‘‘High Biodiversity Wilderness,’’ behind Amazonia and the Congo Forest. Papua also contributes nearly 50 percent of the biodiversity that makes Indonesia the world’s most biodiverse country (Supriatna 1999) according to some estimates. ‘‘Wilderness’’ areas are more than 70 percent intact, and are typically under less pressure from encroaching human populations and landscape scale conversion than areas like ‘‘Biodiversity Hotspots’’ (Mittermeier et al. 2002). Hotspots are areas that continue to harbor exceptional concentrations of endemic species while undergoing exceptional loss and degradation of habitat (Myers et al. 2000). New Guinea is centered between the Wallacea Hotspot in the west, and the East Melanesian Islands Hotspot (encompassing the Bismarck Archipelago and Admiralty Islands, the Solomon Islands, and the islands of Vanuatu) to the northeast and east (See Mittermeier et al 2005). Given that only three major tropical High Biodiversity Wildernesses are left on the planet, this distinction between ‘‘Hotspots’’ and ‘‘Wildernesses’’ may lose its relevance as threats increasingly gravitate to these rich areas. For example, as forests have become depleted in Sumatra and Kalimantan in Indonesia (see World Bank Group 2001), loggers are shifting their operations to Papua (Asia-Pacific Peoples Environmental Network 2000, 2002b; International Crisis Group 2002; Environmental Investigation Agency and Telapak 2005). Recent data indicate that Papua still contains 30.4 million hectares of natural forest cover, which is equivalent to 73% of the province (Forest Watch Indonesia 2004). It is indeed the challenge of conservation and sustainable development in Papua to maintain existing wilderness ecosystem ‘‘connectivity,’’ and to avoid the drift towards becoming a fragmented Biodiversity Hotspot. While the natural factors which have given rise to the great biodiversity of Papua apply to both sides of the island of New Guinea, it is also logical that the threats facing biodiversity across New Guinea are similar. However, in some categories of threat, Papua and Papua New Guinea diverge.

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What Is Threatened? ‘‘Biodiversity’’ is defined, and it is delineated and characterized for Papua, in a number of chapters in this volume. The case for the importance of this biodiversity is also presented elsewhere, and it is taken as a given here. Briefly, biodiversity is important to people because it supports their livelihoods and welfare directly, through use of biological resources, or indirectly, through its contribution towards the maintenance of ecosystem functions (Christie et al 2004; Chapter 5.1) that provide services that benefit humanity. High Biodiversity Wilderness areas like Papua provide services like watershed protection and climate regulation which help maintain this biodiversity at a broader level, across their regions and across the globe. These values are in addition to the notion of an intrinsic value of biodiversity. This chapter looks at threats to Papua’s biodiversity. But what is this ‘‘biodiversity’’ that is threatened? While the general concept of biological diversity evolved from ‘‘number of species present’’ to the notion that biodiversity is much more complex than that (e.g., as defined under the Convention on Biological Diversity), perhaps ‘‘biodiversity is best considered at three levels: ‘genetic’, ‘organismal’, and ‘ecological’ (or ‘community’) biodiversity’’ (Christie et al. 2004, citing Harper and Hawksworth 1995). An assumption is made here that Papua, as more or less half of the New Guinea High Biodiversity Wilderness, encompasses high biodiversity at all three levels. And depending on the type and scale of threat, one or more levels of biodiversity may be adversely affected.

Kinds of Threats Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms (Chapin et al. 2000; J. A. Thomas et al. 2004). While species extinction is a natural phenomenon, currently the rate of extinction has accelerated far beyond ‘‘background’’ rates. An often cited figure is that the current extinction rate is now approaching 1,000 times (or even 10,000 times) the natural background rate of extinction (Stuart 1999; Pimm 1997). And this is often coupled with a prediction that this rate will further increase dramatically (Purvis et al. 2000). As early as 1997, the World Resources Institute reported that more than 80 percent of the earth’s large, intact natural forests (starting from 8,000 years ago) had been destroyed (Bryant, Nielsen, and Tangley 1997). The pressures and threats on the remaining 20 percent of these forests have escalated dramatically in the present decade. Increasingly, global trade and communication are directly contributing to the mixing of faunas and floras that were previously separated by biogeographical boundaries. To describe this new epoch of widespread human influence, some researchers have suggested the term ‘‘Homogocene’’ (IUCN/SSC 2005) or ‘‘Anthropocene’’ (Crutzen 2002). Sadly, myriad threats confront the earth’s remaining biodiversity. Papua is not immune or insulated.

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Biodiversity threats can be grouped and studied in a number of ways, and threats have been variously systematized depending on the needs of those studying or dealing with them. A number of ‘‘threat classifications’’ have been developed (e.g., Thorpe and Godwin 1999). Probably the best known example is the ‘‘threat authority file’’ used in concert with the 2004 IUCN Red List of Threatened Species (IUCN 2004). BirdLife International (2005) uses this file as a basis for its threat classification. A simplified version of the threat authority file classification appears in Box 7.1.1. Threats to biodiversity can come from the ‘‘top-down’’ and collectively from the ‘‘bottom-up.’’ Biodiversity loss and degradation are cumulative and cascading (e.g., food-web relationships can be adversely affected by loss of a particular species), and threats typically work in synergy (Hannah and Lovejoy 2003). There are threats that operate at a global level and those that, taken together collectively, constitute a global threat. An example of a threat that operates globally (i.e., one that is ‘‘top-down’’) is climate change. However, micro-climate can also be affected by a host of local activities. For example, clear-cut logging of a large block of forest increases local temperature, with attendant detrimental impact on certain local biotic and abiotic elements, beyond the primary consequence of habitat loss. Climate change driven by an increase in greenhouse gases (as precipitated and exacerbated by numerous and often disparate activities) is, however, a global phenomenon with global and cumulative consequences for biodiversity (e.g., loss of productive coastal wetlands due to global sea rise). And having a global impact, climate change has consequences all the way down the rest of the top-down continuum, from the landscape level to the local level. So, in perhaps a perverse turn of phrase, the global threat ‘‘acts locally.’’ Landscape threats (e.g., primary forest conversion to oil palm plantations) and local threats (e.g., invasive species, at least at the initial stages of their invasion), may be global in scope and in cumulative impact, but they act first directly on the biodiversity landscape at some component level (albeit affecting variously different size pieces of the mosaic). Measuring biodiversity, and therefore loss of biodiversity, has engendered quite some debate among scientists (Christie et al. 2004; Myers 1997; Willis, Gillson, and Brncic 2004). For example, are all species equal or are some ‘‘more equal’’ (more important) than others (e.g., keystone species)? While these discussions go on it is important to remember that threats that occur at global, landscape, or local scale can have detrimental impact on biodiversity (under particular scenarios) across multiple levels (e.g., genetic, species, ecosystem), given that biodiversity too is measured and distributed unevenly. Most benefits and services that derive from the natural interactions of species in an ecosystem are local and regional. If a keystone species is lost from an area, a dramatic reorganization of the ecosystem can occur. Localized extinctions may be just as significant as the extinction of an entire species worldwide (Mock 2000). A local threat realized upon a species over the territory of its restricted range—in other words, the potential loss of an en-

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SIMPLIFIED IUCN THREAT AUTHORITY FILE (THREATS TO TAXA) 0. No threats 1. Habitat loss/degradation (human induced) 1.1. Agriculture 1.2. Land management of non-agricultural areas 1.3. Extraction 1.4. Infrastructure development 1.5. Invasive alien species (directly impacting habitat) 1.6. Change in native species dynamics (directly impacting habitat) 1.7. Fires 2. Invasive alien species (directly affecting the species) 2.1. Competitors 2.2. Predators 2.3. Hybridizers 2.4. Pathogens/parasites 3. Harvesting (hunting/gathering) 3.1. Food 3.2. Medicine 3.3. Fuel 3.4. Materials 3.5. Cultural/scientific/leisure activities 4. Accidental mortality 4.1. Bycatch 4.2. Collision 5. Persecution 5.1. Pest control 6. Pollution (affecting habitat and/ or species) 6.1. Atmospheric pollution 6.2. Land pollution 6.3. Water pollution

7. Natural disasters 7.1. Drought 7.2. Storms/flooding 7.3. Temperature extremes 7.4. Wildfire 7.5. Volcanoes 7.6. Avalanches/landslides 8. Changes in native species dynamics 8.1. Competitors 8.2. Predators 8.3. Prey/food base 8.4. Hybridizers 8.5. Pathogens/parasites 8.6. Mutualisms 9. Intrinsic Factors 9.1. Limited dispersal 9.2. Poor recruitment/reproduction/ regeneration 9.3. High juvenile mortality 9.4. Inbreeding 9.5. Low densities 9.6. Skewed sex ratios 9.7. Slow growth rates 9.8. Population fluctuations 9.9. Restricted range 10. Human disturbance 10.1. Recreation/tourism 10.2. Research 10.3. War/civil unrest 10.4. Transport 10.5. Fire 11. Other1 12. Unknown1

The categories ‘‘other’’ and ‘‘unknown’’ also occur under each main threat category but have been removed here to save space.

1

Source: After IUCN (2005).

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demic species—has a significant impact on biodiversity at that local scale, as well as at a global (genetic) scale. Table 7.1.1 provides a sample of a number of generalized ways in which threats could be categorized or described. While we often take a dichotomous approach to contrasting threats, in reality threats occur across a continuum, and there are often underlying causes of threats that are not as conspicuous as what appears to be driving them at the surface. Immediate threats to biodiversity often stem from societal problems like human population pressures, material consumption, poverty, and inequitable access to resources (BirdLife International 2004). As previously mentioned, seemingly discrete threats do not usually work in isolation, but often work in perverse synergy, operating across multiple scales. Anthropogenic threats can exacerbate natural threats (e.g., cutting of mangroves removes a layer of natural protection against sea surges, tsunamis, etc.). Whatever the classification of threats, Papua, encompassing half of the world’s third largest expanse of High Biodiversity Wilderness, is not immune. On the contrary, it is clear that the threats facing biodiversity in Papua are manifesting themselves across a scale never seen before, including endogenous threats. While archeological and paleoecological evidence suggests that tropical forests, including those in New Guinea, have been ‘‘disturbed’’ for millennia (Willis, Gillson, and Brncic 2004), the changes stemming from those basic and low-intensity human activities cannot be compared in scale or magnitude to the landscape-scale changes presently occurring in the forests of Papua. It is unequivocal that Papua’s biodiversity is now under assault, from both within and outside. And while many of the threats confronting Papuan biodiversity are typical of other places, there are also those that might be considered, if not uniquely Papuan, perhaps emblematic of Papua.

Table 7.1.1. Ways to contrast biodiversity threats Typical threat

vs.

Unique threat

vs.

Other threats

Global threat

vs.

Landscape threat

vs.

Local threat

Direct threat

vs.

Indirect threat

Direct threat

vs.

Major social, institutional and infrastructural threats (Thaman 2002)

Terrestrial threat

vs.

Marine threat

Natural threat

vs.

Anthropogenic threat

Physical changes

vs.

Human activities behind the physical changes

vs.

Incentives and capacity that determine human activities (Anggraeni 2005)

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Threats in Papua Papua, as we have seen, has been blessed with a superlative natural bounty of vast wilderness areas and incredible landscape diversity giving rise to high biodiversity on land and in its lakes, rivers, and streams. This is matched by spectacular marine biodiversity in its seas. Papua has an abundance of precious metals, natural gas, and oil. It is also Indonesia’s largest and least populated province. By 2000, the population of Papua was just over 2.2 million people, with more than 70% living in rural areas. Within Papua, there are ca 250 ethnic tribes with their own languages and traditional systems (Anggraeni, 2005; Chapter 1.3). But the rapacious appetites that have depleted timber and other resources elsewhere have turned this, until now, relatively untouched bounty into a magnet for a range of threats. The immense potential for profit from extracting these resources creates a tremendous incentive to exploit them (Millennium Institute 2003). The world’s third largest expanse of High Biodiversity Tropical Wilderness, by definition less threatened than areas like Hotspots, is in fact seriously threatened in the medium term, simply because it is one of the last such areas where these resources can still be found in abundance. With that extractive pressure as a backdrop, and the ongoing and ever escalating global climate change crisis, punctuated with a host of more localized but not necessarily less worrisome typical threats, Papuan biodiversity is in for a rough ride. A highly systematized, detailed, and complete analysis of threats is beyond the scope of this chapter. Instead, a number of the most serious threats to the biodiversity of Papua will be discussed in the following sections.

logging: the mad rush to papua Forest loss is raging across the planet (Bryant, Nielsen, and Tangley 1997). Recent figures show that Indonesia alone has lost 18 million hectares of tropical forests between 1985 and 1997 on just three of the major islands—Kalimantan, Sumatra, and Sulawesi (Putz et al. 2000). Environmental Investigation Agency and Telepak (2005) report that forest area equivalent to the size of Switzerland (41,290 km2) is lost every year from Indonesia. Put another way, this is equivalent to a forest loss of nearly one-tenth of Papua’s land area per year. Most of this forest loss has occurred in lowland forest and logging has played a major (but not the only) role in that destruction. See Some Biodiversity Impacts of Forest Loss, next page. In 1997, actual forest cover in Papua was estimated to be just over 33 million hectares (Forest Watch Indonesia/Global Forest Watch 2002, citing Holmes 2000). This had dropped to 30.4 million hectares by 2002 based on GIS/remote sensing measurements (Forest Watch Indonesia 2004). Of all of the threats facing Papuan biodiversity, forest loss is most serious in both the short and long term. ‘‘By 2010, Irian Jaya [Papua] is likely to be the only part of Indonesia with any significant areas of undisturbed natural forest’’ (FWI/GFW 2002). This is the corollary to the predictions of imminent doom for the future of lowland forests in Sumatra and Kalimantan (See World Bank Group 2001).

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SOME BIODIVERSITY IMPACTS OF FOREST LOSS • Loss of species habitat and local extirpation or extinction of species • Fragmentation of species habitat disrupting species cycles and ecological processes • Disruption and degradation of ecosystem services that maintain and support biodiversity • Increased erosion and sedimentation with deleterious effect on aquatic and marine ecosystems and biodiversity • Increased risk of wildfire • Large release of carbon, further exacerbating climate change with adverse effects on biodiversity ‘‘. . . complex non-linear ecological structures of hydrological, climatic, geochemical and biological importance are being lost, in most cases, forever’’ (Thompson 1996).

According to the ‘‘Down to Earth’’ Newsletter (Asia-Pacific People’s Environment Network 2000), quoting the Provincial Environmental Impact Management Agency (BAPPEDALDA: Badan Pengelolaan Pengendalian Dampak Lingkungan Daerah), some 57 timber companies were logging 11 million hectares in Papua. The total area designated for logging was said to be 10 million hectares, indicating that other areas, presumably areas of protected forest, were included in the concessions. (Similarly, the Environmental Investigation Agency and Telapak (2005) cite illegal logging incursions in Salawati Nature Reserve, Bintuni Bay Strict Nature Reserve, and Cenderawasih Bay National Park.) Papua’s Provincial Environmental Impact Management Agency (BAPPEDALDA) named 16 companies with concessions over 200,000 hectares, with a combined total of 6,812,770 hectares under their control. International Crisis Group (2002) reported 13 million hectares under concession (including hundreds of smaller-scale concessions). Meanwhile, Conservation International Indonesia-Papua Program (2004a) research indicated that as of 2000, the area designated as production forest was almost 22 million hectares, with almost 12 million hectares granted to 54 logging concessions (11% of this overlapped with conservation and protected areas). Similarly, Forest Watch Indonesia (2005) lists 58 active forestry and plantation concessions covering 9.6 million hectares, according to data from 2003. The total of all active and nonactive concessions exceeds 13.5 million hectares, according to this data source. While these numbers may vary according to the year and data set, the result is that at a conservative minimum, at least one-third of Papua’s forests are slated for logging under one type of concession or another. While some companies have run into determined resistance from indigenous landowners resulting in inactive concessions (Asia-Pacific People’s Environment Network 2000, 2002b; International Crisis Group 2002), the fact that there is nowhere else to turn for logs

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(World Bank Group 2001) means that pressure on Papua’s forests can only grow. Indeed, if the somewhat controversial figures released by the Indonesian Forestry Department (Departemen Kehutanan 2003) and widely carried in the national press were correct, then Papua was losing a staggering 600,000 cubic meters of logs per month to illegal logging alone in 2003. The situation appears to have perhaps even worsened if one considers new data from Environmental Investigation Agency and Telepak (2005) that illegal export of a just single type of timber— ‘‘merbau’’ (Moluccan ironwood: Intsia bijuga and I. palembanica)—from Indonesia (almost entirely from Papua) is approximately 300,000 cubic meters a month! Not only lowland rainforests have suffered loss. Indonesia probably contains the most extensive mangrove forests in the world (Muller, in prep; Chapter 5.4), estimated at 4.25 million hectares in the early 1990s (Forest Watch Indonesia/ Global Forest Watch 2002). Southern New Guinea, including Papua, has the greatest diversity of mangrove species in the world (Muller, in prep.; Chapter 5.4). But mangrove forests throughout Indonesia have also been under serious pressure. The Indonesian government reported that it lost some 1 million hectares of mangroves between 1969 and 1980 alone (Forest Watch Indonesia/Global Forest Watch 2002, citing BAPPENAS 1993). World Resources Institute (Mock 2000) compiled an estimate of 2.4 million hectares of existing mangroves in Indonesia, representing an approximate 55% loss since the 1980s. Mangroves are found throughout Indonesia, and more than 50% of those remaining are located in Papua (British Petroleum 2002), home of the world’s third-largest mangrove area at Bintuni Bay (The Nature Conservancy 2004). This formerly pristine mangrove measuring about 450,000 ha (BP 2002; The Nature Conservancy 2004) underwent illegal logging in the 1980s (International Crisis Group 2002). The mangrove continues to be threatened by adjacent timber extraction, as well as by local, often small-scale wood chip production, by conversion of mangrove areas to shrimp ponds, by agriculture, or by salt pans, energy exploration, and pollution (BP 2002). Economic development in the Bintuni area is increasing due to the opening of a new natural gas field in Bintuni Bay, and the population is expanding rapidly (The Nature Conservancy 2004). Less than a decade ago the prawn fishing industry in Bintuni Bay was valued at US$ 35 million, but the destruction of a large area of mangrove for wood chip milling resulted in a drastic decline in catch, and the industry shifted elsewhere (Muller, in prep.). See Lose Mangroves . . . Lose Biodiversity, next page, for some impacts to biodiversity from loss of mangroves.

plantations: threat in kind . . . threat in name Logging, both legal and illegal, is often regarded as a primary agent for forest loss in Papua. However, conversion for agriculture, especially oil palm plantations, looms as a threat (see Some Biodiversity Impacts of Plantations, next page). Casson (2000, 2003) found that companies preferred to expand oil palm operations in Sumatra because it is best suited in terms of climate, soil, and infrastructure for

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LOSE MANGROVES . . . LOSE BIODIVERSITY Globally, about half of the world’s mangrove forests may have been lost. Lost: food and refugia for many species Lost: nutrients for the marine environment that support trophic structure Lost: nursery grounds for fish and shellfish species Lost: nesting sites and migratory sites for hundreds of bird species Lost: protection from siltation afforded to seagrass and coral ecosystems Source: Adapted from UNEP (2002).

SOME BIODIVERSITY IMPACTS OF PL ANTATIONS • 80–100 % of mammal, reptile and bird species are lost (when established on primary tropical forest). • Fire is often used as a management tool on plantations. This fire can spread to the surrounding habitats. • Palm oil production brings pollution. Carelessly and copiously applied pesticides, herbicides and fertilizers plus processing waste products endanger aquatic and terrestrial flora and fauna, and even coral reefs (from runoff). Source: Friends of the Earth (2004).

‘‘For most local animals, a plantation is a desert, lacking food, shelter and opportunities for reproduction’’ (Carrere and Lohmann 1996).

cultivating oil palm. While government policy provides for incentives to expand the plantation sector to eastern Indonesia, there is still high demand to open up more land for plantations in Sumatra. With the deficit of land available there and the aforementioned prediction of the effective loss of Sumatran lowland forests by 2005, it is likely that investors will establish large-scale oil palm plantations in East Kalimantan and Papua (Casson 2003). Furthermore, changes in government policy have paved the way for oil palm expansion into logging concession areas or conversion forest, which corresponds neatly to the situation in East Kalimantan and Papua, where large tracts of these forest designations exist. In addition to increasing the area of allocated conversion forest in Papua and Maluku, the government has increased the incentive for companies to establish new plantations in ‘‘nonproductive production forests’’ (Asia-Pacific People’s Environment Network 2002a). These areas are defined as logging concession forests containing less than 20 m3 of timber per hectare. Under this regulation, 60% of the nonproductive area must be converted to timber plantation, and the rest may be used for estate crop

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plantations (FWI/GFW 2002). In areas where profitability can be achieved this regulation provides a perverse incentive (and a major threat to sustainable forestry in Indonesia’s lowland rainforests) for the company to (illegally) reduce the timber density below the aforementioned threshold and then apply for oil palm and softwood plantation permits (Wakker 2000). Oil palm plantations throughout Indonesia have usually been established on conversion forest land rather than on the readily available degraded land in the same province. There is a reason for this. Oil palm companies can offset the cost of plantation establishment with the profits obtained from timber extraction, sometimes including valuable tropical hardwood species. This has been the case particularly in Kalimantan and Papua, where the total area of oil palm plantations had only reached half a million (517,328) hectares by 2000, despite the fact that companies had applied for permits to establish oil palm on more than 3.8 million hectares of forest land (Casson 2003, citing Badan Planologi 1999). Up to 1998, plantation development in Papua reached 128,183 ha consisting of local communityowned plantations (97,159 ha), state-owned plantations (31,024 ha) and privately owned plantations (18,270 ha). Meanwhile, land area licensed to be converted to plantation area was 1,263,742 ha. According to the Plantation Services of Papua, potential land for plantation development in Papua was 6,115,443 ha in 1999. There was one state-owned plantation company operating in Jayapura and Manokwari regencies, and nine private companies operating in Papua at this time (Anggraeni 2005). Wakker (2005) quotes news reports and other sources that about 2.8 million hectares in the districts (regencies) of Jayapura, Manokwari, Sorong, Merauke, Yapen, Waropen, Nabire, and Timika had been reserved for oil palm plantations in May, 2003. Van Gelder (2001) and Wakker (2000) list details of some of the companies and subsidiaries who have established oil palm plantations (or concessions to such) in Papua. Large areas of production forest have also been allocated to oil palm companies in Central Kalimantan, East Kalimantan, and Papua, despite the fact that conversion forest is still available in these three provinces. As with conversion forests, this loophole effectively allows companies to clear-cut production forests, if in particular cases this is deemed profitable. Notwithstanding financial, infrastructure, labor, or even social constraints that may have impeded establishment of some plantations, some companies are apparently only after the profitable timber cover. After all, long term investment in Papua and other places where there is not a long-established palm oil industry is more risky than walking away with the timber profits (Casson 2003). Traditional gardening is also threatening specific pockets of Papuan biodiversity. Increasing encroachment of gardens stimulated by urban drift is clearly visible from Sentani climbing the slopes of the Cyclops Mountains, a small insular mountain range and biodiversity locus on the north coast of Papua near Jayapura (pers. obs.).

mining and energy: scratching the surface Mining can be broken down into five stages, from exploration to closure and postoperation; each stage has its own set of activities and a long list of potential im-

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pacts on biodiversity (see Miranda et al. 2003). Mining has also contributed to the large-scale loss of biodiversity habitat in Papua (see Biodiversity Impacts of Mining, below). Apart from loss and degradation around a mine, the associated development of roads, towns, and ports, the resulting pollutants carried in water courses, and the increased competition for land and resources from an the influx of outsiders, also do damage (Asia-Pacific People’s Environment Network 2002a). Although the area of Papua currently under exploitation is only 1.3 million hectares (by one company, PT Freeport Indonesia), the total area approved for smallscale mining and contracted for commercial mining investigation, exploration, and exploitation, adds up to about 11 million hectares (Anggraeni 2005, citing JATAM 2000), which is over 25 percent of the provincial territory. This involves an additional 13 companies currently undertaking exploration only (Anggraeni 2005). More than 60 percent of the areas contracted are within protected and conservation forest (Conservation International Papua Program 2004a). Almost 25 percent of mining concessions are located within mountain forest area, and more than 65 percent of mining concessions are within lowland forest. Only 15 percent of the mining concessions are actually located within a distance of 10 km from main roads (Conservation International Papua Program 2004b). These mine concessions are situated mostly in the northern parts of Papua, with most areas concentrated in the central and eastern portions. There are also four licensed local small-scale gold mining operations operating along rivers in Jayapura and Nabire regencies and covering over 18,000 ha in concessions (Anggraeni 2005). Because most of the major mineral deposits that can be mined in Papua involve surface mining, it is inevitable that the original landscape will be destroyed, along with the biodiversity and ecosystems hosted there (Anggraeni 2005).

SOME BIODIVERSITY IMPACTS OF MINING Direct impacts • Habitat loss/fragmentation • Disturbance to wildlife communities • Chemical contamination of surface waters resulting in: • Toxicity impacts to organisms (terrestrial and aquatic plants and animals) • Persistent contaminants in surface waters resulting in: • Persistent toxicity to organisms • Loss of original vegetation/biodiversity • Declining species populations

Secondary impacts • Increased colonization due to (road) development • Species loss due to hunting Source: Adapted from Miranda et al. (2003).

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As previously mentioned, PT Freeport Indonesia is the only company currently commercially exploiting its mining concession. Having the world’s largest gold deposit and the second-largest copper deposit, all in one current ore body, the Grasberg mine is a major operation (which followed on from the original Ertsberg body). The licensed concession covers 12,997 km2, a reduction in area (from 26,150 km2) after mining permission was extended in 1997, now due to expire in 2021, with options for two 10-year extensions (Anggraeni 2005, citing Provincial Department of Mining and Energy 2000). The mine is expected to be economically viable until 2041 (van Zyl et al. 2002). Mill throughput from production from PT Freeport Indonesia’s producing mines averaged 219,500 metric tons of ore per day from 1998–2003 (Freeport-McMoRan Copper & Gold, Inc. 2003, 2004). In 1996/ 1997 it was reported that 125,000 tons of mine tailings were dumped into the Ajkwa River per day (International Federation of Chemical, Energy, Mine and General Workers’ Unions 1998). According to a 1999 audit 230,000 tons of ore were processed a day, generating 400,000 tons of waste rock and overburden per day (van Zyl et al. 2002, citing Montgomery Watson 1999). This displacement of waste rock has led to the effective destruction of several high-energy rivers that drain the Grasberg and Ertsberg environs. While the Grasberg mine itself was not situated in a forested area (Asia-Pacific People’s Environment Network 2002a), about 14,600 ha of land below the mine constitutes the ‘‘Modified Ajkwa Deposition Area’’ where tailings are transported by the Ajkwa River system (PT Freeport Indonesia 2004), resulting in the loss and degradation of lowland forest in this area (Asia-Pacific People’s Environment Network 2002a). While levees have been constructed to contain these tailings within this spillway (PT Freeport Indonesia 2004), previously tailings-laden water had spilled over into the neighboring Minajerwi River because of heavy sedimentation and flooding (Kennedy 1998). These mine tailings and other wastes constitute a major environmental problem, and a controversial one. Before extracting metal-bearing ores and producing tailings, all of the ‘‘overburden’’ above the deposit—the vegetation, soil, and rock—must be removed. Overburden exists in a ratio of about three to one with the ore at Grasberg, and the massive overburden is being dumped in two adjacent alpine valleys (Kennedy 1998). The mine waste contains sulfide-bearing minerals and is producing acid drainage (van Zyl et al. 2002). Acid mine drainage can be devastating to the ecology of a river and its associated environment, by raising acidity and releasing poisonous heavy metals (e.g., mercury, lead, and cadmium) into the river system (International Federation of Chemical, Energy, Mine and General Workers’ Unions 1998) once the neutralization potential of surrounding limestone is used up (Kennedy 1998). A sacred alpine lake, Wanagon, is the recipient of 30,000 tons of overburden a day. It can no longer be used for drinking water by local people (Dwiyana 2001). The dam at Lake Wanagon has failed three times: June 1998, March 2000, and May 2000 (Johansen 2002) and there was an overflow in 1999 (Asia-Pacific People’s Environment Network 2003). Mine tailings are dumped continuously into the Aghawaghon River, which

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merges into the Otomona and Ajkwa rivers (Kennedy 1998), flowing down steep mountainsides through rainforest at lower elevations, producing a desolate landscape (Johansen 2002, citing Roberts 1996)—a vast and awesome scene observable from the air on approach to Timika airport. The cause of this devastation is in dispute. The company states that the tailings are alkaline (i.e., not acidic), that samples from the Ajkwa River meet U.S. and Indonesian quality standards for dissolved metals in drinking water, that the estuaries downstream of the tailings deposition area are ‘‘functioning ecosystems,’’ and that native species are easily reestablished in the highlands and that native species as well as agricultural crops grow well in the lowlands on soil containing tailings (Freeport-McMoRan Copper & Gold, Inc 2004; PT Freeport Indonesia 2004). In terms of the habitat destruction seen in the deposition area, company literature focuses on the physical, (i.e., not chemical) effect of heavy sedimentation (tailings). Altered hydrology and adjacent vegetation is characterized as a temporary situation that will return to equilibrium once the mine ceases operation, including implementation of the reclamation phase (Freeport-McMoRan Copper & Gold, Inc. 2002; PT Freeport Indonesia 2004). This is at odds with several other, sometimes cross-quoting sources (e.g., Asia-Pacific People’s Environment Network 2001; Bryce 1996; Dwiyana 2001; International Federation of Chemical, Energy, Mine and General Workers’ Unions 1998; Johansen 2002; Kennedy 1998; van Zyl et al. 2002.). These sources include or cite data and reports that indicate that (in addition to the detrimental effects of heavy siltation) there is significant chemical involvement: that the waters exposed to the tailings are seriously polluted and that the tailings are a major source of toxicity. Interestingly, ‘‘few mines around the world currently utilize riverine tailings disposal for waste management, and all [that do] are located on the island of New Guinea’’ (Miranda et al. 2003).

gas and oil Papua’s first major crude oil deposits were discovered around Sorong, Salawati (Raja Ampat Islands), and Bintuni Bay during the Dutch colonial period preceding World War II. Exploration continued and the first productive wells were developed in Klamono, Salawati, and Wariagar (over an area of 150 km2). These wells have been exploited by the state oil company Pertamina since 1964. Other oil fields were developed at Sele and Linda. All of these fields are concentrated in the Vogelkop region of Papua (Dwiyana 2001). In 1998 a new major natural gas field ‘‘Tangguh’’ was located at WiriagarBintuni. Exploratory drilling indicated that this gas deposit could last 15 to 20 years (Anggraeni 2005). Development of a project commenced, and after a succession of negotiations and corporate mergers, British Petroleum (BP) and Pertamina are now developing the Tangguh LNG (liquified natural gas) project. Construction of the plant was to commence in late 2002 and production is expected to begin in the last quarter of 2005 (BP 2002; Chapters 6.4 and 6.5). The Tangguh project involves offshore production platforms and undersea gas pipelines in Berau-Bintuni Bay, connected to land facilities located on the bay’s

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southern shore. The onshore project site is located between the Saengga and Manggosa rivers and will take up about 3,200 hectares (BP 2002). Located 80 km east of the project is southeast Asia’s most extensive intact mangrove area in the Bintuni Bay Strict Nature Reserve (Tangguh Independent Advisory Panel 2002). Depending on sales, the facilities might require expansion, but this would not require additional land (BP 2005). As such, the project site will be relatively small compared to other resource extraction projects, like the Grasberg mining complex, or large logging concession operations (International Crisis Group 2002). While this will minimize direct impacts on biodiversity, there are a host of potential secondary impacts of the project that could be just as detrimental. Secondary impacts are those that are usually triggered by the operations, but may reach outside project or even concession boundaries and may begin before or extend beyond a project’s life cycle, whereas primary impacts result directly (and usually immediately) from project activities. Secondary impacts tend to result from government decisions and the actions and practices of nearby communities or immigrants, in response to the presence of the project, rather than from the operational decisions and the activities of project personnel (Energy and Biodiversity Initiative 2004). There is existing industrial infrastructure surrounding the Tangguh project site, including substantial fishing and shrimp fishing in the Berau-Bintuni Bay. Extensive land has been cleared for oil palm, agriculture, and transmigration settlements in the area directly south of Tangguh. The Tangguh site has poor soils and limited accessible water supplies, making the development of a new town with related industries unsustainable. Furthermore, no roads to the site currently exist and none are planned. The highly automated facility will require a final workforce of just 500 people (BP 2002), but up to 5,000 workers will be employed in the Bintuni Bay area during the construction phase, and this will be the most ecologically sensitive period in the life of the project (Tangguh Independent Advisory Panel 2002). In an effort to alleviate the social conflicts associated with the aforementioned extractive industries, the project will attempt to control these secondary impacts with the added benefit of protecting the area’s biodiversity. The project’s solution is to restrict the workers on the site and to use Sorong, Fakfak, and Manokwari as leave sites (to which workers return when they are off duty, and where they are paid) and as centers for supplies, administration, and other ancillary aspects of the project. Workers will be obliged to leave Bintuni Bay once the work is over. In this way BP seeks to spread the positive and negative impacts thinly over a wide area rather than allowing them to pile up around the Tangguh project site (International Crisis Group 2002). A similar stringent effort to control secondary (exogenous) impacts from oil and gas production in a sensitive rainforest area along the Kikori River in Papua New Guinea, was observed to be successful (Diamond 1999). Still the threat of direct impacts exists. A potential significant impact is the handling of drilling fluids and cuttings. The project will seek to re-inject all drilling fluids and cuttings into underground formations, with a view to having a zero

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discharge operation. Only in an emergency situation would drilling fluids and or cuttings be discharged into Berau-Bintuni Bay, in strict compliance with Indonesian and World Bank environmental standards. The natural gas in the Tangguh fields has a CO2 content of approximately 10%, which is relatively high. The Tangguh project is looking at options for reducing greenhouse gas emissions (BP 2005). All of the liquified natural gas will be exported, leaving no residual product for local use (BP 2002), lessening the potential for threats from associated industries. See Factors That May Lead to Secondary Impacts on Biodiversity, below.

infrastructure development: dreams versus destruction As alluded to previously, large-scale infrastructure development also constitutes a serious threat to biodiversity in Papua, not only from direct habitat loss and fragmentation, but also from a host of accompanying secondary impacts. Road density (ratio of length of road to size of area) in Papua is one of the lowest in Indonesia, at 0.04 km/km2, far below the national average of 0.17 km/ km2 (Anggraeni 2005; Chapters 6.4 and 6.5). However, after national decentralization, transportation was listed as a separate budget item, accounting for 21.51% of the total Papua provincial development budget in 2002, the largest of all such

FACTORS THAT MAY LEAD TO SECONDARY IMPACTS ON BIODIVERSITY Immigration and new settlements are often stimulated by a large energy project (such as a mine or other large development). In addition to attracting people looking for work, an oil and gas operation can also provide entry into an area that was previously inaccessible for other purposes. Access is usually facilitated by the construction or improvement of linear infrastructure, such as roads and pipelines, into these environments. Some potential secondary impacts on biodiversity associated with these factors include: • Deforestation from clearing of land for agriculture, building housing and other infrastructure, and collection of wood for construction, cooking, and heating • Increased demand on water resources and generation of wastes and other pollution • Increased demand for public services such as schools, law enforcement, and health care, that reduces the resources available to address biodiversity concerns • Commercial and illegal logging • Extraction of non-timber forest products, such as fibers, medicinal plants, and wild foods • Increased hunting and fishing, for subsistence or trade in bushmeat • Poaching for skins, exotic pet trade, or other uses, such as folk remedies. Source: The Energy and Biodiversity Initiative (2004).

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budget items. Much of this earmarked amount goes toward linking strategic road segments in Papua. According to official plans, major roads will extend 4,016 km in Papua, of which 2,303 km have already been paved (Anggraeni, 2005). According to a spatial analysis conducted under Conservation International’s Rapid Assessment for Conservation and Economy (RACE) project in Papua in 2002 (refer to Mertens 2002), if the planned regional-scale roads were added to existing regional-scale roads, the total affected area of land would increase dramatically (Anggraeni, 2005). In order to estimate the extent and influence of the planned road network, theoretical ‘‘zones of influence’’ of various widths were mapped, paralleling the existing and planned road infrastructure. These zones were delineated with a 10 km, 20 km, or 50 km width from both sides of the road network. The analysis showed that approximately 15, 22, and 51 percent of the area of the province is located within 10, 20, and 50 km, respectively, of the planned and existing regional-scale roads (Conservation International Papua Program 2004b). Almost 25 percent of conservation and protection forests are located within less than 20 km of existing regional roads. Considering the area affected by planned regional roads with a 20 km zone of influence, 35 percent of protection forest and almost 30 percent of conservation areas would be affected (CIPP 2004b). The amount of mountain forests affected would double if the EnarotaliWamena segment in the eastern part of Paniai were to be constructed, because most of these areas are located within 10 or 20 km from proposed roads (Anggraeni 2005). In recent years, two other major infrastructure developments seriously threatened Papuan biodiversity. The first was a proposed ‘‘road for logs’’ deal in which the Indonesian central government would grant a foreign consortium a contract to construct the trans-Papua highway for the length of 11,280 km. In return, the developers would be granted rights to all of the logs within five kilometers on both sides of the highway and they would be given the rights to manage plantation and other forestry projects in the surrounding areas. This was supposed to commence in 2001 but the project has apparently not materialized. The other case involved the proposed ‘‘Mamberamo mega-project’’ which involved constructing a series of dams in the 8 million hectare and seismically active catchment in order to generate 12,000 mW of electricity mainly for mining and smelting. In addition, major agro-industries, logging, and major downstream developments were planned (Anggraeni, 2005). These plans were ‘‘shelved’’ when investors lost confidence in the viability of the project. While these plans have been put on hold, they have not been forgotten, and represent potential catastrophic threats to the biodiversity of Papua. Each would result in massive, direct forest loss (logging, conversion, or flooding) and habitat fragmentation, and each would bring the manifold secondary detrimental impacts that attend such huge projects.

threats to the marine environment: awash in a sea of troubles Paralleling the threats to its terrestrial natural wilderness endowment are the threats confronting Papua’s marine heritage. Papua contains some of the world’s

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most biodiverse coral reef faunas, with its Raja Ampat Islands sitting as they do at the heart of the Coral Triangle, the world’s center of marine biodiversity. And this natural bounty is also under assault. Major causes of coral reef decline are tourism overuse, destructive fishing practices, runoff and land-based pollution, climate change, and, in particular, the associated effect of coral bleaching. The most alarming threats involve destructive and wasteful fishing practices, especially using homemade bombs to harvest fish. Some of the world’s most spectacular and productive coral reefs have been effectively destroyed for the medium term by blasting, all for the short term gain of filling a boat with fish—once. During Conservation International’s Marine Rapid Assessment Program (RAP) survey work in Raja Ampat in 2001, teams noted that overall reefs were in good condition but over 13% of reefs surveyed had experienced blast damage (McKenna, Allen, and Suryadi 2002). Similarly, the fairly widespread practice of using potassium cyanide and other piscicides to capture particular marine species (e.g., large carnivorous fishes like groupers, serranids, Napoleon wrasse, and lobsters) destroys coral reefs and kills other fauna. Complaints were heard in villages throughout the survey area about blasting and poisoning fish ‘‘by outsiders’’ which was said to have increased significantly in the preceding two years. Overharvesting of fish (for example Napoleon wrasse for the live fish food trade) and invertebrates is also another threat to marine biodiversity which was identified in the 2001 Marine RAP (McKenna, Allen, and Suryadi 2002). A place like Raja Ampat is particularly vulnerable to these kinds of threats, being a porous archipelago of hundreds of islands, approachable from all directions. Another direct threat that devastates these rich habitats is the practice of coral reef excavation for building materials. The Nature Conservancy (TNC) identified commercial harvest as a major threat to turtle populations in Raja Ampat (noting 68 carapaces found in one small village), and mentioned the trade in shark fin and the dearth of shark observations (Donnelly, Neville, and Mous 2003). Threats also come from the terrestrial realm where agricultural runoff of fertilizers, pesticides, and herbicides can have deleterious effects on marine life. Logging often results in erosion which can lead to siltation, smothering nursery areas like mangroves and seagrass beds, and productive coral reefs (Chapters 5.2–5.4).

invasive species: our uninvited guests One of the most pernicious threats confronting biodiversity on the planet is invasive species. Whether intentionally carried by humans as pets or as stock (e.g., fish) and released, or accidentally introduced, as stowaways on ships, planes, or heavy equipment, these invaders are slowly wreaking havoc in ecosystems across the globe, by mixing of faunas and floras across heretofore minimally permeable biogeographical boundaries. Global trade and communication is increasingly an agent in this spread. Alien species suddenly dropped into new environments may fail to survive, or they may thrive and become invasive in situations where they outcompete the native species (IUCN/SSC 2005).

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Invasive species successfully establish themselves in, and then overcome, otherwise intact, pre-existing native ecosystems. Factors which may facilitate invasion include: a species finds itself suddenly without the normal pressures of predators or parasites of its native habitat; a species has habits or cycles that enable it to invade (e.g., a generalist diet or short generation time); a species fills a niche that was previously not occupied; or a species is introduced into an ecosystem already disturbed by humans or some other factor, which enhances its ability to invade. But whatever the causes, the consequences of such invasions—including alteration of habitat and disruption of natural ecosystem processes—are often catastrophic for native species (IUCN/SSC 2005). Furthermore, small islands, while not more susceptible to invasions by alien species than larger landmasses, are particularly vulnerable to the impacts of such invasions. Compared with species found on the mainland, the space-constrained island species are generally less capable of moving elsewhere, are comprised of fewer populations, and have smaller total population sizes. These characteristics, coupled with isolation and endemism, make island ecosystems especially sensitive to disturbance, and island species typically more vulnerable to extinction, than those of continental systems. Invasive alien species are perhaps the most significant driver of the decline of plant and animal populations and species extinctions in island ecosystems (Subsidiary Body on Scientific, Technical and Technological Advice 2003). Species invasions, together with habitat destruction, have been a major cause of extinction of native species throughout the world in the past few hundred years. While the extent to which such losses have gone unrecorded in the past is unknown, today there is an increasing realization that biological invasions play a major role in terms of irretrievable loss of native biodiversity (IUCN/SSC 2005). Papua too has been invaded. A search of the Global Invasive Species Database (ISSG 2005) reveals 18 records of invasive species (14 alien and four native species) in Papua (see Invasive Species in Papua, next page). An analogous search for Papua New Guinea revealed 46 invasive species. Depending on their position within PNG, it would seem that Papua might be threatened additionally by some of these (and vice versa as in the case of the alien macaques in eastern Papua).

Aquatic Invasives in Papua: Foreign Fish In relation to its overall size, the New Guinea region exhibits a remarkably low incidence of invasive freshwater species, but in 1991 the presence of 22 invasive species representing 19 genera, 11 families, and six continents had already been recorded in New Guinea (citing Allen 1991). Since then at least six more introductions have been noted, and more can be expected, especially in Papua (Polhemus, Englund, and Allen 2004; Chapter 5.5). In Papua, the Mamberamo River system is ‘‘badly contaminated’’ with invasives. Carp inhabit the upper Baliem River, Tilapia are in the Timika area, predatory Snakeheads (Channa striata) are currently found on Waigeo Island in the Raja Ampat Islands, in streams near Bintuni, and in the Timika region. Walking Catfish (Clarias batrachus) are now in parts of the Timika region, after first having

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INVASIVE SPECIES IN PAPUA REGISTERED IN THE GLOBAL INVASIVE SPECIES DATABASE Alien 1. Achatina fulica (mollusk/snail) 2. Anoplolepis gracilipes (insect/ant) 3. Chromolaena odorata (herb) 4. Clarias batrachus (fish) 5. Eichhornia crassipes (aquatic plant) 6. Lantana camara (shrub) 7. Leucaena leucocephala (tree) 8. Limnocharis flava (aquatic plant) 9. Macaca fascicularis (mammal) 10. Mimosa pigra (shrub)

11. Piper aduncum (shrub, tree) 12. Pomacea canaliculata (mollusk/ snail) 13. Rattus exulans (mammal/rat) 14. Solenopsis geminata (insect/ant)

Native Species 1. Adenanthera pavonina (tree) 2. Boiga irregularis (reptile/snake) 3. Cupaniopsis anacardioides (tree) 4. Platydemus manokwari (flatworm)

Source: ISSG (2005).

been introduced into New Guinea in the Lake Sentani region, and are now also found in the Vogelkop. Timika also now hosts the exotic Blue Panchax (Aplocheilus panchax). Climbing perch (or climbing gouramies) are now common across central southern New Guinea having been first introduced to Papua. Two families and three species of gourami are found in New Guinea (Polhemus, Englund, and Allen 2004, citing Allen and other authors). Many of the invasive fish species now present in New Guinea have been documented to cause extinction or severe range reductions of native fish and invertebrate taxa in other areas of the world, but their specific impacts within New Guinea ecosystems are at present conjectural (Polhemus, Englund, and Allen 2004). The fact that no evidence of invasive invertebrate species was encountered during recent faunal surveys in New Guinea led the authors to conclude that most if not all of the current freshwater invasives on the island of New Guinea are fishes.

Macaques in Papua While there are many theories, it is presently not known how or when (or even perhaps, why) Macaca fascicularis was introduced in Papua (Kemp and Burnett 2003). Even if these facts were known they would be of limited value in dealing with the invasive species problem (and this particular species can symbolize as a kind of perverse ‘‘flagship invasive’’ for Papua). In Chapter 7.7, Kemp and Burnett discuss this issue in detail. The essential point is how easy it is for a destructive alien species introduction to take place in Papua. The prevailing perception seems to be that all local species are ‘‘Indonesian’’ rather than from particular islands (e.g., from Java, from Sulawesi, from Papua), and awareness of basic biogeography or concepts of endemic or alien species is almost nonexistent. Certainly there is

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very little local consciousness among local residents, military, or policy makers of the possible negative consequences that released or escaped exotic pets might have on local ecosystems (Kemp and Burnett 2003).

trade in species Another all-too-common threat to the biodiversity of Papua comes from the trade in species of flora and fauna. Across Papua in towns and villages one can see a plethora of caged birds (lories, parrots, birds of paradise, cockatoos, cassowaries, crowned pigeons) as pets or on sale. Marsupials and pythons are also frequently on display. Crocodile skulls and sea turtle shells can be bought. Orchid stands line the province’s highways (pers. obs.). Persistent anecdotal information indicates substantial trade in species by soldiers departing Papua at the end of their tours of duty. A study of illegal wildlife trade was carried out by Conservation International and the government conservation agency, in Manokwari and Jayapura (20 November 2002–10 January 2003). The study targeted markets, seaports, airports, arts and crafts shops, and nearby forest areas thought to be the source of some fauna in trade (Suryadi, Wijayanto, and Wahyudi 2004). The study, although brief, documented an extensive and thriving trade in animals, plants, and their products from Papua.

social and policy threats Papua is in the midst of social and policy flux brought on by the strange brew of Special Autonomy status (bestowed on it by Law No. 21/2001 but never fully implemented) and the de facto division of Papua province into two provinces (not three, as was intended by resurrection of the Law No. 45/1999 by legislatureendorsed Presidential Decree INPRES 1/2003) resulting from a court decision overturning the three-way division, but letting the new province of West Irian Jaya stand, because it was a fait accompli (Chapter 6.1). This confusing state of affairs is built on the foundation of contradictory laws and interpretations, and competing jurisdictions (i.e., province versus regency) (YPMD 2004) and sometimes fickle and fluid constituencies. The indecision and ambiguity this situation engenders makes Papua ripe for unbridled and unsustainable resource exploitation and profiteering, all at the expense of the public and of the biodiversity of Papua.

Redistricting Fever Perhaps a uniquely Papuan threat, or maybe more precisely, a threat emblematic of Papua, is the current wave of redistricting fever washing across the land. While the splitting of the province and its parallel splitting and spawning of new regencies (kabupaten) and districts has its genesis in Indonesian central government plans and decisions, it has also created aspirations for new regencies (kabupaten) among local communities. Under the Regional Autonomy Law No. 22/1999 numerous provinces and regencies have been created, often as a result of successful lobbying by those regions concerned. The creation of new administrative units

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under the General Autonomy Law is most often driven by indigenous groups. The interest in becoming a new political jurisdiction in places like Fakfak, Merauke, Serui, Biak, Nabire, and the highlands is conditional—so long as they become the new administrative center (Chauvel and Bhakti 2004). In addition to the four regencies (kabupaten) created via Law No. 45/1999 (and its successors), another 14 of these jurisdictions were born by the enactment of Law No. 26/2002. By statute, a potential regency must meet a number of criteria in order to become an official regency (kabupaten). One of these is attainment of a threshold human population. Nevertheless, there are machinations going on across Papua to cleave off yet more regencies (kabupaten) even in areas with sparse populations that drastically fall short of the threshold population but have high timber density. There is also a notion that regency (kabupaten) status will bring a windfall of central funding even though a pie does not increase in size the more it is sliced. Perhaps more worrisome than this apparent naivete´ is the lure of rich natural resources, especially forests. The access to resources and position that comes with partition is an attractive force (Chauvel and Bhakti 2004). While a new regency (kabupaten) does receive initial outlays of central government funding, it must eventually assume a greater share of the cost of operations itself, as central funds are decreased. In order to fund these administrative operations, and most notably development of infrastructure, a new regency (kabupaten) often has nowhere else, or perhaps no easier place to turn, than to its forests. Even a regency (kabupaten) that does meet minimum startup criteria faces significant challenges. Detrimental effects can be compounded, for example, when the new administration does not have the capacity to manage the exploitation it has embarked upon. This only exacerbates the unsustainability of this course of action, and is conducive to uncontrolled and illegal logging, and so on. The unbridled expansion of new jurisdictions in Papua is a major potential threat to biodiversity. Another social threat to the biodiversity of Papua stems from government and other entities failing to properly ‘‘socialize’’ plans and activities among traditional communities. For the Papuan people, the advent of Special Autonomy was a nonevent if one considers a poll conducted in 2002. According to the results, an average of ‘‘83% of all Papuans have never heard of something called Special Autonomy Law’’ (International Foundation for Election Systems 2003). However, the Special Autonomy Law reinforces the role of traditional (adat) law (International Crisis Group 2002), and nowadays the expression of pride in and the rights of traditional (adat) communities is the rule. Traditional communities are asserting their traditional rights over the forests where they have lived for generations. Often coupled with this expression of rights (hak) is a skepticism or a distrust of outsiders and institutions, built up over years of not being consulted or being cheated. Now more than ever it is imperative that the government, nongovernmental organizations, and others treat traditional communities on an equal footing or a footing commensurate with their status as traditional owners of the forests. That is if the desire is to maintain the heritage of Papuan biodiversity. Trust is slowly built and hard won, but it can be lost in an instant. Traditional

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communities must be full partners in this endeavor to safeguard Papuan biodiversity.

natural threats The world is dynamic, bringing forth seasons and cycles, and movements of earth, wind, and fire. These interactions support biodiversity and, at times, threaten it. This has been the case since time immemorial and life has thrived and diversified into its myriad forms. So while natural threats do threaten biodiversity, it is the greater threat of exacerbation from anthropogenic interactions that is most worrisome, a phenomenon the Worldwatch Institute has termed ‘‘unnatural disasters’’ (see Abramovitz 2001). These are those disasters that are made more frequent or more severe due to human actions. And these altered threats from nature are the greater threat to biodiversity. Many ecosystems and species are adapted to natural disturbance, and indeed disturbances are necessary to maintain their health and vitality, and even the ecosystem’s continued existence (Abramovitz 2001). It is when these natural perturbations are compounded by the actions and changes wrought by humans, that the natural becomes unnatural. Typical natural threats include volcanic eruptions, earthquakes, and tsunamis, as well as cyclones, extreme and cyclical weather phenomena, periods of drought, and lightning-induced fire. These events can precipitate various additional secondary effects, such as landslides, flooding, drought, and habitat destruction. Miranda et al. (2003) pointed out that mines are particularly vulnerable to landslides and floods, especially in seismically active areas. And should they topple or wash away tailings or waste overburden, then the impacts on biodiversity are compounded. Papua, given its position along tectonically active plate boundaries, is seismically active, experiencing frequent, occasionally major, earthquakes (e.g., the Nabire earthquakes of 2004). Likewise undersea earthquakes have spawned tsunamis in Papua on occasion (e.g., in Biak in 1996, and at nearby Aitape, PNG, in 1998). Interestingly enough, Papua does not have any active volcanoes but is bounded on both east and west by many. The most significant natural disaster for biodiversity in Papua occurred during the 1997–1998 El Nin˜o event. Prolonged drought affected areas in the central highlands, and exacerbated fires which consumed 1,000,000 ha of forest in Papua (Boissie`re 2002). Some 6,000 ha inside Lorentz National Park were burned because of El Nin˜o conditions (UNEP-WCMC 2001).

global climate change Of all of the threats examined thus far, none have a scale, scope, or reach that matches the escalating crisis in global climate change. This reach extends to New Guinea and Papua, and what happens in New Guinea or Papua contributes appreciably to the global effect. And while climate is a natural phenomenon which has oscillated greatly over time, there is compelling evidence that the current episode of climate change is not natural.

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Globally, 1998 has proven to be the warmest year on record, with 2002 and 2003 coming in second and third, respectively. There is a recognition that the earth is now out of energy balance, with 0.85  0.15 W/m2 more solar energy coming in than terrestrial heat radiation going out to space (Hansen et al. 2005). One result of this imbalance is that it makes it likely that global temperatures in 2005, aided also by a weak El Nin˜o, will exceed those of 2003 and 2004 and perhaps even the temperature of 1998, which had stood out far above the temperature of any year in the preceding century (Goddard Institute for Space Studies 2005). The amount of CO2 in the atmosphere in 2003 reached an all time high of 376 ppm (World Wildlife Fund 2005). Many species of flora and fauna will probably not be able to survive climate change. Recent modeling suggests that 15–37% of a sample of 1,103 terrestrial species of plants and animals would eventually succumb to climate changes by 2050 and are ‘‘committed to extinction.’’ Some species will simply not find suitable habitat available and others will be unable to reach places where the climate is suitable (C.D. Thomas et al. 2004). Climate change exerts a perverse synergy, interacting with other threats to make them worse. Climate change may affect species directly, for example through changes in temperature and precipitation. But often the indirect effects are even more important. Secondary impacts may be increased pressure from competitors, predators, parasites, diseases, and disturbances (e.g., fires or storms). Climate change will often act in combination with major threats such as habitat loss and alien invasive species, making their impacts considerably worse (BirdLife International 2004). But the effects of climate change are not distributed evenly geographically and altitudinally. How will climate change adversely affect biodiversity in Papua? A World Wildlife Fund study of climate change in globally significant terrestrial ecosystems provides the following generalizations that might be applicable to Papua as a tropical region. Changes in habitats from global warming will be more severe at high latitudes and altitudes than in lowland tropical areas. In Southeast Asia, insular areas tended to show stability, whereas mainland areas tended to be more vulnerable. In non-glaciated regions, where previous selection for high mobility has not occurred, species may suffer disproportionately. Therefore, even though obligately high migration rates are not as common in the tropics as in colder regions, they may still have strong impacts in terms of species loss. Finer subdivision of biomes led to much higher migration rates in warm temperate and subtropical areas; only the lowland tropics still have relatively low migration rates. Generally lower levels of habitat change and required migration rates in tropical compared to temperate regions may not be particularly relevant if tropical species have lower migration capabilities, for example. Of course, the enormously high species richness in tropical regions makes these ecosystems of special concern regardless. In tropical regions, losses of just a few percentages of the biota potentially translate into tens of thousands of species or more (Malcolm et al. 2002). In PNG’s country report to the United Nations Framework Convention on

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Climate Change (UNFCCC), Kaluwin, Ashton, and Saluei (2002) reported that the country’s coastline, coastal villages, and rural coastal population would be vulnerable to sea level rise and other weather-related manifestations of climate change. Specifically, the report added: the main impacts will be inundation of coastal wetlands and foreshore areas, bleaching of corals, which will weaken the coral reefs as barrier protection systems. Loss of wetlands, freshwater sources due to seawater intrusion, and lands may eventually lead to displacement of communities, resulting in aggrivated [sic] future social problems. The permanent or periodic inundation of deltaic flood plains, swamps, and low-lying areas could affect up to 50% of the Papuan Coastlines, and 10% of the northern shorelines (for a 1 m sea level rise—IPCC’s highest estimate). This may result in damage to mangroves and swamp forest ecosystems, as well as human productive systems. More than 90% of the coastlines of Gulf and Western Provinces are likely to be impacted. Flooding is also expected to affect the lower Sepik-Ramu region. Approximately 4500 kilometers out of a total of 17100 kilometers of shoreline are expected to be moderately to severely inundated, affecting up to 30% of Papua New Guinea’s population. In addition, there is a danger that some very low-lying islands, including barrier islands, will be completely submerged. Evidences of this are already occurring, especially in the outer lying atoll islands of Mortlock, Tasman and the Duke of York Islands (Kaluwin, Ashton, and Saluei 2002: 41).

The report went on to mention the vulnerability of PNG’s substantial coral reefs. Given its adjacent location and that PNG’s aforementioned geographical features find many counterparts in Papua, climate change impacts manifested in PNG could also be expected to occur in Papua in regions endowed with similar topography. Such significant changes would radically impact the biodiversity of Papua. As climate change is coming from without Papua, what can be done to ameliorate these effects? Papua, with one-half of the world’s third-largest contiguous tracts of tropical forest, is an enormous global store of carbon. Release of that carbon through massive forest loss contributes significantly to global climate change. Alternatively, wise stewardship of those forests not only contributes to the maintenance of global climate stability, but also contributes to the maintenance of globally significant biodiversity. In Papua, one of the most telling signs that climate change is upon us is the steady retreat of the Carstensz/Mt Jaya glaciers, one of three equatorial glacier complexes on the planet (UNEP-WCMC 2001).

Ameliorating the Threats Threats to the biodiversity of Papua are manifold. Many of these threats appear to come from afar or are of such scale and magnitude that trying to do anything

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about them seems an exercise in futility (e.g., climate change). However, given the negative synergy that accompanies biodiversity threats, it is essential that efforts to ameliorate the most serious of them are undertaken rapidly. The influence of Papua’s forests on climate is of global significance, so that what happens in them and to them does have a direct (albeit ‘‘boomerang’’) effect on Papua’s climate and its biodiversity. And many threats to biodiversity occur closer to home. Of course, the broad characterization of particular activities as threats to Papuan biodiversity is made to highlight general issues and is not intended as a call for a total moratorium on these activities. But it is intended to draw attention to these issues, in the hope that they will be examined more closely and in a more holistic fashion. It is inevitable that there are some ecological sacrifices to developmental processes, but it must be determined if this sacrifice of biological diversity is actually benefiting overall economic return (see Thompson, 1996; Chapter 6.5). It is the daunting but imperative task of Papua’s planners, decision makers, the business community, conservationists, and communities to steer a course that provides for social and economic welfare but does not undermine it by destroying its genetic reservoir, its biodiversity base, and its ecosystem service infrastructure. Inevitably robust prioritization will be necessary, but should not be driven by short term economic considerations, as has typically been the case in the past. In many instances, remedial or preventive measures will necessarily have to be of landscape scale to truly prove effective. Initiatives such as the recently launched Forum for Conservation and Development in Papua (FKPTP: Forum untuk Konservasi dan Pembangunan di Tanah Papua) provide a structure and mechanism to bring together diverse stakeholders in order to collectively affect meaningful change. Comprehension and acceptance of the importance of Papua’s biodiversity by the broadest possible constituency is the crucial first step in ameliorating the threats to Papuan biodiversity. The success of such efforts ultimately depends on their sustained longevity and effectiveness, however these efforts must yield tangible progress in the short term, given the immediacy of the biodiversity crisis in Papua.

Acknowledgments Dessy Anggraeni routinely provided valuable insights and information, both directly and indirectly assisting me with this chapter. Special thanks to some superb colleagues in the Conservation International Papua office who assisted me in many aspects of this work.

Literature Cited Abramovitz, J. 2001. Unnatural disasters. Worldwatch paper 158. October 2001. Worldwatch Institute. Anggraeni, D. (2005). Rapid Assessment for Conservation and Economy (RACE) in Papua. Conservation International.

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Threats to Biodiversity / 1225 Asia-Pacific People’s Environment Network. 2000. Communities confront loggers. Down to Earth Newsletter No. 45, May 2000; and Contracts and communities. Down to Earth Newsletter No. 45, May 2000. Asia-Pacific People’s Environment Network. 2001. Court orders Freeport to clean up its act. Down to Earth Newsletter No. 51, November 2001. Asia-Pacific People’s Environment Network. 2002a. Forests, people and rights. Down to Earth Special Report: June 2002. Asia-Pacific People’s Environment Network. 2002b. The logging of West. Papua. Down to Earth Newsletter No. 55, November 2002. Asia-Pacific People’s Environment Network. 2003. Protests over fatal collapse at Freeport/ Rio Tinto West Papua mine. Down to Earth Newsletter No. 59, November 2003. BirdLife International. 2004. State of the World’s Birds 2004: Indicators for Our Changing World. BirdLife International, Cambridge. BirdLife International. 2005. Datazone. www.birdlife.org/datazone/species/terms/ threats.html. Accessed 15 February 2005. Boissie`re, M. 2002. The impact of drought and humanitarian aid on a Yali village in West Papua, Indonesia. Asia Pacific Viewpoint 43 (3), December 2002. British Petroleum. 2002. BP Indonesia Biodiversity Action Plan. British Petroleum. 2005. www.bp.com/sectiongenericarticle.do?categoryId2011026& contentId2016211. www.bp.com/sectiongenericarticle.do?categoryId2011028& contentId2016235. Accessed 26 February 2005. Bryant, D., D. Nielsen, and L. Tangley. 1997. The last frontier forests: ecosystems and economies on the edge. WRI/WCMC/WWF. Bryce, R. 1996. Spinning gold. Mother Jones September/October 1996. Casson, A. 2000. The hesitant boom: Indonesia’s oil palm sub-sector in an era of economic crisis and political change. Occasional Paper 29. CIFOR, Bogor. Casson, A. 2003. Oil palm, soy bean and critical habitat loss. Review prepared for the WWF Forest Conversion Initiative. World Wildlife Fund, Switzerland. Chapin, F.S., E.S. Zalaleta, V.T. Eviner, R.L. Naylor, P.M. Vitousek, H.L. Reynolds, D.U. Hooper, S. Lavorel, O.E. Sala, S.E. Hobbie, M.C. Mack, and S. Dı´az. 2000. Consequences of changing biodiversity. Nature 405: 234–242. Chauvel, R., and I.N. Bhakti. 2004. The Papua Conflict: Jakarta’s Perceptions and Policies. East-West Center, Washington, D.C. Christie M., J. Warren, N. Hanley, K. Murphy, R. Wright, T. Hyde, and N. Lyons. 2004. Developing measures for valuing changes in biodiversity: final report to Department for Environment, Food and Rural Affairs (DEFRA). London. Conservation International Indonesia-Papua Program. 2004a. Rapid Assessment for Conservation and Economy (RACE) in Papua Province. Factsheet. Conservation International Indonesia-Papua Program. 2004b. Spatial implications of roads and other types of development for conservation in Papua. Factsheet. Crutzen, P.J. 2002. Geology of mankind. Nature 415: 23. Departemen Kehutanan. 2003. Siaran Pers._No. 51/Ii/Pik-1/2003. Press release on controlling illegal logging. 15 January 2003. Diamond, J. 1999. Paradise and oil: oil exploration in the jungle of New Guinea. Discover Magazine March 1, 1999. Donnelly, R., D. Neville, and P.J. Mous. 2003. Report on a rapid ecological assessment of the Raja Ampat Islands, Papua, Eastern Indonesia, held October 30–November 22, 2002. Final Draft. November 2003. The Nature Conservancy. Dwiyana, A. 2001. Oil, gas and mining development and decision making process in

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1226 / scott frazier Papua. Paper prepared in the context of Rapid Assessment of Conservation and Economy (RACE). Conservation International Indonesia, Jayapura. Energy and Biodiversity Initiative. 2004. Negative secondary impacts from oil and gas development. Discussion paper. www.theebi.org/pdfs/impacts.pdf. Accessed 26 February 2005. Environmental Investigation Agency and Telapak. 2005. The last frontier: illegal logging in Papua and China’s massive timber theft. Environmental Investigation Agency, London. Forest Watch Indonesia. 2004. Landsat7 ETM Satellite’s Land Use Land Cover (LULC) mapping of Mamberamo and Raja Ampat in Papua Province. Unpublished report. Forest Watch Indonesia–Conservation International Indonesia Papua Program. Forest Watch Indonesia. 2005. Database Perusahaan Kehutantan. www.fwi.or.id/cgi-bin/ perusahaan.cgi. Accessed 17 February 2005. Forest Watch Indonesia/Global Forest Watch. 2002. The State of the Forest: Indonesia. Forest Watch Indonesia, Bogor, and Global Forest Watch, Washington, D.C. Freeport-McMoRan Copper & Gold, Inc. 2001. Working to produce value. 2000 annual report. Freeport-McMoRan Copper & Gold, Inc. 2002. Response from Freeport-McMoRan Copper & Gold Inc. to the Draft Report of Minerals, Mining and Sustainable Development (MMSD) Inc. Correspondence to MMSD project posted on IIED website. www.iied.org/mmsd/draftreport/rcv_comments.html and www.iied.org/ mmsd/mmsd_pdfs/comments_freeport.pdf. Accessed 24 February 2005. Freeport-McMoRan Copper & Gold, Inc. 2003. Real assets/real value. 2002 annual report. Freeport-McMoRan Copper & Gold, Inc. 2004. The strength of our metals. 2003 annual report. Friends of the Earth. 2004. Greasy palms—palm oil, the environment and big business. Summary report. Goddard Institute for Space Studies. 2005. Global temperature trends: 2004 summation. www.giss.nasa.gov/data/update/gistemp/2004/. Accessed 15 February 2005. Hannah, L., and T.E. Lovejoy (eds.). 2003. Climate Change and Biodiversity: Synergistic Impacts. Advances in Applied Biodiversity Science No. 4, August 2003. Center for Applied Biodiversity Science. Conservation International, Washington, D.C. Hansen, J., L. Nazarenko, R. Ruedy, M. Sato, J. Willis, A. del Genio, D. Koch, A. Lacis, K. Lo, S. Menon, T. Novakov, J. Perlwitz, G. Russell, G.A. Schmidt, and N. Tausnev. 2005. The earth’s energy imbalance: confirmation and implications. Science 308: 1431–1435. International Crisis Group. 2002. Indonesia: resources and conflict in Papua. ICG Asia report no. 39, 13 September 2002. International Crisis Group, Jakarta/Brussels. International Crisis Group. 2003. Dividing Papua: how not to do it. Indonesia Briefing, 9 April 2003. International Crisis Group, Jakarta/Brussels. International Federation of Chemical, Energy, Mine and General Workers’ Unions. 1998. Rio Tinto Tainted Titan. The Stakeholders Report 1997. International Foundation for Election Systems. 2003. Public opinion survey Papua Indonesia. February 28, 2003. International Union for the Conservation of Nature and Natural Resources. 2004. 2004 IUCN Red List of Threatened Species. www.iucnredlist.org. Accessed 15 February 2005. International Union for the Conservation of Nature and Natural Resources. 2005. The IUCN Red List of Threatened Species. Threats Authority File (Version 2.1). www.iucn.org/themes/ssc/sis/authority.htm. Accessed 15 February 2005. International Union for the Conservation of Nature and Natural Resources/Species

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Threats to Biodiversity / 1227 Survival Commission. 2005. The IUCN/SSC Invasive Species Specialist Group (ISSG). www.issg.org. Accessed 15 February 2005. Invasive Species Specialist Group. 2005. ISSG Global Invasive Species Database. www.issg.org/database. Accessed 15 February 2005. Johansen, B.E. 2002. Indigenous Peoples and Environmental Issues: An Encyclopedia. Irian Jaya/Papua New Guinea. www.ratical.org/ratville/IPEIE/IJ_PNG.html. Kaluwin, C., J. Ashton, and S. Saluei (eds.). 2002. Papua New Guinea. Initial national communication under the United Nations Framework Convention on Climate Change. November 2002. 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. December 2003. Indo-Pacific Conservation Alliance, Washington, D.C. Kennedy, D. 1998. Risky business: the Grasberg gold mine. An independent annual report on P.T. Freeport Indonesia. Project Underground, Berkeley, California. Malcolm, J.R., C. Liu, L.B. Miller, T. Allnutt, and L. Hansen. 2002. Habitats at Risk: Global Warming and Species Loss in Globally Significant Terrestrial Ecosystems. World Wildlife Fund–World Wide Fund for Nature, Gland, Switzerland. McKenna, S.A., G.R. Allen, and S. Suryadi (eds.). 2002. A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, D.C. Mertens, B. 2002. Spatial analysis for the Rapid Assessment of Conservation Economics (RACE) in Papua. 1st Task Report, July 10, 2002. Center for International Forestry Research (CIFOR), France. Millennium Institute. 2003. Threshold 21 model project for Conservation International: strategic options for Papua, Indonesia. Miranda, M., P. Burris, J.F. Bingcang, P. Shearman, J.O. Briones, A. La Vina, and S. Menard. 2003. Mining and Critical Ecosystems: Mapping the Risks. World Resources Institute, Washington, D.C. Mittermeier, R.A., P.R. Gil, M. Hoffman, J.D. Pilgrim, T.M. Brooks, C.G. Mittermeier, J. Lamoreux, and G.A.B. da Fonseca. 2005. Hotspots Revisited: Earth’s Biologically Richest and Most Threatened Terrestrial Ecoregions. CEMEX, Mexico City. Mittermeier, R.A., C.G. Mittermeier, P.R. Gil, J.D. Pilgrim, W.R. Konstant, G.A.B. da Fonseca, and T.M. Brooks. 2002. Wilderness: Earth’s Last Wild Places. CEMEX. Mexico City. Mock, G. (ed.). 2000. Linking people and ecosystems. In World Resources 2000–2001: People and Ecosystems: The Fraying Web of Life. World Resources Institute, Washington, D.C. Muller, K. (in prep.). The biodiversity of New Guinea. On-line version. http:// papuaweb.org/dlib/up/muller-ngb/index.html. Myers, N. 1997. The rich diversity of biodiversity issues. In Marjorie L. Reaka-Kudla, Wilson, D.E., and Wilson, E.O. (eds.) Biodiversity II. Understanding and Protecting Our Biological Resources. National Academy of Sciences. Joseph Henry Press, Washington, D.C. Myers, N., R.A. Mittermeier, C.G. Mittermeier, G.A.B da Fonseca, and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853–858. Nature Conservancy. 2004. Bintuni Bay. Protecting the world’s third largest mangrove area. Factsheet. http://nature.org/wherewework/asiapacific/indonesia/work/ art13456.html. Accessed 20 February 2005. Pimm, S.L. 1997. Extinctions, geographic ranges, and patterns of loss. Plenary address at ‘‘Humans and other catastrophes’’ symposium. American Museum of Natural History, New York. www.amnh.org/science/biodiversity/extinction/Day1/bytes/PimmPres.html.

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1228 / scott frazier Polhemus, D.A., R.A. Englund, and G.R. Allen. 2004. Freshwater biotas of New Guinea and nearby islands: analysis of endemism, richness, and threats. Bishop Museum Technical Report 31. Prepared for Conservation International. Polhemus, D.A., and J.T. Polhemus. 1998. Assembling New Guinea: 40 million years of island arc accretion as indicated by distributions of aquatic Herteroptera (Insecta). Pp. 327–334 in Hall, R., and J.D. Holloway (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden. PT Freeport Indonesia. July 2004. Riverine tailings transport. Freeport-McMoRan Copper & Gold Inc. Purvis, A., P.-M. Agapow, J.L. Gittleman, and G.M. Mace. 2000. Nonrandom extinction and the loss of evolutionary history. Science 288: 328–330. Putz, F.E., K.H. Redford, J.G. Robinson, R. Fimbel, and G.M. Blate. 2000. Biodiversity conservation in the context of tropical forest management. World Bank Environment Department working paper no. 75. World Bank, Washington, D.C. Stuart, S. 1999. Species: unprecedented extinction rate, and it’s increasing. International Union for the Conservation of Nature and Natural Resources (IUCN) press release. www.iucn.org/info_and_news/press/species2000.html. Accessed 15 February 2005. Subsidiary Body on Scientific, Technical and Technological Advice. 2003. Pilot assessments: the ecological and socio-economic impact of invasive alien species on island ecosystems. Note by the Executive Secretary. UNEP/CBD/SBSTTA/9/INF/33. 5 November 2003. Convention on Biological Diversity. Supriatna, J. (ed.). 1999. The Irian Jaya Biodiversity Conservation Priority-Setting Workshop. Biak, 7–12 January 1997. Final Report. Conservation International, Washington, D.C. Suryadi, S., A. Wijayanto, and M. Wahyudi. 2004. Survey Pasar/Monitoring Perdagangan Hidupan Liar di Kabupaten Manokwari dan Jayapura. Conservation International Indonesia dan Seksi Konservasi Wilayah I Manokwari Balai Konservasi Sumber Daya Alam Papua II. Jakarta. Tangguh Independent Advisory Panel. October 2002. First report on Tangguh LNG Project. Thaman, R.R. 2002. Threats to Pacific Island biodiversity and biodiversity conservation in the Pacific Islands. Development Bulletin 58: 23–27. Thomas, C.D., A. Cameron, R.E. Green, M. Bakkenes, L.J. Beaumont, Y.C. Collingham, B.F.N. Erasmus, M.F. De Siqueira, A. Grainger, L. Hannah, L. Hughes, B. Huntley, A.S. Van Jaarsveld, G.F. Midgley, L. Miles, M.A. Ortega-Huerta, A.T. Peterson, O.L. Phillips, and S.E. Williams. 2004. Extinction risk from climate change. Nature 427: 145–148. Thomas, J.A., M.G. Telfer, D.B. Roy, C.D. Preston, J.J.D. Greenwood, J. Asher, R. Fox, R.T. Clarke, and J.H. Lawton. 2004. Comparative losses of British butterflies, birds, and plants and the global extinction crisis. Science 303: 1879–1881. Thompson, H. 1996. Indonesia: development, degraded rainforests and decreasing global biological diversity. AntePodium, Victoria University of Wellington. Thorpe, J., and B. Godwin. 1999. Threats to biodiversity in Saskatchewan. Saskatchewan Research Council Publication No. 11158-1C99. UNESCO. 2005. The World Heritage List. http://whc.unesco.org/pg.cfm?cid31& id_site955. Accessed 15 February 2005. United Nations Environmental Program–WCMC. 2001. Lorentz National Park. Natural site data sheet. www.wcmc.org.uk/protected_areas/data/wh/lorentz.html. Accessed 15 February 2005.

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Threats to Biodiversity / 1229 van Gelder, J.W. 2001. European banks and palm oil and pulp and paper in Indonesia. Research paper by Profundo, prepared for World Wildlife Fund International. van Zyl, D., M. Sassoon, C. Digby, A.-M. Fleury, and S. Kyeyune. 2002. Grasberg Riverine disposal case study. Appendix J in Mining for the Future. Institute for Environment and Development and World Business Council for Sustainable Development. Wakker, E. 2000. Funding forest destruction: the involvement of Dutch banks in the financing of oil palm plantations in Indonesia. AIDEnvironment, in co-operation with Jan Willem van Gelder, Contrast Advies, and the Telapak Sawit Research Team, Amsterdam and Bogor. Commissioned by Greenpeace Netherlands. Wakker, E. 2005. Greasy palms: the social and ecological impacts of large-scale oil palm plantation development in Southeast Asia. Friends of the Earth, Washington, D.C. Willis, K.J., L. Gillson, and T.M. Brncic. 2004. How ‘‘virgin’’ is virgin rainforest? Science 304: 402–403. World Bank Group. 2001. Indonesia. Environment and natural resource management in a time of transition. World Wildlife Fund. 2005. Global warming. Changing a delicate balance. http:// panda.org/about_wwf/what_we_do/climate_change/problems/warming. cfm. Accessed 28 February 2005. YPMD (Irian Jaya Rural Community Development Foundation: Yayasan Pengembangan Masyarakat Desa). 2004. Prospect and constraints of natural resource management in Papua’s Special Autonomy: conflicts of laws and regulations of NRM in Special Autonomy. Draft report.

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7.2. Setting Priorities and Planning Conservation in Papua john burke burnett o n se r v a ti o n p ri o r i ty s e t ti n g (CPS) is a form of spatial planning based on the overall objective of maximizing preservation of the natural environment and its biota using biological, economic, cultural, and social criteria, and making practical and scientifically sound land-use recommendations to decision makers and natural resource owners. Biodiversity conservation requires substantial human and financial resources, both of which are in limited supply. Papua is globally recognized as a high-priority conservation area (Sujatnika et al. 1995; Supriatna 1999; Wikramanayake et al. 2000), but the limited availability of funding for conservation efforts, as well as the relative scarcity of motivated and trained individuals to implement them, present practical constraints to implementing the level of conservation activities that would be required to adequately protect Papua’s biodiversity. Because of these limitations, it is necessary to plan conservation activities carefully and strategically. In short, conservation must have a strategy for maximizing overall protection for the fewest dollars. The ways that conservationists create priorities and the methods they use are both varied and complex. This chapter attempts to summarize priority-setting processes, beginning with a general overview, followed by an examination of the process in Papua. The discussion is primarily limited to terrestrial conservation. Marine conservation priority setting at this time is in its infancy (Lourie and Vincent 2004; Olson 1997).

C

Challenges to Conservation Priority Setting

the ‘‘patchiness’’ of biodiversity Biodiversity is not evenly distributed. The patchy nature of species and habitat distributions is well known to biologists and biogeographers. Some areas have high concentrations of species, others have low concentrations. Which areas are most important to protect also depends greatly on the criteria used to define ‘‘importance.’’ For example, certain areas may be extremely rich in one taxonomic group, such as butterflies, but not in another, such as mammals. Or, as is frequently the case, some areas, such as lowland forests, may have the highest species richness, but those species may have relatively large ranges. Compare this with another patch of forest in the mountains that has fewer species but more endemics. Which area is more important to protect? Moreover, species richness, endemism, and rarity are not the only criteria that should be considered. Genetic uniqueness, habitat diversity, and degree of threat are also important, as will be discussed later. Marshall, A. J., and Beehler, B. M. (eds.). 2006. The Ecology of Papua. Singapore: Periplus Editions.

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issues of scale To make matters more complicated, the degree of patchiness in the distribution of species or habitats varies with scale. For example, a priority-setting assessment at a global scale may identify particular areas as critical ecoregions or hotspots, while a smaller-scale regional or local level exercise using similar criteria will reveal much finer details, allowing clarification of which areas are of greater or lesser importance. This influences how conservation priorities are defined, because a broad-scale classification scheme will identify only a few types of critical ecoregions, and thus only a relatively few areas will be identified for protection. A localscale exercise, on the other hand, will distinguish among habitat subtypes. This allows a more refined, targeted approach that can more accurately represent the full complexity of the larger ecoregion, with specific conservation measures tailored to each local area. Analysis at different scales reveals differences in habitat resolution or species distribution, including areas of high richness or areas that have already been converted to other uses (e.g., urban or agricultural areas) and are thus nearly devoid of conservation importance. The term ‘‘ecosystem’’ is applied to a variety of scales, so conservation decision makers cannot always readily compare ecosystem units (Perlman and Adelson 1997). For example, World Wildlife Fund (WWF) has defined Papua and PNG’s Trans-Fly region as a single Ecoregion, based the predominance of savanna habitat (Chapter 5.12). However, WWF also recognizes that the Trans-Fly also has numerous areas of dry evergreen forest, seasonal swamplands, and lakes (Wikramanayake et al. 2000). Although the Ecoregion approach is primarily a means of representation rather than prioritization, conservation priority setting must be carried out at a finer scale of analysis, as WWF has indeed done for this area (Chapter 7.6). It is important to note that any system of classification runs into the scale problem, but the Ecoregion approach is nonetheless very valuable for understanding broad patterns of biodiversity, and for protecting representative habitat and species within Papua. But given the degree of scientific uncertainty about species and their habitat distributions and ecological requirements, and the knowledge that there are other development realities in Papua, biological criteria cannot be the only factor used to identify the specific areas that are the most important for intensive conservation efforts.

limitations imposed by lack of scientific knowledge about papua Conservation priority setting exercises are carried out using data on geology, forest cover, known species distributions, vegetation types, habitation, economic development plans, and so on. Although it is impossible to obtain complete data on the biology or ecology of any area, the data available for Papua are especially poor. Biodiversity field surveys and ecological studies in Papua have been very limited in comparison with those for other parts of Indonesia or Papua New Guinea (PNG). Thus, for most species, scientists do not have sufficient information to

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know which areas of Papua are the most important for richness or endemicity, or what long-term ecological conditions allow species to persist. This lack of adequate biological information makes it more challenging to set conservation priorities for Papua than for other better documented areas. As a result, conservation priority setting typically uses biodiversity surrogates (e.g., indicator species, or other proxy information such as forest type, that may indicate the overall levels of species richness or endemism) as substitutes for comprehensive biodiversity inventory data. For some taxonomic groups (particularly birds and other vertebrates) enough is known to make scientifically sound estimates. For other less well-documented taxa, conservation priority setting relies on basic ecological and biogeographic principles to estimate priority habitat areas.

limitations imposed by social, economic, and political context Biodiversity is not the only criterion considered in conservation priority setting. Conservation is, in the end, a political issue. It requires political support—by landowners as well as local, provincial, and national levels of government—for land use other than unmitigated economic development. All these stakeholders have critical roles in decisions about the disposition of land and natural resources. Conservation priority setting must take these perspectives and factors into account. As Margules and Pressey (2000) note, conservation priority setting ‘‘is an activity in which social, economic, and political imperatives modify, sometimes drastically, scientific prescriptions.’’ Ranking conservation priorities must take into consideration factors such as which areas which are under active logging, where local conservation partners are weak or absent, and where there is political discord. This does not mean that ecologically critical habitat is not ranked as a high conservation priority where these social and political conditions exist; rather, conservation priority setting must strike a pragmatic balance. Obviously it would make little sense to invest scarce conservation resources in an area that is certain to be converted to agriculture. On the other hand, conservation priority setting provides supporting material for the construction of more sound policies, policy implementations, and interventions, and it may provide sufficiently strong arguments to stop or reverse some planned land uses (such as the now-shelved Mamberamo hydroelectric dam project; Supriatna 1999).

The Conservation Priority-setting Process Margules and Pressey (2000) provide a comprehensive and incisive overview of the conservation priority-setting process. Since ancient times and in many cultures, humans have established reserves both to preserve the animals within them and to slow or halt encroachment from external processes such as logging, agriculture, or human settlement (Harrison 1992). However, reserves—including those created recently in the developed world—have often been established on the basis

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of scenic beauty (the so-called ‘‘rocks and ice’’ syndrome) rather than on the basis of their contribution to the protection of representative patterns of biodiversity (Margules and Pressey 2000; Terborgh 1999). Tragically, the lack of rational reserve-planning based on biodiversity criteria has led, in many parts of the world, to an unfortunate triage approach to conservation in which extensive development has largely destroyed or fragmented much original habitat and caused many species to become endangered or extinct. In the United States, for example, protecting remaining threatened and endangered species requires expensive interventionist approaches to protect habitat critical to their survival. This might have been avoided or at least mitigated had the natural reserve system been designed and applied using biodiversity conservation planning principles. Papua is fortunate in that these types of ‘‘last ditch’’ approaches are not yet necessary. But to avoid slipping into this situation, it is necessary to make informed decisions today that are based on sound, scientifically robust, and achievable criteria about which areas to protect in order to maximize the representation of Papua’s biodiversity. The goal of conservation priority setting is to provide an objective and realistic spatial template that can be used to guide conservation and development planning.

stages of conservation priority setting Conservation priority setting takes place in several stages. The general framework described here applies to CPS at global, regional, or local scales. Though details of the requirements of each scale vary, and a great deal of technical complexity may be involved, the basic process for setting conservation priorities (adapted from Groves 2003; Margules and Pressey 2000; The Nature Conservancy et al. 2003) is summarized below.

Stage 1: Measuring and Mapping Biodiversity • Compile existing data (physiographic, geological, meteorological, hydrological, forest cover, biological specimen collection point data and range maps, demographic, economic, etc.). • Review all data sets for accuracy and assess existing biological data sets for appropriateness as surrogate measures of biodiversity.

Ensuring the maximal integrity and accuracy of data sets is essential. Because biodiversity is highly complex, biodiversity surrogates (indicator species, habitat types, etc.) should be used carefully to ensure they accurately represent this complexity and the differences and similarities among areas. Margules and Pressey (2000) note that ‘‘there is no best surrogate’’: the use of higher-level biological information (e.g., species assemblages, habitat types) loses some biological precision but may add valuable information on ecosystem processes that would otherwise not be included. Museum and herbarium collections contain important information (usually in the form of species point data) on the known distributions of species. This is especially valuable for local-scale planning where more extensive surveys have been carried out (e.g., Cyclops Mts). However, because most of

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Papua is notoriously undercollected, using information on species’ distributions will be of limited utility until more biological surveys are undertaken. One general issue is how to define biodiversity. Biodiversity can be analyzed in several different ways, including phylogenetic diversity (in which taxonomic rarity—species with less genetic similarity to other species—is prioritized), species richness, species endemism, community assemblages, and habitat types. However, as these targets or priorities are often (though not always) complementary, most conservation priority-setting exercises strike a pragmatic balance among them.

Stage 2: Identifying Conservation Targets • Set specific, quantitative conservation targets for species, vegetation types, or other environmental features. (For example, identify areas with the highest species richness; ensure the protection of 95% of endemic birds; all major vegetation types and species assemblages must be represented in at least one area with minimum size of 20,000 ha, etc.). • Set quantitative targets for minimum size, connectivity, or other reserve design criteria. • Identify qualitative targets, such as that priority areas must be free of previous logging, planned transmigration settlements, oil or gas development, and so on.

Conservation targets can be defined in a variety of ways, including preventing the extinction of threatened or endangered species (using estimates of minimum viable population size), maximizing representation of biodiversity, maintaining the provision of ecological services, or maintaining evolutionary processes. These are not mutually exclusive targets, but the choice will shape the priorities set and the conservation outcomes, and so goals should be carefully selected and defined. Most conservation plans are based on various ecological indices, including the occurrence of species (e.g., species richness, endemic species, or threatened species), habitat types, known species population corridors, forest or vegetative cover, and abiotic information (e.g., soils, rainfall, hydrology, topography). These data sets are mapped and entered into a computer-based geographic information system (GIS) which represent these various elements as layers. This allows layers to be overlaid in order to visualize the spatial distribution of biodiversity, land system, roads, and other features. GIS layers also include data such as land use patterns, logging concessions, land tenure, and other characteristics that indicate the level of threat to particular areas. Priority areas are those which maximize representation of target indices. Soule´ and Sanjayan (1998) assert that setting targets based on percentage of land area to be conserved (typically 10–12%) should be avoided. This is because it is relatively easy to include areas with low economic and biodiversity value (e.g., previously logged areas or other least productive or least threatened landscapes). As Margules and Pressey (2000) note, more is not necessarily better. Moreover, conservation targets in the range of 10% of land area are inadequate to protect both wide-ranging and rare/restricted-range species, especially in the tropics. Tar-

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gets closer to 20–50% of land area may be required to protect the full spectrum of biodiversity in a region (Soule´ and Sanjayan 1998). Recent evidence suggests that the choice of diversity indexes used to guide conservation priorities largely determine the outcome of the effort (Orme et al. 2005). It was previously thought that there would be broad overlap of areas despite use of different criteria (viz., ‘‘species richness,’’ ‘‘threatened species,’’ or ‘‘endemic species’’), but there may be less congruence in priority areas than anticipated. Intriguingly, the study by Orme et al. (2005) noted that the selection of priority areas using ‘‘endemic species’’ as the criterion actually protected a greater proportion of overall species richness and of threatened species than either of those two indexes alone. Possingham and Wilson (2005) speculate that this phenomenon may be associated with large-scale topography. Possingham and Wilson also note that caution is necessary when using this method for conservation priority setting, because without good data on the distributions of poorly known taxa it can yield spurious results. (The study by Orme et al. was based on data on birds, which is one of the best-known groups.) This observation seems expecially pertinent to Papua, where the paucity of data for nearly all taxonomic groups (not to mention a high degree of taxonomic uncertainty) is striking. In general, however, this finding lends some support for the use of endemism as a criterion for determining priority areas, although Orme et al. (2005) recommend using multiple indices to ensure comprehensiveness and accuracy. Emerging field research, although still in the preliminary stages, is showing that New Guinea species display very high beta diversity along both north-south (i.e., elevational) and east-west (cross-island) transects. Mack (pers. comm.) suggests that early results, and other biogeographic data, indicate low levels of gene flow even among populations that appear to be connected by continuous forest. The result is a lack of mixing between populations and species—contrary to what might be assumed upon seeing thousands of square kilometers of continuous forest. New Guinea’s high species turnover reflects the island’s highly complex geological history. This history has created numerous isolating mechanisms (i.e. geographical barriers and elevational gradients) that generate diversity and has also created isolated pockets of stable landforms that affect the extinction rates of small-range species over evolutionary time (Possingham and Wilson 2005). As a result, a network of protected areas that covers centers of endemism is critical to preserve the full spectrum of New Guinea biodiversity (A.L. Mack, pers. comm.).

Stage 3: Review Existing Reserves and Select Additional Reserves • Assess the extent to which quantitative conservation targets identified in Stage 2 have been fulfilled in existing conservation areas. • Identify and assess level of threats to (a) areas under-represented in existing conservation areas and (b) areas that are vital to fulfilling quantitative conservation targets set in Stage 2. • Consider existing reserves as fixed ‘‘constraints.’’ Then (a) prioritize expansions of

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1236 / john burke burnett existing reserves rather than create new areas, or (b) create new reserves, based on principles of reserve design (e.g., proximity, connectivity). • Identify new reserve areas, taking into consideration such factors as local partner institutions and capacity, likelihood of stakeholder approval, existing or planned development projects, conservation budgets, opportunity costs of land uses other than conservation.

Stage 3 of the conservation planning process is typically conducted in workshops led by experts. Because various conservation target indices both within and among taxonomic groups will often result in identification of different priority areas, one of the first objectives is to integrate these priority areas to achieve consensus in the planning group. It is often difficult to achieve consensus without some friction. For example, some taxa, such as plants, may have many species with extremely small ranges, whereas other taxa may require much larger areas. Inevitably, a pragmatic approach to reaching consensus on conservation goals is required. If good range maps are available, GIS analysis can be used to show areas of overlap in conservation target areas. Pragmatism is also required to determine priorities among areas, by assessing threat levels to different areas, and identifying the areas that offer the greatest likelihood of success. Gap analysis is used to determine which areas that achieve conservation targets occur within existing parks and reserves, and which important targets are not met. The gap analysis algorithms yield complementarity scores, which are measures of how much the conservation target would gain (e.g., how many new, previously unprotected species would be protected) by adding each new reserve area. To facilitate gap analysis, the conservation priority-setting process selects areas by using an algorithm that incorporates spatial constraints reflecting the costs (both real costs and opportunity costs) that would be involved in adding each new area. Because formal establishment of new reserves is highly bureaucratic and requires a significant investment of time and labor, precedence is usually given to biodiversity priority areas that occur inside existing reserves. Gap analysis criteria are different for fragmented areas and for areas with large blocks of intact habitat, as occurs in Papua. Fragmented habitat requires consideration of additional factors such as species corridors and minimum viable population assessments. Because Papua contains more intact habitat than most tropical forest regions, current conservation options for Papua are much less constrained than for other regions of the world. Because of this, as well as the relative paucity of biological data, conservation priority-setting analysis in New Guinea can emphasize basic principles of reserve design to maximize conservation targets.

setting conservation priorities Reserve Design for Terrestrial Areas The clearance of tropical forest for logging or agricultural conversion results in the loss of the habitat that is essential to maintain biodiversity. Forest destruction leads to the fragmentation of habitat, creating forest ‘‘islands.’’ Each of these ‘‘is-

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lands’’ can support fewer varieties and lower abundances of species than can unfragmented forests. This is especially true for species that require large areas to sustain viable long-term populations, such as the New Guinea Harpy Eagle (Harpyopsis novaeguineae), and perhaps cassowaries (Casuarius). Forest fragments are subject to lower rates of in-migration (i.e., colonization) and higher rates of population extinction than larger areas. To avoid extinction, populations must be of sufficient size (i.e., minimum viable population, MVP) to maintain genetic variability. In addition, random events, such as changes in reproductive or survival rates, weather changes, disease outbreaks, changes in food availability, fires, drought, and so on, will inevitably occur, and these events affect the long-term viability of a population in a particular area. Metapopulations (sets of populations that may replace one another if one population becomes locally extinct) can persist only when forest patches of a minimum size are present, and these patches are located close enough to allow (re)colonization. Finally, ‘‘edge effects’’ may create unsuitable conditions for populations. Perimeters of forest blocks are subject to higher rates of predation, changes in micro-climates, wind damage, and increased anthropogenic activities; isolated blocks of fragmented forest have longer perimeters than a single block of forest containing the same area. In short, habitat fragmentation inevitably leads to greater risk of population extinctions. The larger the protected area, the more species and larger, more viable populations it can contain. Biogeographic theory has established general principles for the design of a protected area system to minimize the impact of fragmentation and to maximize the ability of species and habitat to persist for long periods (Diamond 1975; Diamond 1986; Dobson 1996; Perlman and Adelson 1997; Petocz 1989; see Figure 7.2.1). MacKinnon et al. (1986) describe the criteria for the size, shape, and distance between protected areas, which can be summarized as: big reserves are better than small reserves; one large reserve is better than several small reserves; reserves close together are better than those spaced further apart; and reserves clustered in a circular pattern are better than those spaced linearly. Any of these principles are further enhanced if areas of minimally disturbed (and sustainably managed) habitat surround the reserves (i.e., single or multiple reserve areas are located within a matrix of minimally disturbed habitat).

Marine Conservation Priority Setting Conservation priority setting for marine areas requires a different approach than for terrestrial systems because the physical, biological, and socio-political characteristics of marine systems are different. Biogeographic data are harder to obtain for marine areas and biogeographic boundaries are more difficult to define (Lourie and Vincent 2004). A recent quantitative analysis has revealed that a high percentage of reef fish, corals, snails, and lobsters have highly restricted ranges, clustered into centers of endemism similar to terrestrial hotspots (Roberts et al. 2002; Chapter 5.2). The restricted ranges of these taxa means that they are more vulnerable to extinction. As Roberts et al. (2002) note, focusing conservation action on these

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Figure 7.2.1. Biogeographic principles in reserve design. Source: after MacKinnon et al. (1986).

centers of endemism could be highly effective in preventing species loss, but it is not yet clear whether this focused strategy would work for protecting more widespread species. Despite ongoing efforts to develop more quantifiable biogeographic criteria for marine conservation priority setting, most approaches have been largely opinion based and ad hoc (Lourie and Vincent 2004). This does not negate the value of these efforts, but because of the magnitude of current and future threats to coral reefs and other marine ecosystems, a more scientifically

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rigorous approach to marine conservation priority setting is needed to focus efforts more effectively. Beck (2003) suggests the most effective approach involves analytical methods to determine and classify biogeographical boundaries, identifies a range of potential conservation sites using specific target criteria, and follows a consensus-building process involving stakeholders (particularly local communities).

Global and Regional Approaches to Conservation Priority Setting

conservation international’s hotspots and wilderness areas Conservation International (CI) has devised two global-scale conservation priority setting approaches, one aimed at Hotspots and the other at Wilderness Areas. Because global biodiversity is unevenly distributed, these strategies identify areas that contain the highest concentrations of global biodiversity, and prioritize these areas on the basis of presence of endemic species and degree of threat (defined as percentage of original habitat remaining). The conservation strategy implicit in the Hotspots approach is to identify and protect first the areas that are richest in species and most threatened. Conservation International’s Hotspots approach is based on a seminal paper by Norman Myers (1988) that identified ten tropical forest areas characterized both by exceptional levels of plant endemism and by high levels of habitat loss. In 1989, CI adopted and adapted Myers’s method to include specific quantitative criteria, and then conducted an extensive global review. To qualify as a Hotspot, a region must contain at least 1,500 endemic species of vascular plants (or at least 0.5 percent of the world’s total), and must have lost at least 70% of its original habitat. The first review in 1999 identified 25 biodiversity Hotspots, including Sundaland (Java, Sumatra, Bali, Borneo, Peninsular Malaysia) and Wallacea (Sulawesi, Maluku, Nusa Tenggara), while a second review in 2005 identified 34 priority regions. Covering only 2.3% of the earth’s land surface, these 34 sites contain at least 150,000 endemic plant species (50% of the world’s total), at least 11,980 endemic terrestrial vertebrates (42% of all terrestrial vertebrate species), and 22,022 total terrestrial vertebrate species (77% of the world’s total) (Mittermeier 2005). While Conservation International prioritizes terrestrial conservation efforts in the Hotspots approach, they have also identified five high-biodiversity Wilderness Areas, which are ‘‘vast regions of relatively pristine habitat that sustain large numbers of plant and animal species found nowhere else’’ (Mittermeier 2003). Although some have questioned whether the term ‘‘wilderness’’ is appropriately applied to New Guinea (Mackie 2003), Conservation International defines Wilderness Areas as areas of at least 10,000 square kilometers (1,000,000 hectares or 3,861 square miles) with at least 70% of their original habitat, containing at least 1,500 endemic species of vascular plants (i.e., at least 0.5 percent of the world’s total), and harboring fewer than five people per square kilometer. By these criteria, the

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three largest and most important Wilderness Areas in terms of total and endemic species are Amazonia, the Congo forests in west Africa, and New Guinea.

world wildlife fund’s ecoregions World Wildlife Fund’s (WWF) Ecoregions approach is a method to classify biogeographic areas and help guide conservation priorities by providing a blueprint for broad representation of the world’s major ecosystems. WWF defines an Ecoregion as a ‘‘large unit of land or water containing a geographically distinct assemblage of species, natural communities, and environmental conditions’’ (Olson et al. 2001). By identifying the broadest variety of the world’s habitats, the Ecoregions strategy is designed to conserve the broadest variety of the world’s species and endangered wildlife, as well as whole communities and ecosystems. The emphasis is on protecting representative areas in each of the world’s Ecoregions, to ensure inclusion of the earth’s important biomes, ecosystems, and species. To achieve conservation results that are ecologically viable and maximize the representation of a region’s biodiversity, it is necessary to conserve networks of key sites, migration corridors, and the ecological processes that maintain healthy ecosystems (Wikramanayake 2000). The Ecoregion approach is a system for representing patterns of biodiversity rather than a method for prioritizing conservation action, and has become among the most prominent and widely used such approach (Mittermeier et al. 2005; Wikramanayake et al. 2002). The Nature Conservancy (TNC) and Wildlife Conservation Society (WCS), among others, employ a modified version of the Ecoregion approach to help identify priority areas. One of the strengths of the ecoregional approach is that both conservation strategies and government spatial planning can be developed within a framework that addresses ecological processes that maintain biodiversity and protect wide-ranging species (Groves et al. 2003; Wikramanayake et al. 2002).

endemic bird areas and other approaches BirdLife International uses birds as a biodiversity surrogate for setting conservation priorities (Chan et al. 2004; Sujatnika et al. 1995). BirdLife has two categories of priority classification, Important Bird Areas (IBAs) and Endemic Bird Areas (EBAs). IBAs are key conservation sites that are small enough to be conserved in their entirety and are often already part of a protected area system. They contain significant numbers of one or more globally threatened species, are part of a set of sites that together hold a suite of restricted-range or biome-restricted species, and have exceptionally large numbers of migratory or congregatory species. IBAs are determined using standardized, quantitative criteria based on the occurrence of key bird species that are vulnerable to global extinction or whose populations are otherwise irreplaceable (Chan et al. 2004). EBAs are based on the recognition that a significant percentage ( 25%) of birds are restricted to ranges smaller than 50,000 km2, and that areas where the distributions of two or more of these restricted-range species overlap are particularly rich in both endemic and wide-

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ranging bird species. BirdLife estimates that approximately 70% of EBAs overlap with areas that are similarly important for endemic plants, and possibly other species as well (Sujatnika 1995). On the other hand, Jepson and Whittaker (2002) note that existing data do not necessarily support the assumption of congruence between the distribution of endemic birds and taxa other than small mammals and certain butterflies. WWF’s Ecoregions strategy to represent biogeographic patterns in New Guinea drew heavily on the EBA classification system (Wikramanayake et al. 2002). Other conservation priority-setting approaches are possible, and several have been tested in Papua New Guinea. The BioRAP program, a CPS exercise carried out by Commonwealth Scientific and Industrial Research Organisation (CSIRO) in the 1990s, used abiotic information (soil types, moisture, detailed physiographic information), along with predictive or probabilistic modeling of species distributions, to help determine conservation priority areas (Faith et al. 2001; Margules and Redhead 1995). However, the BioRAP team had the benefit of access to far more detailed and accurate data sets on forest cover, rainfall, and other biological information than exist for Papua, and so the use of this approach for Papua is unrealistic at this time.

Conservation Priority Setting in Papua The most common approach to conservation priority setting, and probably the most appropriate for Papua, with such limited availability of scientific data, is to select areas based on the known occurrences of vertebrate species (Diamond 1986), known centers of endemism (especially for birds, which are relatively well known; Sujatnika 1995; Chapter 2.4), species assemblages and habitat (Wikramanayake 2000), or a combination of all these targets (Supriatna 1999).

scientific data for papua The biodiversity of Papua is very poorly documented. This fact is a significant constraint on the ability of conservation priority-setting exercises to set targets and priorities for Papua. Mack and Alonso (2000) note that as many as 510 species of amphibians and reptiles are known from PNG, compared with 330 for Papua; for mammals the numbers are 227 known in PNG compared to 164 in Papua. This difference reflects the relative lack of collecting in Papua compared to PNG (PNG is itself highly underdocumented, compared to other tropical forests), rather than a significant difference in the levels of biological diversity between the two sides of the island. This lack of basic biological inventory work in Papua is the result of the difficulty many foreign researchers have had in obtaining permission to conduct scientific research, as well as the lack of opportunity and high research costs for many Indonesian scientists. In short, there is a glaring shortage of biological and ecological data for Papua. This gap means that the locations and distributions of many species cannot be identified with precision. Compounding this is the inadequate number of taxonomists (locally, nationally, and internation-

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ally) who can accurately identify Papuan taxonomic specimen material. Lack of coordination between institutional biological information management systems means that much information that does exist is inaccessible, particularly for local researchers. Finally, there have been few ecological studies on particular taxa. Thus there is insufficient information on the behavior, movements, or habitat requirements for most New Guinea species. Ongoing research, mainly on the PNG side, in programs such as Wildlife Conservation Society’s Crater Mountain Research Station, is helping to address this gap, but a great deal of work remains to be done, especially on the Papua side of the island. Data on tenure patterns and land ownership are critically important, but these may be more difficult to obtain in Papua due to the legal uncertainties currently surrounding this issue. Other key types of data, which are available in varying degrees of accuracy, include physiography (geology and terrain), climate and rainfall, hydrology (watersheds), soil types, planned or existing road and transport development, extractive (oil, gas, mining) areas, logging concessions, oil palm plantations, demographic patterns and socioeconomic data related to threats, linguistic and cultural areas, forest cover, and others. As Jepson and Whittaker (2002) have noted, the Regional Physical Planning Program for Transmigration (RePPProT) maps published in 1986 are not very useful for conservation planning purposes, though they do indicate areas of agricultural suitability, hydrological data, and soil type (RePPProT 1986).

basic reserve design in papua Petocz (1989) describes the following biological factors as having been the criteria for the basic design of the protected area system in Papua (ca 1975–1989): coverage of the entire altitudinal spectrum of the province, and inclusion of the complete altitudinal gradient in each mountain reserve; inclusion of each major center of endemism and diversity as known at the time; coverage of representative crosssections of habitats within each area of that particular ecoregion; inclusion of substantial tracts of lowland rainforest in order to secure the floral diversity which is highest in the lowlands; protection for species with large area requirements, by linking, as closely as possible, the Mamberamo-Foja, Jayawijaya, and Lorentz reserves; sufficient coverage of special or important habitats such as mangroves, special forest communities, major vegetation transition zones, and the driest regions of the province; inclusion of important habitat for migratory species; inclusion of small offshore islands that are critical to colonies of nesting birds; inclusion of major nesting beaches and feeding areas for marine turtles.

the 1996–1998 irian jaya priority-setting program Beginning in 1996, Conservation International undertook a three-year program known as the Irian Jaya Priority-setting Program (PSP) to define conservation priorities for Papua (then known as Irian Jaya). CI’s approach was to develop a participatory, expert-driven, consensus-based process that incorporated the best available knowledge for the region, to create a set of recommendations for conser-

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vation priorities in Papua, and to integrate this information into the provinciallevel spatial planning (tata ruang) carried out by the Provincial Development Planning Bureau (BAPPEDA: Badan Perencanaan Pembangunan Daerah), a function that is now carried out at the district or regency (kabupaten) level with assistance from the provincial government. The following summary is based on CI’s report (Supriatna 1999). Conservation International began collecting data to develop a comprehensive biodiversity information system for Papua. These data included biological information based on museum specimen collections; ecoregions, physiographic, hydrological, and rainfall information; forest cover; protected area boundaries; demographic and sociolinguistic patterns; and development plans (oil and gas, mining, logging concessions, transmigration). This information was digitized and entered as GIS layers. In early 1997, Conservation International convened a six-day workshop in Biak that included over 100 Indonesian and international scientific experts, local Papuan community representatives, government planners, and other stakeholders, to seek consensus on priority conservation areas. Participants were split into nine thematic groups based on their area of expertise: Plants, Insects and Arthropods, Reptiles and Amphibians, Birds, Mammals, Freshwater Systems, Marine/Coastal, Socioeconomic, and Conservation Implementation. In Stage One of the workshop (days 1–3), the taxonomic groups used biological and ecological criteria to assess and define conservation priority areas on maps. Taxonomic group recommendations were digitized and priority area designations were entered into the GIS system. The Socioeconomic Group analyzed demographic trends, cultural patterns, and the socioeconomic implications of various land use and conservation strategies. The Conservation Implementation Group assessed local institutional capacity for carrying out conservation goals as well as the most effective strategies for implementing the conservation targets identified by the taxonomic groups. Plenary sessions were held daily to allow cross-disciplinary discussion and to facilitate understanding and consensus. In Stage Two (day 4) of the workshop, participants were re-assigned to four geographical interdisciplinary regional groups: Northern, Southern, Vogelkop, and Offshore Islands. These groups were gathered to discuss the various biological, socioeconomic, and cultural criteria defined in the thematic group sessions, and to reach consensus on priority conservation areas that met the targets defined in Stage One. Each area was assessed using four broad criteria containing numerous subtargets. These four evaluation criteria were: 1. Level of biological importance of the area, based on: species richness, species endemism, phylogenetic diversity, phylogenetic endemism, rare or endangered species, beta diversity, and biological uniqueness. 2. Degree of human pressures and threats, based on: habitat conversion, logging, overhunting/poaching, alien invasive species, development projects, human population growth, and the loss of traditional values and indigenous knowledge. 3. Urgency for conservation action, based on: the need for a new conservation area

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1244 / john burke burnett necessary to meet particular conservation targets, improving protected area management, the re-design of a conservation area, implementing sustainable natural resource use, preventing environmental degradation, increasing public awareness, and/or to restoring habitats and increasing species populations. 4. Importance of additional research effort in the area, based on: the current level of scientific knowledge, and gaps in that knowledge with respect to biological resources or ecological processes, human ecology, and natural resource use.

Because of insufficient data for some taxonomic groups, ‘‘best scientific judgment’’ was sometimes used in assessing the criteria and setting priorities. A semiquantitative scale ranking relative importance was used to score each site. The resulting consensus on conservation priority areas is reflected on a map produced based on Conservation International (see color insert) which ranked conservation priorities into six categories: (1) areas of documented importance and requiring conservation protection; (2) areas inadequately surveyed but thought to be biologically significant; (3) areas of biological significance but under human pressure, thus requiring integrated management of biological resources; (4) marine areas of documented significance; (5) mangrove areas requiring special protection; and (6) lower priority areas reflecting the lack of data to verify priority status. The last category engendered particularly contentious debate. Conservation in Papua needs a clear set of priority areas based on defined metrics. However, there was understandable discomfort in categorizing an area as ‘‘lower priority’’ (which, because most areas are unsurveyed, may or may not be accurate by biological criteria). The concern was that this could be misconstrued or misused by outside planners or commercial enterprises as an invitation to unbridled development. This ‘‘lower priority’’ designation, accurate or not, could inadvertently subject an area, its inhabitants, and its biodiversity to inappropriate forms of development. This was partially resolved by defining ‘‘low priority’’ areas as lower in priority ‘‘assuming effective conservation has been achieved in all the designated priority areas’’ (i.e., categories 1–5; Supriatna 1999). The scientific consensus at the workshop was that the existing protected area system in Papua was relatively well designed, containing a high percentage of the most important ecosystems and areas of endemicity (Supriatna 1999). Other biologically critical areas not included in the original protected areas system were identified, including most notably the karst areas in the Vogelkop (Bird’s Head), the lowland forest at the southern base of the central mountains, the southern Bomberai Peninsula, and the Tami River valley adjacent to PNG. To preserve biologically critical habitat not currently included in the protected areas system, reserve extensions were proposed in the Tamrau, Arfak, Fakfak, and Foja mountains, and the Mamberamo reserves. Several areas, including the Baliem Valley, Asmat region, Temongsir lowland forests, and Waigeo, Batanta, and Misool islands, were proposed as ‘‘Integrated Biological Management Areas’’ which should be carried out in partnership with local communities to sustainably manage natural resources. There was strong consensus that the management of protected areas in Papua

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was inadequate, existing only as ‘‘paper parks’’ and so in great need of additional staff, legal enforcement, and outreach to communities for support. There was also strong agreement that many areas are only proposed, and so their legal status should be clarified (Supriatna 1999). Other specific recommendations of the Priority-setting Program included: including biodiversity considerations into district- and provincial-level spatial planning, both to preserve species’ requirements and to preserve ecosystem services necessary for sustainable economic development; ensuring that social and environmental assessments are thorough, transparent, and publicly available prior to implementation of oil, gas, mining, agricultural, and other development projects, and critically, ensuring robust community involvement in the decision making processes; greater emphasis in development projects on the socioeconomic and cultural needs and aspirations of local communities; additional biological inventory work and ecological studies; capacity-building training for university students in Papua, and the local establishment of specimen collections and herbaria; enhanced cooperation and collaboration among the Indonesian Institute of Sciences (LIPI), the Directorate of Conservation and Protected Areas (PHKA: Perlindungan Hutan dan Konservasi Alam), Papua’s universities, and international institutions and nongovernmental organizations; enhanced public understanding and awareness of environmental priorities and issues; greater commitment in the donor community to long-term planning horizons (up to ten years) for conservation and development projects; establishing a Papua Conservation Trust Fund to help fund local conservation initiatives. Conservation International’s 1996–1998 Irian Jaya [Papua] Priority-setting Program was successful in establishing a process for productive dialogue and for reaching consensus among various levels of stakeholders, which are crucial for successful implementation of the protected areas system in Papua (Supriatna 1999). It also brought the biodiversity of Papua into the international spotlight, and generated much interest in expanding the documentation and protection of the province’s unique biological and cultural diversity. The Priority-setting Program recommendations also directly facilitated the establishment in 2002 of the Papua Conservation Fund (PCF). Shortcomings of the Priority-setting Program included: incomplete data capture in taxonomic group meetings; lack of consistent criteria used by the crossdisciplinary regional groups; lack of relative ranking of priority areas based on biological importance, degree of threat, or the identification of specific conservation opportunities (e.g., potential local community partners with sufficient local capacity); and incomplete follow-through implementation in integrating the workshop results into government spatial planning (pers. obs.). To be sure, however, the last was particularly challenging because of the task’s complicated political nature and the relative lack of access by conservation practitioners to levels of government at the time (ca 1997–1998).

world wildlife fund’s ecoregional approach in papua Ecoregions are not conservation priority areas. They are a system of classifying biodiversity in a framework that recognizes broad ecological patterns and proc-

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esses, which can be used as a tool for maximizing the representation of land systems and biodiversity in conservation planning. The World Wildlife Fund’s Global 200 list used Ecoregions to identify areas essential for the conservation of irreplaceable and threatened biodiversity (Olsen and Dinerstein 1998; Wikramanayake et al. 2002). WWF’s approach to conservation in Papua includes establishing representative networks of protected areas within each Ecoregion and identifying priority actions to apply WWF’s system of Ecoregion-based management strategies. WWF has identified twelve terrestrial Ecoregions in Papua (see color insert for map): Biak-Numfoor rainforests, Central Range montane rainforests, Central Range subalpine grassland, Yapen Island rainforests, northern New Guinea lowland rain and freshwater swamp forests, northern New Guinea montane rainforests, southern New Guinea freshwater swamp forests, Southern New Guinea lowland rainforests, Vogelkop montane rainforests, Vogelkop-Aru lowland rainforests, Trans-Fly savanna and grasslands, and New Guinea mangroves. With the exception of the rainforests of Biak-Numfoor, Yapen, Vogelkop, and VogelkopAru, these Ecoregions also occur in PNG. WWF is also developing a separate Ecoregion classification for freshwater systems. WWF’s Global 200 Ecoregions are a complementary approach that prioritizes a subset of the most important and threatened global Ecoregions. Global 200 sites in Papua are: southern New Guinea lowland forests, New Guinea montane forests, Trans-Fly savannas, Central Range subalpine grasslands, New Guinea mangroves, New Guinea rivers and streams, and Lake Sentani (with Lake Kutubu in PNG). The WWF program seeks to integrate the Ecoregions approach into provincial and regency level land use planning. In addition, WWF advocates refining and clarifying of park boundaries, identifying conservation and resource management priorities, aligning the activities of conservation organizations with government agencies, identifying common ground in approaches to conservation strategies at island and ecosystem levels, identifying priority stakeholder groups, and establishing and implementing field-level action plans for priority sites (WWF 2002).

birdlife international’s endemic bird areas in papua BirdLife International’s Endemic Bird Areas (EBAs) are well known and used by local land use planning authorities in other parts of Indonesia such as Maluku and Nusa Tenggara Timur (Jepson and Whittaker 2002), but it is unclear whether planning authorities in Papua are aware of the usefulness of the EBAs. Eight Endemic Bird Areas have been identified in Papua (Sujatnika et al. 1995): west Papuan lowlands (Raja Ampat Islands, Vogelkop lowlands, Bird’s Neck area); west Papuan mountains (Tamrau and Arfak mountains of the Vogelkop, Fakfak and Kumawa mountains of the Bomberai, and Wandammen Peninsula); Biak-Numfoor (Biak, Numfoor, and Num islands); north Papuan lowlands (lowland forests from Baropasi and Nabire in the east, to the Mamberamo, Rouffaer [Tariku], Idenburg [Taritatu] rivers, to Jayapura); north Papuan mountains (Foja, Gauttier, and Cyclops mountains); central Papuan ranges (Central Mountain Range, including the Snow [Sudirman] and Star [Jayawijaya] mountain ranges); south Papuan lowlands

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(lowlands and foothills south of the Central Mountains, from Etna Bay through the upriver areas of the Digul River); Trans-Fly (southern lowlands from the Digul to Fly rivers).

Priorities and the Way Forward The increasing intensity of threats to biodiversity in Papua means that there is little time to waste. Papua’s biodiversity is certain to come under increasingly serious threat. Unless field conservation actions, in authentic partnership with local communities and other stakeholders, are undertaken now, it is possible that this biodiversity will disappear. Furthermore, conservation priority setting in Papua cannot progress much further without additional biological and ecological data on which to base further analyses. Nevertheless, the need for more data to better ascertain conservation priorities should not be an excuse for delaying onthe-ground conservation and policy actions. The global significance of Papua’s biodiversity has been amply demonstrated by Conservation International, World Wildlife Fund, and other organizations in several programs to identify conservation priorities (Supriatna 1999; Petocz 1989; Wikramanayake 2000). There is sufficient scientific knowledge to make informed and accurate assessments of what blocks of habitat are worthy of immediate conservation action to conserve species, ecosystems, and ecological processes. While further refinements and modifications to the priority area list may prove necessary, a large number of important priority areas have already been identified. Given the escalating intensity and scope of the threats, the need for immediate and expanded site-level conservation activities is clear. Perhaps the most practical way forward is to prioritize field conservation implementation that incorporates biological survey work and ecological studies (including the establishment of field stations) to fill in information gaps. New data will allow further refinements in conservation priority setting at both the site-level (e.g., a particular patch of forest in the Arfak reserve contains a high density of food plants necessary to the long-term persistence of certain endemic birds) and ecoregion-level (e.g., understanding patterns of species turnover in order to refine the protected area network and add new biodiversity management areas). This research would allow more accurate prioritization of particular areas within the existing reserve system, or, if necessary, extensions or boundary modifications to existing reserves. Although research should certainly not be limited to conservation project areas, site-level conservation often provides key opportunities for biological studies that can be used both to support the site level effort and to refine conservation priority setting. It is not too late to save Papua’s unique biological and cultural heritage, but time is running out. As Supriatna (1999) noted, it is relatively easy to decide on a region’s importance to global biodiversity, but it is much more difficult to decide how to implement a conservation strategy in the region. Field-level implementation may be even more complex and arduous, requiring not only the precise iden-

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tification, monitoring, and assessment of specific conservation targets, but also a continuous process of engagement to ensure the active support of local communities and government partners. How effectively communities are engaged as genuine partners—and incorporating as far as possible their perspectives and aspirations—will dictate success or failure in site-based conservation.

Acknowledgments Thanks to Allen Allison for his comments, to Sara Haney for proofreading this paper, and to John Morrison at WWF–U.S. for providing the Ecoregions map.

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1250 / john burke burnett Wikramanayake, E., E. Dinerstein, C. Loucks, D. Olson, J. Morrison, J. Lamoreux, M. McKnight, and P. Hedao. 2002. Ecoregions in ascendance: reply to Jepson and Whittaker. Conservation Biology 16 (1): 238–243. World Wildlife Fund. 2002. Ecoregion Meeting IV Handout Book. Unpublished meeting agenda. WWF-US, WWF-South Pacific, WWF-PNG, WWF-Indonesia.

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7.3. The Protected Area System in Papua yance de fretes h i s c h a p te r addresses the protected area system in Indonesia, with special focus on Papua, and includes a historical review of the process of establishment and current status of the protected areas of Papua. Also, in this chapter I provide a brief analysis of the legal status, size, habitat, and altitudinal coverage of Papua’s protected areas. Lastly, I provide a brief discussion on the future of protected area management and coverage for Papua. For protected area management constraints in Papua, please refer to Chapter 7.5. The modern protected area system was started in the late 19th century with the establishment of the Hot Spring Reservation and Yellowstone National Park in the United States. The decision to set aside areas of unspoiled land or wilderness was a response to the rapid conversion of the natural land in the western part of North America that led to the demise of forest land and grassland (Miller 1988). But the concept of protected areas can be traced back to the 4th century b.c. in India and about the 3rd century b.c. in Sri Lanka (Chape et al. 2003). It is important to point out that early preservation efforts through the establishment of forest reserves and wildlife sanctuaries were closely linked to local religious beliefs and practices or often for exclusive enjoyment of kings and colonial rulers, and did not always meet biological conservation needs. Increased public understanding of the importance of habitat conservation and the need to protected biological diversity has led governments worldwide to designate terrestrial and marine habitats for protection. In the 1960s about 10,000 protected areas were created that covered some 2 million km2. Today the number of protected areas has increased tenfold, to more than 100,000 protected areas, covering more than 18 million km2 (Chape et al. 2003; see Figure 7.3.1 a,b). Although this is a very encouraging trend for conservation, it is important to note that tropical regions, where majority of biodiversity is found, are less advanced than European and North American regions in creating of new protected areas (Chape et al. 2003). Conservation efforts have long existed in Indonesia, but formal initiatives for creation of protected areas were introduced by the Dutch colonial administration. Setiawan (2001) reports that development of conservation areas was initiated in the period 1863–1919 by Koorders, founder and leader of the Netherlands Nature Conservation Association (Nederland Indische Vereeninging tot Natuurbescherming). Prior to that, in 1714 a protected forest was established in Depok, between Jakarta and Bogor (Setiawan 2001). The first reserve was established in 1889 on Mt Gede Pangrango (West Java) and the first ordinance was issued in 1905 by the Dutch administration (Cribb 1988). During the same period, the forest around Cibodas was declared as a Strict Nature Reserve (Cagar Alam; Setiawan 2001).

T

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

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Figure 7.3.1. Increase in a) number and b) area encompassed by protected areas in the world. Source: IUCN (2003)

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However, conservation-friendly management practices have been long practiced in the country. It can be argued that these practices may not be necessary for conservation purposes, but apply conservation principles. The first comprehensive Indonesian protected area system was developed in the late 1970s and early 1980s. The Directorate General of Forestry (then within the Ministry of Agriculture) with technical assistance and financial support from the Food and Agriculture Organization/United Nations Development Program (FAO/ UNDP), World Wildlife Fund (WWF), and International Union for the Conservation of Nature and Natural Resources (IUCN) formulated national conservation plans for Indonesia. The joint project produced a conservation plan, based on the biogeographic provinces of Indonesia (each of the seven biogeographic provinces presented in a separate volume, with an introductory volume presenting the overall objectives and methodology). Maluku and Irian Jaya (now Papua) were presented in Volume VIII. The 8-volume plan reviewed Indonesia’s biodiversity and proposed a system of protected areas (FAO 1981). The plans also briefly describe the proposed protected areas, reasons for their protection, protection status and priority, and overall scoring (which includes genetic value, socioeconomic justification, and management viability), and threats (FAO 1981).

Protected Areas Systems in Indonesia Several government departments deal with nature conservation and environmental issues in Indonesia: the Department of Forestry (Departemen Kehutanan), the Department of Marine Affairs and Fisheries (DKP: Departemen Kelautan dan Perikanan), and the Ministry for the Environment (KLH: Kementerian Lingkungan Hidup). Only the Department of Forestry has jurisdiction over conservation work in the field, on management of protected areas, both terrestrial and marine. In a recent development, however, the Department of Marine Affairs and Fisheries (DKP) is taking an increasing role in marine conservation. Two main Acts regulate the protected area systems and management: the Conservation of Living Resources and their Ecosystems Act, Law No. 5/1990 (henceforth, Conservation Act) and the Forestry Act, Law No. 41/1999 (Chapter 7.4). According to the Conservation Act, conservation areas have been put under two broad classifications: Natural Protected Areas (Kawasan Suaka Alam) and Nature Conservation Areas (Kawasan Konservasi Alam) and fall into several management regimes as described below. Throughout this chapter, the term of ‘‘conservation area’’ is used interchangeably with ‘‘protected area’’ although the term of ‘‘protected area’’ often refers to Strict Nature Reserve (Cagar Alam), Wildlife Sanctuary (Suaka Margasatwa), and National Park (Taman Nasional). When a specific conservation area (or protected area) is mentioned, the proper protected area category will be used. Appendix 8.7 provides detailed descriptions on the IUCN Protected Area Category (IUCN 1994). Natural Protected Areas (Kawasan Suaka Alam) include Strict Nature Reserves (Cagar Alam), Wildlife Sanctuaries (Suaka Margasatwa), and Biosphere Reserves (Cagar Biospher). Nature Conservation Areas

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(Kawasan Konservasi Alam) include National Parks (Taman Nasional), Grand Forest Parks (Taman Hutan Raya), and Recreational Parks (Taman Wisata). Nature Protected Area (Kawasan Suaka Alam) status is given to areas with special features, either terrestrial or marine, with the goal of preserving and maintaining floral and fauna diversity, natural ecosystems, and life support systems. Nature Conservation Area (Kawasan Konservasi Alam) status is given to areas with specific features, either terrestrial or marine, with the goal of conserving life support systems and floral and fauna diversity, yet allowing for sustainable use of natural resources and ecosystems. Strict Nature Reserve (Cagar Alam) status is given to an area that contains specific flora and fauna, or to particular ecosystems that must be protected and allowed to evolve naturally. Designated to preserve and maintain ecological and natural processes. Level of protection: high/strict (IUCN Category Ia). Wildlife Sanctuary (Suaka Margasatwa) status is given to an area with specific features such as fauna richness or uniqueness that must be managed. Designated with specific conservation goals, such as to protect certain taxa or animal groups. Level of protection: varied from medium to high (IUCN Category Ib). Biosphere Reserve (Cagar Biospher) status is given to an area that contains natural ecosystems, unique ecosystems, or degraded ecosystems to be protected and conserved for research and education. Level of protection: varied from low to medium. National Parks (Taman Nasional) are usually undisturbed areas with outstanding natural values, high potential for recreation, and easy access to visitors. A national park will be managed through varied zoning systems, such as core zone for nature protection. Level of protection: varied depending on zone types, from very minimal to very high/strict (IUCN Category II). Grand Forest Park (Taman Hutan Raya) is a conservation area established for plant and animal collections, either in their natural or human-managed state, with native or exotic species, with the main objective of supporting research, knowledge, education, husbandry, cultural value, tourism, and recreation. Level of protection: low to medium. Recreational Park (Taman Wisata) is usually a natural area with high recreational value, designated primarily for recreational purposes. Level of protection: minimal to moderate (IUCN Category III). Game Reserves are natural or semi-natural areas, managed for specific recreation (hunting) purposes. Level of protection: moderate.

Figure 7.3.2 depicts the number of marine and terrestrial protected areas in Indonesia and Figure 7.3.3 depicts the area included in Indonesian protected areas. Protected Forests (Hutan Lindung) are often cited as protected areas in Indonesia, but they are not officially designated as such. Protected Forests are normally established to protect water catchments and to prevent landslides and erosion; such Protected Forests are managed by the Forestry Bureau (Dinas Kehutanan), under the Ministry of Interior. The large size of Protected Forests and the importance of their functions for local communities suggests that these areas should be managed

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Figure 7.3.2. The total number of protected areas in Indonesia. and protected in the same way as formal Natural Protected Areas (Kawasan Suaka Alam). Therefore, Protected Forests (Hutan Lindung) should be formally designated as Natural Protected Areas and placed under the management of the Department of Forestry or its relevant representatives in the region, such as the Natural Resources Conservation Bureau (BKSDA: Balai Konservasi Sumber Daya Alam) or the National Park Bureau (BTN: Balai Taman Nasional). It is unfortunate that, because of intensive lobbying and economic hardship, the government has allowed much mining exploration in nine Protected Forests (Hutan Lindung) in Indonesia in recent years (including in two Protected Forests in Papua). Within the Department of Forestry, the jurisdiction of Conservation and Protected Areas issues falls under the Directorate General of Conservation and Protected Areas (Dirjen PHKA: Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam). However, it is important to note some aspects of protected area management are not directly under the Dirjen PHKA. For instance, mapping and boundary delineation of protected areas is under the Directorate General of Forest Inventory and Mapping. This arrangement often creates difficulties in the management of protected areas in Indonesia. Prior to the enactment of Law No. 20/1999, the Department of Forestry established its representative in the province through the Papua Provincial Forestry Office (Kanwil: Kantor Wilayah Kehutanan Papua), whose main function was to administer the work of the Department of Forestry

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Figure 7.3.3. Percentage protected area coverage (see Table 7.3.1 for detailed numbers). within the province. In addition, Dirjen PHKA maintained two main representatives in the province or region: the Natural Resources Conservation Bureau (BKSDA: Balai Konservasi Sumber Daya Alam), and the National Park Bureau (BTN: Balai Taman Nasional). After the enactment of Law No. 20/1999, all Provincial Forestry Offices (Kantor Wilayah Kehutanan) were abolished, and all Natural Resources Conservation Bureaus (BKSDA) were reassigned to provincial level. National Park Bureaus (Balai Taman Nasional), however, remained unchanged and are present wherever there is a National Park. All other protected areas (e.g., Strict Nature Reserves, Wildlife Sanctuaries) are managed by Natural Resources Conservation Bureaus (BKSDA). Both representatives are often referred as Technical Implementation Units (UPT: Unit Pelaksana Teknis), while all units in Jakarta (containing the ten National Park Bureaus (BTNs) in Java) are referred to as Central Offices (Kantor Pusat). Currently there are 44 National Park Bureaus (BTNs) and 32 Natural Resources Conservation Bureaus (BKSDAs) in Indonesia. These figures may vary as many new provinces

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and districts are created after enactment of Law No. 20/1999. A few Natural Resources Conservation Bureaus (BKSDAs) have been split into new BKSDAs following the creation new provinces. These Natural Resources Conservation Bureaus are responsible for 486 protected areas that cover 23,122,431 hectares, or 12% of the total land and marine area of Indonesia (Table 7.3.1). The amount of habitat in protected areas is actually higher than the figure given in Forest Classification in Indonesia, below, which includes only protected areas on land. Certain human activities are allowed in particular Nature Conservation Areas (Kawasan Konservasi Alam), but almost nothing is permitted in a Strict Nature Reserve (Cagar Alam) or in the ‘‘protected zone’’ of a National Park (Taman Nasional). For research activities within these conservation areas, written permission must be obtained from the Directorate General of Conservation and Protected Areas (Dirjen PHKA), and this permission is issued only at the Ministry of Forestry Office in Jakarta. See Human Activities Permitted and Prohibited in Protected Areas of Indonesia, next page.

Table 7.3.1. Conservation areas by management regime in Indonesia, 2002 Nature Conservation Areas (Kawasan Konservasi Alam)

Size (km2)

Habitat

Number

Strict Nature Reserves (Cagar Alam)

Terrestrial Marine Total

173 8 181

27,186 2,116 29,301

Wildlife Sanctuaries (Suaka Margasatwa)

Terrestrial Marine Total

53 3 56

35,480 62,220 36,132

National Parks (Taman Nasional)

Terrestrial Marine Total

35 6 41

112,918 36,809 149,727

Recreational Park (Taman Wisata Alam)

Terrestrial Marine Total

87 18 105

2,839 7,658 10,496

Grand Forest Park (Taman Hutan Raya)

17

3,343

Game Reserve (Taman Buru)

14

2,224

Total terrestrial conservation areas

379

183,990

Total marine conservation areas

35

47,235

414

231,224

Total conservation areas Source: Dirjen PHKA (2003).

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FOREST CL ASSIFICATION IN INDONESIA According to the Forestry Act (Law No. 41/1999), forest has three main functions: conservation, protection, and production. To ensure forest management will retain these functions, forest was further classified into six categories.

Forest Classifications

Size (ha)

Percentage of total land

Conservation forest and recreation forest (Hutan Suaka Alam and Hutan Wisata)

19,471,000

10.36

Protected forest (Hutan Lindung)

29,961,000

15.95

Limited production forest (Hutan Produksi Terbatas)

25,629,000

13.65

Permanent production forest (Hutan Produksi Tetap)

35,332,000

18.81

Convertible production forest (Hutan Produksi yang dapat dikonversi)

22,735,000

12.11

Forest for other usage (Areal Penggunaan Lain)

54,656,000

29.1

Total land area

187,784,000

100

Source: Departemen Kehutanan (2003).

Protected Areas Management and Development in Papua

brief history and recent developments As in other Indonesian provinces, management of protected areas in Papua follows the standard rules set out for the management of national protected areas. Nevertheless, the enactment of Law No. 20/1999, and especially Law No. 21/2001, which gave Special Autonomy to Papua, may influence the protected area management systems in the future. With Regional Autonomy, provincial and regency (kabupaten) governments are already demanding more involvement in forestry management, which includes protected area management. But until now, there has been no special regulation or serious attempt to reconcile these demands and discrepancies between related Acts (Law No. 20/1999, Law No. 21/2000, Law No. 5/1990 [the Conservation Act], and Law No. 41/1999 [the Forestry Act]). The Department of Forestry (Departemen Kehutanan) held series of workshops and meetings regarding the impact of Law No 20/1999, and outlined a matrix for future forestry planning and management (Departemen Kehutanan dan Perkebunan 2000), but there is no clear legal and political solution to accommodate this proposal. A detailed account of the development of protected areas in Papua is provided

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HUMAN ACTIVITIES PERMITTED AND PROHIBITED IN PROTECTED AREAS OF INDONESIA Type of Protected Area Human Activity

CA

SM

TW

TB

HL

兹

兹

兹

TN(ZI)

TN(ZR) TN(ZP)

Growing food crops Growing tree crops Human settlement Commercial logging Collecting herbs and firewood

兹 兹

兹

兹

兹

兹

Hunting Fishing Camping

*

兹

兹

兹

Scientific collection with permit 兹

兹

兹

兹

兹

Active habitat management

兹

兹

兹

Introduction of non-exotic species

兹

兹

Collection of rattan and poles with permit

兹 *

兹

兹

兹

兹

兹

兹

兹

兹

兹

兹

兹

兹

兹

兹

Mineral exploitation Wildlife control

兹

兹

兹

兹

兹

兹

Visitor use

*

兹

兹

兹

*

兹

Introduction of exotic species Notes: Protected areas: CA  Strict Nature Reserve (Cagar Alam); SM  Wildlife Sanctuary (Suaka Margasatwa); TW Recreational Park (Taman Wisata); TB  Game Reserve (Taman Buru); HL  Protection Forest (Hutan Lindung); TN  National Park (Taman Nasional); ZI  Protected Zone (Zona Inti); ZR  Transition Zone (Zona Rimba); ZP  Use Zone (Zona Pemanfaatan). 兹 indicates permitted activities, * indicates limited activities (permit is required), blank cells indicate prohibited activities. Source: MOF and FAO (1990).

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by Petocz (1989). As in other parts in the country, it started during the colonial administration. In 1919, the Dutch administration declared the Snow Mountain area as the Lorentz Strict Nature Reserve. Additional comprehensive conservation work was initiated in the late 1970s, when the Indonesian government and FAO/ UNDP launched its conservation project (FAO 1981), later refined by IUCN/ WWF (Petocz 1983, 1989). The result of this work was the establishment of a comprehensive protected area system for Papua that designated approximately 20% of Papua’s terrestrial and marine habitats as conservation areas (Figure 7.3.4) under various management regimes. This coverage is by far the highest in Indonesia, and the area is much larger than that for Papua New Guinea (see Protected Area Management in Papua New Guinea, below). Currently, about 54 areas are either designated or proposed as protected areas in the province that have been placed under a variety of management schemes, among which are Strict Nature Reserve (Cagar Alam), National Park (Taman Nasional), and Wildlife Sanctuary (Suaka Margasatwa). Important to note is that total numbers of protected areas listed tend to vary from count to count, and there is also variation in reported management designations. Different authorities list anywhere from 54 to 70 protected areas (Appendix 8.8 provides the list of existing protected areas in Papua). These protected areas are managed by two Natural

Figure 7.3.4. Map of protected areas in Papua.

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PROTECTED AREA MANAGEMENT IN PAPUA NEW GUINEA Because almost all land and natural resources in Papua New Guinea are under the customary control of communities and land owners, the protected area systems are different from those in Papua, where almost all land is under the control of the government. The existing protected area systems were based on the following Acts: The Fauna Protection and Control Act 1966. Sanctuary, Protected Areas (PA), and Wildlife Management Areas (WMA) were voluntarily established by land owners. Thus almost all Sanctuaries, Protected Areas, and Wildlife Management Areas were established in lands owned by traditional communities, managed by land owners with assistance from the Department of Environment and Conservation, PNG (DEC). The Conservation Areas Act 1978. No protected areas were established under this Act. The Act applies to land under customary control and under state control or leased land. The National Park Act 1982. Intended to provide a legal basis for designation of various categories of national park on land under government control, or land under lease. Currently there are about 52 protected areas in PNG, consisting of WMAs, National/Provincial Parks, Nature Reserves, and Wildlife Sanctuaries, the majority of which were established for terrestrial habitats or mixed terrestrial and marine habitats; only one was exclusively for marine habitats. The total land and marine habitats under protection in PNG is 1,642,826 ha or 3.63% of land (compared to Papua, where about 20% terrestrial land is designated as protected areas). The size of individual PNG protected areas varies from 2 ha (mostly memorial parks) to the 590,000 ha of the Tonda Wildlife Management Area. Source: DEC and WWF (1993).

Resources Conservation Bureaus (BKSDAs) and two National Park Bureaus (BTNs). The Natural Resources Conservation Bureau (BKSDA) Papua 1 Office (formerly known as BKSDA Irian Jaya I) located in Jayapura, the Papuan capital, is responsible for eight protected areas located in the north, south, and central mountains of the province. The Natural Resources Conservation Bureau (BKSDA) Papua 2 (formerly BKSDA Irian Jaya II) Office, located in Sorong, is responsible for 19 protected areas found in the Vogelkop region (Manokwari, Sorong, Raja Ampat, and Fakfak). The Cenderawasih Bay National Park Bureau (Balai Taman Nasional Teluk Cenderawasih), located in Manokwari, is responsible for the management of

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Cenderawasih Bay (marine) National Park. Wasur National Park Bureau (Balai Taman Nasional Wasur), located in Merauke, is responsible for the management of Wasur National Park. A National Park Bureau (Balai Taman Nasional) for Lorentz National Park still does not exist, so management of Lorentz National Park is still under the Natural Resources Conservation Bureau (BKSDA) Papua 1.

number and status of protected areas in papua Of the 70 Nature Conservation Areas that currently exist in Papua, only 29 (41%) have been gazetted and have full legal status (Figure 7.3.5). Within the subset that have been gazetted, only a handful are currently under some sort of management in the field, or have a management plan; a few examples are Lorentz National Park, Mt Arfak Strict Nature Reserve, and Wasur National Park. Figure 7.3.5 indicates that the majority of protected areas in Papua, both proposed and gazetted, are classified as Strict Nature Reserves (Cagar Alam), Recreational Parks (Taman Wisata), and Wildlife Sanctuaries (Suaka Margasatwa). Of these three management regimes, gazetting has occurred for 50% of the Strict Nature Reserves, 40% of the Recreational Parks, and 29% of the Wildlife Sanctuaries. Though National Parks (Taman Nasional) form a relatively small subset of the protected areas in Papua, all have been granted official legal status (i.e., been gazetted). Although it may seem promising that there are more Strict Nature Reserves than any other type of protected area and that they form the largest number of gazetted areas, this particular management system also presents greater chal-

Figure 7.3.5. Number of protected areas by management and status category.

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lenges than the others. As indicated previously, the Strict Nature Reserve is designed to offer more substantial protection to forests and biodiversity than other protected areas. However, these Strict Nature Reserves are regularly used by local communities, either as hunting grounds or as pathways, and enforcement of the Strict Nature Reserves’ status is likely to generate conflicts between government and traditional communities. The primary reason that such a small proportion of protected areas have been granted clear legal status is that the convoluted and ambiguous bureaucratic procedures involved in the legalization process usually thwart even the best efforts to gazette these protected areas (World Bank 2001). Limited budgets and staff capacity may also be factors that are inhibiting the legalization of protected areas. However, since many of these protected areas are in remote and sparsely populated locations, where the scale of past economic activities was relatively low, the legalization of the majority of Papua’s protected areas may not have been a priority. Problems concerning the legal status of protected areas will intensify as the national government increases the decentralization of control to the provincial level. Decentralization is spawning political changes in Papua; local governments are attempting to exercise their new control over natural resources (e.g., forests, minerals, and aquatic ecosystems) while local communities are simultaneously attempting to regain their traditional control over the same lands and resources. Despite these power struggles, decentralization offers more substantial opportunities to enhance the management of existing protected areas than was possible under the strict control of a centralized government through the increased commitment and involvement of local stakeholders. Among others, Petocz (1989) and Diamond (1986) have argued that, in the past, the designation and legalization of protected areas in Papua was controlled largely by the central government and failed to engage the participation of all relevant stakeholders. While these arguments may be justified, the protected areas were laid out in accordance with sound biological criteria (FAO/UNDP 1981, Diamond 1986, Petocz 1989, Conservation International 1999). Thus at this point, it is necessary to finish securing the status of the proposed areas (through more democratic processes than were used in the past), and to marshal the efforts of the conservation communities in garnering the support and active participation of local communities and governments in the management of these areas.

size of protected areas in papua Size and placement are two of the most important factors when designing a protected area. The size of a conservation area will determine the number, composition, and population sizes of species within it. Sufficiently large populations of target species must be secured in any conservation area to ensure the populations’ long-term viability; that is, to prevent inbreeding, intra-species competition, and to guard against population collapse during stochastic events such as disease outbreaks and natural disasters. There is no simple or consistent answer to how large

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a conservation area must be. The size of an effective reserve depends largely on such factors as the long-term objectives of the protected area, characteristics of the species needing protection, the management systems that can be adopted at the site, and the socioeconomic conditions surrounding the area. For instance, a relatively small reserve may be sufficient to protect small and highly localized populations of plants, but relatively larger reserves are needed for species with historically wider ranges (e.g., the New Guinea Harpy Eagle, Harpyopsis novaeguineae). Island biogeography theory (MacArthur and Wilson 1967) suggests that the number of species found on an island or isolated habitat (e.g., a protected area surrounded by cropland) depends upon the size of the island and extinction and immigration rates on the island. Some conservationists have used this theory to estimate the ‘‘appropriate size’’ of a given reserve; others have based their estimates upon the minimum viable population (MVP) size of a particular species. MVP is the smallest population size of a species in a given habitat that is needed for that species to survive for a specified length of time (Gilpin and Soule´ 1986; Shaffer 1987; Soule´ 1987). By calculating the MVP of a species, the minimum habitat area (i.e., reserve size) required for its long-term survival can also be determined (assuming that range requirements for the species are known). This approach is most appropriate when the main purpose of a reserve is to protect a single species, but can be made to encompass the needs of several groups by selecting an ‘‘umbrella’’ species at the top of the trophic network (Monk et al. 1998). Umbrella species not only require more habitat area than species at underlying trophic levels, but are dependent on them as well; thus, protecting umbrella species may potentially serve as a proxy for direct protection of species at lower trophic levels. Whether designing reserves based upon the number of species present or the population of these species, both approaches require substantial amounts of data, including the distribution, extinction and immigration rates, natural history, and life history data about the relevant species. Faced with the numerous difficulties in obtaining the accurate and comprehensive data necessary for determining the appropriate size of a protected area, conservationists have been forced to look for more practical methods. Shafer (1990), among others, concluded that reserves should be larger than 2,500–3,000 ha, and suggests that reserves for tropical birds should be larger than 10,000 ha. Diamond (1976) and Terborgh (1976), however, suggest that a minimum of 25,000 ha of natural habitat is required; their calculations stem from a number of field observations and mathematical models designed for bird populations. MacKinnon and MacKinnon (1986) offer an even larger estimate based on the Minimum Critical Size theory, and state that 50,000 ha is necessary for protecting tree species in species-rich lowland rainforest habitat. The mean size of conservation areas in Papua is 162,041 ha, with the median at nearly 300,000 ha. It may seem promising that the average size of conservation areas in Papua is well over the suggested minimum reserve sizes listed above, but this is a false representation. Lorentz National Park, by far the largest protected area in the province at 2,505,600 ha, strongly skews the calculations (Appendix

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8.8). Figure 7.3.6 clarifies the distribution of reserve sizes and indicates that 52% of all existing conservation areas are smaller than the 25,000 ha minimum reserve size suggested by Diamond (1976) and Terborgh (1976). It is interesting to note that nearly all Strict Nature Reserves (72%) are larger than 25,000 ha and that existing National Parks are larger than 150,000 ha. In contrast, approximately 50% of all Wildlife Sanctuaries and 56% of Recreational Parks are smaller than 5,000 ha. Using MacKinnon and MacKinnon’s (1986) minimum suggested size of 50,000 ha, only 39% of existing reserves meet this criterion (Figure 7.3.7). When compared to the size of conservation areas across Indonesia, it becomes clear that protected areas in Papua tend to be larger, and perhaps also more viable, than other Indonesian protected areas. For instance, only 17% of forest reserves in Maluku and Nusa Tenggara meet this minimum viable size (Monk et al. 1998).

protected area coverage and placement Like the size of a protected area, its geographic placement is a critical factor in ensuring the achievement of its long-term objectives, particularly if the conservation area is meant to protect endemics and other species (FAO/UNDP 1981). Due to a paucity of detailed data on species richness and spatial distributions, it is difficult to determine at this time whether Papua’s existing protected areas are appropriately placed to meet individual species’ conservation priorities. However,

Figure 7.3.6. Sizes of current protected areas in Papua using the minimum critical size (25,000 ha) as suggested by Diamond (1976) and Terborgh (1976).

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Figure 7.3.7. Sizes of protected areas in Papua using the minimum critical size (50,000 ha) suggested by MacKinnon and MacKinnon (1986). based on the known distributions of some taxa and piecemeal information on the habits of others, this issue can be initially addressed. On a larger scale, the areas necessary for conserving multiple, related species (i.e., a higher taxon) can also be inferred from the existing data and used to determine the suitability of existing protected areas. Areas critical for the protection of specific taxa (i.e., areas that harbor high species richness, endemics, and threatened species) were identified during the Irian Jaya Biodiversity Conservation Priority-setting Workshop in 1997 led by Conservation International (see The Irian Jaya Biodiversity Conservation Priority Setting Workshop, next page). A spatial gap analysis between existing protected areas (Figure 7.3.4) and priority areas for biodiversity conservation (as identified by Conservation International 1999; Figure 7.3.8) indicated that 50% of the priority conservation areas were outside of the existing protected areas (Figure 7.3.9). Conversely, nearly all of the existing protected areas fall within the biodiversity priority areas. The priority areas for biodiversity conservation are a composite of areas critical for the conservation of certain taxa—birds, plants, and mammals. Gap analyses between the locations of protected areas and the locations of priority areas for specific taxa indicate that only 39% of bird areas, 40% of plant areas, and 43% of mammal areas are covered by existing reserves. More detailed gap analysis of the distribution of Papua endemic rainbow fishes indicated that, although habitat for 16 spe-

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THE IRIAN JAYA BIODIVERSITY CONSERVATION PRIORITY SETTING WORKSHOP In 1997 Conservation International and its partners held the Irian Jaya Biodiversity Conservation Priority Setting Workshop (PSW) to determine priority areas for biological conservation. The workshop included over two years of data compilation and involved about 90 scientists (CI 1999). The workshop participants were organized according to taxonomic groups and were headed by two senior scientists. The task of each group was to determine priority areas for conservation for each respective taxon. Participants at the workshop used a set of biological/ conservation criteria to determine priority areas for conservation of specific taxa. The criteria included: • • • •

Level of biological importance Degree of human pressures and threats Priority for conservation action Priority for research

Later, these taxonomy-based priority areas (maps) were overlaid to produce priority areas for conservation for Papua Province. These ‘‘biological inputs’’ were then combined with information on socioeconomic conditions, existing and proposed development, and capacity of institutions, and used to reach consensus among participants as to which areas should be prioritized and what conservation actions must be taken. The total area recommended as priorities for biological diversity conservation in Papua was 24,770,660 ha. This figure is almost twice the total size of current protected areas in Papua.

cies (of 30 endemic rainbow fishes) are covered under the protected areas, the coverage is only about 27% of their entire habitat. Interestingly, habitat of 14 species are not covered at all under the existing protected areas, these include the habitat for the threatened species, Chilatherina bleheri (Vulnerable). Of the 14 species not under protected area coverage, 11 species have very limited ranges of less 10,000 ha (Ohee 2005). Habitat consideration (i.e., altitude) is another important factor in the spatial placement of reserves. For example, Petocz (1989) points out that with increasing altitude, species richness decreases while the proportion of endemic species increases. In addition, studies of avian altitudinal distribution and seasonal activities in Papua New Guinea (Pruett-Jones and Pruett-Jones 1986) indicate that many species require a variety of habitats and altitudinal ranges to complete their life cycles. The high incidence of endemic species in isolated montane habitats, the high species richness seen at lower elevations, and the dependence of many species on the presence of multiple habitat types means that the entire range of terrestrial altitudes must be represented within protected areas and that adequate amount of each habitat must lie within the reserves. Thus, the proportion of each elevation-

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Figure 7.3.8. Priority areas for biodiversity conservation in Papua as identified during Irian Jaya Biodiversity Conservation Priority-setting Workshop (Conservation International 1997). specific habitat within Papua’s existing protected areas must be determined to assess whether these habitats are adequately represented. Table 7.3.2 indicates that the majority of the protected areas cover lowland habitats, while only eight protected areas incorporate habitats at altitudes above 3,000 m above sea level and only three reserves (Lorentz National Park, Jayawijaya Wildlife Sanctuary, and Enarotali Strict Nature Reserve) contain true alpine habitats. A few of the larger reserves (i.e., Mamberamo-Foja Wildlife Sanctuary, Jayawijaya Wildlife Sanctuary, Mts North and South Tamrau Strict Nature Reserves, Mt Wondiwoi Strict Nature Reserve, and Lorentz National Park) are distributed across a fairly wide range of elevations. At a larger scale, several biogeographic units—composed of sets of habitats— have been identified that span the province and that should also be represented within the protected areas. Petocz (1989) has identified five main biogeographic units on the Papua mainland: northern plains and foothills (including the Cyclops, Foja-Gauttier, and Van Rees mountains); western lowlands and hills (also known as the Vogelkop area); the northwest area (including the Arfak Mts, Tamrau Mts, Wondiwoi Peninsula, and Wandammen Mts); the southwest region (including the Fakfak and Kumawa mts); the Central Cordillera; and southern lowlands. Petocz

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Figure 7.3.9. Gap analysis between priority areas for biological conservation, as identified in the Irian Jaya Biodiversity Conservation Priority-setting Workshop, and existing protected areas in Papua. Results indicate a 50% gap between priority areas for biological conservation and existing Protected Areas. further identifies two biogeographic units covering Papua’s satellite islands: continental (or land bridge) islands (including Waigeo, Batanta, Misool, and Yapen islands); and oceanic islands (including Biak-Supiori and Numfoor Islands). The existing conservation areas cover portions of all the known biogeographic units and a few of the largest reserves (such as Lorentz National Park, Jayawijaya Wildlife Sanctuary, Mamberamo-Foja Wildlife Sanctuary, and Weyland Strict Nature Reserve) span portions of two biogeographic units. Oceanic islands such as Biak and Numfoor—which have the highest number of endemic birds per hectare—contain only two protected areas. Recent forest cover analysis of 39 protected areas (only three of which were marine protected areas) shows that lowland rainforest has the highest coverage (39%) follow by upper montane/alpine forest (11%) and lower montane forest (10%). Figure 7.3.10 shows detailed habitat coverage within the 39 protected areas analyzed, and detailed data are presented in Appendix 8.9 (Forest Watch Indonesia et al. 2001; Conservation International and Forest Watch Indonesia 2003). If the percentage of habitat coverage within the 39 protected areas (Figure 7.3.10 and Appendix 8.8) is compared to the size of Papua’s terrestrial habitats, then the

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50–1134

CA

75

Batanta Barat Island

CA

0–1183

Biak Utara Island

CA

0–695

Misool Selatan Island

CA

0–565

Numfoor Island

CA

100

Pombo Island

CA

sea level

Salawati Utara Island

CA

0–931

Supiori Island

CA

0–1034

Waigeo Barat Island

CA

0–777

Yapen Tengah Island

CA

500–1496

Pantai Mubrani-Kaironi

CA

sea level

Pantai Sausapor

CA

sea level

Pantai Sidei-Wibain

CA

sea level

Pantai Wewe-Koor

CA

sea level

Cyclops/Dafonsoro Mountains

CA

90–2160

Fakfak Mountains

CA

275–1620

Kumawa Mountains

CA

0–1442

Tamrau Selatan Mountains

CA

0–3825

Weyland Mountains

CA

900–3892

Tamrau Utara Mountains

CA

0–3825

Wandammen/Wondiwoi Mountains CA

0–2,222

Rawa Biru

0–90

CA

Kais River

CA

0–150

Bintuni Bay Mangrove

CA

sea level

Arfak Mountains

CA

1500–2941

Bian Lake

SM

50

Inggresau

SM

sea level

Ajoe Islands

SM

sea level

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⬎4,000 m

CA

Kumbe-Merauke

⬎3,000 m

Gunung (Pantai) Wagura-Kote

⬎2,000 m

1750–4000

⬎1,000 m

CA

⬎500 m

Enarotali

⬎100 m

Protected Area

Altitude range Type (m)

sea level

Table 7.3.2. Altitudinal coverage of conservation areas in Papua

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The Protected Area System in Papua / 1271 Asia Islands

SM

sea level

Asia dan Ajoe Islands

SM

sea level

Raja Ampat Islands

SM

sea level

Mamberamo/ Foja Mountains

SM

0–2103

Anggrameos Island

SM

0–205

Mapia Island

SM

sea level

Sabuda Island and Tataruga Island SM

sea level

Pantai Jamursba-Medi

SM

sea level

Jayawijaya Mountains

SM

180–4646

Dolok Bay (Kimaam Island)

SM

150

Rouffaer River

SM

200

Misool Selatan

SML 0–565

Lorentz

TN

0–5031

Wasur

TN

0–90

Teluk Cenderawasih

TNL

sea level

Sayang Island

TW

sea level

Parieri (Biak)

TW

50

Meja Mountain

TWA 50–250

Yotefa Bay

TWA sea level

Note: CA: Strict Nature Reserve (Cagar Alam); SM: Wildlife Sanctuary (Suaka Margasatwa); SML: Marine Wildlife Sanctuary (Suaka Margasatwa Laut); TN: National Park (Taman Nasional); TNL: Marine National Park (Taman Nasional Laut); TW: Recreational Park (Taman Wisata); TWA: Nature Recreation Park (Taman Wisata Alam).

highest percentage habitat coverage is only 7.53%. This is a very interesting result in light of the recent debates on the suggested 10% protected area coverage (Soule´ and Sanjayan 1998, Faith et al. 2001), and will be discussed below.

Discussion As indicated earlier, there has been a significant increase in terrestrial and marine habitat protection worldwide over the past two decades. This has mainly occurred in Europe, Antarctica, and the New World. The increase for countries in the tropical region, where most biodiversity is found, has been relatively smaller. Only about 5% of habitats in rainforest biome are protected (Soule´ and Sanjayan 1998). Yet habitat degradation and loss are continuing at an alarming rate. It is thus important to enlarge the size of protected areas in the tropical regions. We still have the opportunity to do so in the countries or regions termed ‘‘Global Wilderness,’’ in which 75% of natural habitat remains intact, with low human population

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Figure 7.3.10. Percentage of habitat coverage within 39 Protected Areas (20 Strict Nature Reserves, 10 Wildlife Sanctuaries, 6 Recreational Parks, and 3 National Parks), including three Marine Protected Areas. Habitat (forest) classifications follow Whitmore (1990) and Forest Watch Indonesia et al. (2001). density and significant biodiversity (Mittermeier et al. 2003). Unfortunately, this opportunity is unique in human history and failure to capture this opportunity may threaten half of the known biodiversity on earth. Papua, which is classified as one of the Major Tropical Wilderness Areas, could play an essential role in this conservation endeavor. The percentage of terrestrial and marine habitats under protection in Papua is significant, and almost double the 10% target. In fact, the total area under protection in Papua is by far the largest in Indonesia. Yet there is still a significant gap between the existing protected areas and conservation priorities for certain taxa (e.g., rainbow fishes). Future expansion should carefully design reserves or reserve extensions to capture these gaps, especially for freshwater ecosystems, which have traditionally received less attention in Papua and worldwide (Conservation International 2004; Chapter 5.5). Although coverage of protected areas in Papua has already surpassed the 10% target overall, an analysis of major terrestrial habitat representation within Papua’s 39 protected areas indicates that for some major habitat types less than 10% are found in protected areas. Protected area management problems are discussed in Chapter 7.5. It is important, however, to point out that many of the protected area management regimes discussed in this chapter should be adopted or expanded to accommodate local realities. In Papua, as in many other parts of Melanesia, forest

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land is claimed by traditional communities, despite the fact that under national law all land within the country is controlled and managed by the state. Many communities still depend on forests and other natural resources to meet their daily needs. The establishment and management of protected areas should accommodate these needs and yet accomplish critical conservation needs as well. Current management systems prohibit human activities, except in designated zones in national parks and recreational parks. Therefore, we should look for possible means to institute a new management system which allow for low impact and traditional human uses. Such multipurpose parks have been discussed in many meetings, but to date no formal steps have been taken to implement such parks.

Acknowledgments I am indebted to Michelle Brown for comments on the first draft, and Hendi Sumantri for helping with maps and calculations on habitat coverage. Special thanks to Budi Iraningrun (Ira), who was always ‘‘on call’’ for literature research and administrative assistance.

Literature Cited Chape, S., S. Blyth, L. Fish, P. Fox, and M. Spalding. 2003. 2003 United Nations List of Protected Areas. International Union for the Conservation of Nature and Natural Resources and UNEP, Gland. Conservation International. 1999. Lokakarya Penentuan Prioritas Konservasi Keanekaragaman Hayati Irian Jaya. Laporan Akhir [Irian Jaya Biodiversity Priority Setting Workshop, Final Report]. Conservation International, Jakarta. Conservation International. 2004. Conserving Earth’s Living Heritage: A Proposed Framework for Designing Biodiversity Conservation Strategies. Conservation International, Washington, D.C. Conservation International and Forest Watch Indonesia. 2003. Landsat7 ETM Satellite’s land use land cover (LULC) mapping of Mamberamo and Raja Ampat in Papua Province. Final report. Bogor Cribb, R. 1988. The Politics of Environmental Protection in Indonesia. Monash University, Clayton, Australia. Departemen Kehutanan. 2003. Statistik kehutanan Indonesia, 2002 [Indonesian Forestry statistics 2002]. Departemen Kehutanan, Jakarta, http://www.dephut.go.id/ INFORMASI/STATISTIK/Stat2003/Baplan/IV1102.pdf. Departemen Kehutanan dan Perkebunan (Dephutbun). 2000. Laporan: Gugus Tugas Kelembagaan Kehutanan Dalam Rangka Desentralisasi, Hasil Kerja Desember 1999–Februari 2000 [Report: Task Force of Forestry institution in decentralization, December 1999–February 2000]. Departemen Kehutanan dan Perkebunan, Jakarta. Department of Environment and Conservation (DEC) and World Wildlife Fund. 1993. Papua New Guinea: Conservation Areas Strengthening Project 1994–2000. Project document. Diamond, J. 1976. Island biogeography and conservation: strategy and limitations. Science 193: 1027–1029. Diamond, J. 1986. The design of a nature reserve system for Indonesian New Guinea. In

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1274 / yance d e fretes Soule´, M.E. (ed.) Conservation Biology: The Science of Scarcity and Diversity. Sinauer and Associates, Sunderland, Massachusetts. Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam. 2003a. Statistik perlindungan dan konservasi alam Tahun 2002 [Nature protection and conservation statistics 2002]. Departemen Kehutanan, Jakarta. Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam. 2003b. Pusat Informasi Konservasi Alam, Desember 2003 (Map 1:5.000.000), www.dephut.go.id/INFORMASI/ STATISTIK/Stat2003/Baplan/IV1102.pdf. Faith, D.P., C.R. Margules, P.A.Walker, J. Stein, and G. Natera. 2001. Practical application of biodiversity surrogates and percentage targets for conservation in Papua New Guinea. Pacific Conservation Biology 6: 289–303. FAO/UNDP. 1981. National Conservation Plan for Indonesia, Vol. VII: Maluku and Irian Jaya. Field Report of UNDP/FAO National Park Development Project INS/78/061. FAO, Bogor. Forest Watch Indonesia, BAPLAN Departemen Kehutanan dan Perkebunan, and Conservation International. 2001. Updating Papua’s biodiversity habitat information by using remote sensing technology. Final report, Bogor. Gilpin, M.E., and M.E. Soule´. 1986. Minimum populations: processes of species extinction. In Soule´, M.E. (ed.) Conservation Biology: The Science of Scarcity and Diversity. Sinauer Associates, Sunderland, Massachusetts. International Union for the Conservation of Nature and Natural Resources (IUCN).1994. Guidelines for protected areas management categories. International Union for the Conservation of Nature and Natural Resources, Cambridge, UK, and Gland, Switzerland, www.iucn.org/themes/wcpa/. MacArthur, R.H., and E.O. Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, New Jersey. MacKinnon, J., and K. MacKinnon. 1986. Review of the protected area systems in the Indo-Malayan realm. Prepared for the International Union for the Conservation of Nature and Natural Resources/United Nations Environmental Program, England. Miller, G.T. 1988. Living in the Environment: An Introduction to Environment Science. Wadsworth Publishing, California. Ministry of Forestry and Food and Agriculture Organisation. 1990. Indonesian forestry action programme: country brief. Minister of Forestry and the Food and Agriculture Organisation, Jakarta. Mittermeier, R.A., C.G. Mittermeier, T.M. Brooks, J.D. Pilgrim, W.R. Konstant, and G.A.B. da Fonseca. 2003. Wilderness and biodiversity conservation. PNAS 100 (18): 10309–10313. Monk, K., Y. de Fretes, and G. Reksodihardjo-Lilley.1998. The Ecology of Nusa Tenggara and Maluku. Periplus, Singapore. Ohee, H.L. 2005. Pendekatan penilaian status konservasi jenis pada ikan pelangi endemik Papua dan konservasi habitatnya [New approach in the assessment of Papua’s endemic rainbowfishes conservation status and their habitat conservation]. Master’s thesis, Jakarta. Petocz, R. 1983. Recommended reserves for Irian Jaya Province: statements prepared for the formal gazettement of thirty-one conservation area. World Wildlife Fund/ International Union for the Conservation of Nature and Natural Resources, Jayapura. Petocz, R.1989. Conservation and Development in Irian Jaya: A Strategy for Rational Resource Utilization. E.J. Brill, Leiden. Pruett-Jones, S.G., and M.A. Pruett-Jones. 1986. Altitudinal distribution and seasonal activity patterns of birds of paradise. National Geographic Research 2 (1): 87–105.

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The Protected Area System in Papua / 1275 Setiawan, A. unpublished. Sistem kawasan konservasi di Indonesia [Conservation area systems in Indonesia]. Shafer, C.L. 1990. Nature Reserve: Island Theory and Conservation Practice. Smithsonian Institution Press, Washington, D.C. Shaffer, M.L. 1987. Minimum viable populations: coping with uncertainty. In Soule´, M. (ed.) Viable Populations for Conservation. Cambridge University Press, Cambridge. Soule´, M.E. (ed.). 1987. Viable Populations for Conservation. Cambridge University Press, Cambridge. Soule´, M.E., and M.A. Sanjayan. 1998. Conservation targets: do they help? Science 279: 2060–2061. Terborgh, J. 1976. Island biogeography and conservation: strategy and limitations. Science 193: 1029–1030. Whitmore, T.C. 1990. An Introduction to Tropical Rainforests. Clarendon Press, Oxford. World Bank. 2001. Indonesia: environment and natural resources in a time of transition. World Bank, Washington, D.C.

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7.4. Conservation Laws, Regulations, and Legislation in Indonesia, with Special Reference to Papua suer suryadi, agustinus wijayanto, and james b. cannon t r on g d o me s t i c c o n s er v a t io n and natural resource management laws exist that—in theory—govern the use of Indonesia’s forests and wildlife and ensure their survival. In practice, the application of these laws is flawed, and activities such as illegal logging and wildlife trading are widespread. Illegal logging is a major cause of forest degradation and loss in Indonesia. Various estimates indicate that as much as 50% to 88% of Indonesian timber forest products comes from illegally logged trees (Greenpeace 2003; Palmer 2001; Scotland and Ludwig 2002). According to data issued by the Indonesian Forum for the Environment (WALHI: Wahana Lingkungan Hidup Indonesia), the total annual capacity of the country’s pulp mills, sawmills, and plywood factories is 63 million cubic meters, while the total legal timber produced is 12 million cubic meters annually, meaning that approximately 51 million cubic meters of illegal timber are processed annually (Kurniawan 2003a). In the last 50 years, Indonesia has lost 50 million hectares of forest cover, from 162 million ha in 1950 down to 98 million ha in 2000. Illegal logging costs the Indonesian government approximately Rp 30 trillion per year (US$ 3.4 billion) in lost forestry revenues. The value of the damaged and destroyed ecosystem services has not been estimated. Illegal logging has contributed to an increased deforestation rate in Indonesia, which has risen from 1.6 million ha/year in 1998 to 2.4 million ha/year in 2002 (Ministry of Forestry 2003). Papua contains more than 40 million hectares of forest, the largest remaining contiguous forest resource in Indonesia and one of the world’s last remaining tropical environments where the majority of original forests remain (Mittermeier et al. 2002). These globally unique forests are under great threat from illegal logging. The Ministry of Forestry (2003) stated in an official press release that a total of 10 million m3 of timber a year was being smuggled out of Indonesia, with Papua contributing 600,000m3/month (7.2 million m3/year, or 72% of the total). While this is perhaps an overestimate of actual illegal take, it points out the seriousness of the situation. Illegal wildlife trading is also rampant across Indonesia, with charismatic animals such as primates, turtles, and birds particularly at risk. Over one thousand orangutans were reportedly poached from Kalimantan’s forests in 2002, nine thousand turtles were recorded being traded in just four months in 2001, and surveys reveal nearly 50% of the birds found in the marketplace are protected

S

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

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species under Indonesian law (Nursahid 2003). The domestic and international trade in wildlife and wildlife products targets many animal species in Papua, some of which are globally rare, endemic to Papua, or endangered. According to the Directorate General of Conservation and Protected Areas (Dirjen PHKA: Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam), the value of wildlife smuggling is approximately US$ 600 million/year, whereas the Gibbon Foundation estimated that the value is between US$ 547.5 million/year to US$ 1 billion/year (Dursin 2004; Kurniawan 2003b). Wildlife is smuggled to Malaysia, Singapore, China, Hong Kong, and European countries. The unrecorded financial loss may be more that stated above since law enforcement is very weak in the region. Current regulations (e.g., Government Regulation (PP: Peraturan Pemerintah) No. 8/1999) allow trade in unprotected species according to quotas released by the Ministry of Forestry (the management authority) based on recommendations from the Indonesian Institute of Sciences (LIPI; the scientific authority). The hunting and collection of protected wildlife is restricted for research and rehabilitation purposes only and general trading or keeping these species as pets is not allowed. However, particular species listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES) can be traded if they are shown to be the offspring of captive breeding or farming. The environment is only one of the victims of illegal logging and wildlife trade. Illegal logging fuels corruption, political competition, and resource conflicts, thus helping to perpetuate poor governance and worsen the security challenges facing Indonesia. Although strong domestic conservation and natural resource management laws exist, in practice these laws are proving ineffective because of extremely low compliance. Several causes are blamed for these low levels of compliance with natural resource laws, among them: demand for the products domestically and internationally; poverty; insufficient alternative legal livelihoods; disputes over land tenure; the army running businesses to cover its costs; grand and petty corruption; overlapping and contradictory policies and legislation; policies and management regimes that fail to respect community rights; and a general breakdown in the rule of law (Barber and Talbott 2003; Patlis 2002; Poffenberger 1995; Repetto 1988; Sembiring et al. 2003). Success in combating many of these causes may be necessary for a long-term solution, and amelioration of these conditions may be warranted in and of themselves, but it is hard to envisage compliance with resource management laws increasing to tolerable levels without significant parallel improvements in enforcement. This chapter starts by providing an overview of the Indonesian laws governing conservation of biodiversity and management of natural resources, explaining the changes that have been made in recent years. An overview is then given of the illegal activities that most threaten the biodiversity and resources of Papua. The poor performance of enforcement is described. This is followed by a discussion of how to strengthen enforcement and carry out other activities to improve compli-

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ance with the law, such as introducing reform laws to improve clarity, as well as increasing equity and community participation in conservation and natural resource management. The chapter closes with some brief conclusions about how to face the challenge of reforming laws and improving compliance in Papua.

Legislation Sembiring et al. (2003) concluded that no single law specifically defines illegal logging or illegal wildlife trade. Instead there are many different laws governing conservation, forestry, use of biodiversity, and others that outlaw specific types of logging or wildlife hunting, collection, and trade. Among these laws, only a few mention the prison sentences or fines, mostly at the level of Government Laws and Government Regulations (PP: Peraturan Pemerintah). The most recent basic laws governing conservation of biodiversity and management of natural resources in Indonesia are:

law (uu) no. 5/1990: conservation of living resources and their ecosystems The main principle of Law No. 5/1990 is sustainable use of natural resources to support human welfare and quality of life. It regulates preservation and conservation of flora and fauna, ecosystems, conservation areas, and sustainable use of natural resources, and describes the investigation process, penalties, and sanctions for crimes established in the act. However, Law No. 5/1990 required that rules for implementation be issued in the format of Government Regulations (PP: Peraturan Pemerintah). Until February 2001, only 8 out of 13 implementation rules were issued, contributing to a failure to provide the details of law enforcement procedure in the conservation and forestry sectors.

law (uu) no. 16/1992: plant, fishes, and animals quarantine In response to increased wildlife trade among Indonesian provinces and internationally, and to the risk of transferring pests and diseases, the Government of Indonesia promulgated this Law No. 16/1992 to replace the previous Act of Quarantine issued by the Dutch Colonial authorities. Law No. 16/1992 addresses quarantine requirements, defining vectors and pests, actions to be taken, investigation processes, penalties, and fines.

law (uu) no. 23/1997: environmental management This Law No. 23/1997 was issued on September 19, 1997, to replace Law No. 4/1982. Law No. 23/1997 states that environmental management should be implemented by the state with highly responsible, accountable, and sustainable practices to ensure environmentally sound sustainable development for Indonesia. It includes material regulating the participation and role of communities, responsibilities for environmental management, conflict resolution, communities’ and organizations’

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right to sue, and their involvement in enforcement of environmental management law (class action).

law (uu) no. 41/1999: forestry This Law No. 41/1999 was issued to replace Law No. 5/1967 on Basic Forestry. The principle of this later Forestry Act (Law No. 41/1999) was to establish good governance of forestry management by considering and combining utilization and conservation, accounting for the needs of local peoples, clarifying investigation process and penalties and sanctions, and promoting transparency. Law No. 41/1999 shifted operational authority to provincial and regency governments, leaving the central government to address macro strategic issues.

government regulation (pp) no. 45/2004: forest protection Government Regulation No. 45/2004 was an implementation regulation derived from the Forestry Act (Law No. 41/1999) to protect forests from various human activities and forest exploitation. Government Regulation 45/2004 was issued on October 18, 2004, to replace Government Regulation No. 28/1985. Exclusive mandate was given to the Ministry of Forestry to maintain ecological functions. The role and responsibilities of Forest Rangers and Forest Civil Investigators was also emphasized, to increase law enforcement. Communities and the private sector were also deemed to have certain responsibilities for forest protection. The contents of this Government Regulation 45/2004 included implementation of forest protection, protecting the forest from fire, and the penalties and sanctions for contravening the Government Regulation.

government regulation (pp) no. 13/1994: wildlife hunting This Government Regulation No. 13/1994 derives from the Conservation Act (Law No. 5/1990), as an implementation rule to regulate hunting of targeted unprotected wildlife. The regulation defines wildlife hunting, hunting areas, seasons, equipment, licenses, and the rights and obligations of hunters.

government regulation (pp) no. 68/1998: nature sanctuaries and nature conservation areas This Government Regulation No. 68/1998 is derived from the Conservation Act (Law No. 5/1990) and provides technical guidance to manage conservation areas and sanctuaries. Government Regulation No. 68/1998 provides definition of types of conservation and protected areas in Indonesia, their requirements, functions, and management, including preservation and utilization.

government regulation (pp) no. 6/1999: forest utilization in production forests This Government Regulation No. 6/1999 was issued before the Forestry Act (Law No. 41/1999) was approved. Government Regulation No. 6/1999 was issued at the

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time of rapid decentralization and reformation of forest management, and gave authority to Governors and Heads of Regency (Bupatis) to issue licenses for smallscale logging concessions. Traditional communities (masyarakat adat) were given the right to log community-owned forest.

government regulation (pp) no. 7/1999: plant and animal preservation Government Regulation No. 7/1999 derives from the Conservation Act (Law No. 5/1990), defining protected species of flora and fauna and their habitats, and provides rules for implementation, including preservation efforts, conservation institutions, rules on shipping and transporting protected species, and overall control and monitoring. Government Regulation No. 7/1999 requires that control and monitoring be conducted by authorized enforcement agencies using both preventive and suppressive enforcement actions. Preventive actions include, but are not limited to, awareness raising, training staff of law enforcement agencies, and publishing identification guidelines for protected species. Suppressive actions include law enforcement actions to bring suspects into the justice system and to court.

government regulation (pp) no. 8/1999: wildlife utilization This Government Regulation No. 8/1999 derives from the Conservation Act (Law No. 5/1990) and provides rules on how to implement the act with respect to research and development, farming, hunting, trading, exhibition, exchange, medicine plant culture, pet, shipping and transporting wildlife, crime sanctions, and classification and quotas. In addition to relying on the penalties and sanctions as mentioned by Law No. 5/1990, Government Regulation No. 8/1999 also mentions administrative sanctions.

government regulation (pp) no. 34/2002: forest management plan, forest use, and forest area management Government Regulation No. 34/2002, derived from the Forestry Act (Law No. 41/1999) and issued to replace GR 6/1999, governs the use of forests and the procedures for getting permits from the provincial and central governments. Forest was generally grouped into three types: Nature Conservation Area (Kawasan Konservasi Alam), Natural Protected Area (Kawasan Suaka Alam), and Production Forest (Hutan Produksi). Various forest-related activities are regulated under this Government Regulation No. 34/2002, although some activities require further regulation to be issued by the Ministry of Forestry.

Changes and Uncertainties in Legislation There is a considerable lack of clarity between what is legal versus illegal. This ‘‘gray area’’ in Indonesian forest laws and regulations exists because of problems

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with the Indonesian legal framework, poorly drafted laws, and flexible interpretation and implementation (Patlis 2002). Patlis (2002) concluded that even if licensed logging concession holders were to attempt to comply with the laws and implementing agencies were fully capable of enforcing the laws as written, significant conflicts, gaps, and overlaps in the laws would remain, making implementation extremely difficult if not impossible. Significant legal reform to reduce the ‘‘gray area,’’ including clarifying jurisdiction between regional and central government authorities, is seen by many as a necessary precursor to improving compliance and reducing corruption in forestry and forest management in Indonesia. Changes in the laws governing decentralization of forest management in production and protection forests is one of the major causes of uncertainty over the legality of different logging acts. The Ministry of Forestry (MoF) Decree (SK: Surat Keputusan) No. 677/1998 enabled people in traditional communities (masyarakat adat) to get a License to Log Traditional Community Forests (IPKMA: Ijin Pemungutan Kayu Masyarakat Adat). This right was strengthened by Government Regulation (PP) No. 6/1999 regarding Forest Utilization in Production Forest, and Ministry of Forestry Decree (SK: Surat Keputusan) No. 317/1999, which allowed traditional communities (masyarakat adat) to get small-scale logging concession rights in Production Forest or inside existing logging concessions, a License to Log Forests Based on Traditional Community Rights (IHPHHMHA: Ijin Hak Pemungutan Hasil Hutan Masyarakat Hukum Adat). Government Regulation (PP) No. 6/1999, Article 11 states that the Ministry of Forestry may delegate authority to the governor to issue a standard Logging Concession License (HPH: Hak Pengusahaan Hutan) for less than 10,000 hectares, and Article 22 gave authority to the Heads of Regency (Bupati) to issue Licenses to Log Forests (IHPHH: Ijin Hak Pemungutan Hasil Hutan) for 100 hectare forest concessions. Further guidance on issuing permits was provided in Ministry of Forestry Decree (SK) No. 50.1/2000, which gave authority to the Governor and Heads of Regency (Bupati). In order to better govern the large numbers of active locally issued licenses and permits to log and clear forests, the Governor of Papua issued Decree (SK) No. 522.2/2002 to regulate forest product licenses called License to Log Traditional Community Forests (IPKMA: Ijin Pemungutan Kayu Masyarakat Adat), which was identical to the License to Log Forests Based on Traditional Community Rights (IHPHHMHA: Ijin Hak Pemungutan Hasil Hutan Masyarakat Hukum Adat). Up to December 2003, 114 Licenses to Log Traditional Community Forests (IPKMAs) were issued for total area of 111,250 hectares. Misunderstandings occurred because the License to Log Forests Based on Traditional Community Rights (IHPHHMHA) described in Ministry of Forestry Decree (SK) No. 317/1999 specifically stated that products of IHPHHMHAs are for local use, not for commercial trade. Under Law (UU) No.22/1999 regarding Regional Autonomy, regencies developed their own regulations and issued their own permits for logging in Production Forests (Hutan Produksi) and for managing Protection Forests (Hutan Lindung), but in many cases this legislation did not comply with national laws, as was re-

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quired by the Regional Autonomy Act (Patlis 2002). The Forestry Act (UU) Law No. 41/1999 covers further aspects of decentralizing forest management across Indonesia, and the Special Autonomy for Papua Province Act (UU) Law No. 21/2001 adds further details specific to Papua. Government Regulation (PP) No. 34/2002 sought to reassert central government power by decreeing that only the Minister of Forestry can issue logging permits, and disallowing the various kinds of permits developed by regional governments, such as Timber Extraction Licenses (IPK: Ijin Pemungutan Kayu) (Patlis 2002). In Papua, such permits were generally known as Community Cooperatives (Kopermas: Koperasi Peranserta Masyarakat) and land was owned by communities. However, by this time concern had already grown about lack of control and the environmental impacts of the Community Cooperatives (Kopermas). Government Regulation (PP) No. 34/2002 attempted to reduce further the impacts of smallscale logging concessions by making Community Cooperatives (Kopermas) illegal. Ministry of Forestry Decree (SK) No. 50.1/2000 was amended two years later by Ministry of Forestry Decree (SK) No. 541/2002, which prevents the Governor and Heads of Regency (Bupati) from awarding more such licenses, and retroactively withdrew and made invalid all licenses issued after December, 2000. The end result of all these legal changes was a de facto increase in illegal logging, as Community Cooperatives (Kopermas) operations that were possibly legal under local regulations were later determined to be illegal under national law. Considerable controversy continues over what rights regional governments actually have, or should have, over forest resources. Despite Government Regulation (PP) No. 34/2002, regional governments continued to issue logging permits and the Ministry of Forestry struggled to implement and enforce Government Regulation No. 34/2002. Finally, Ministry of Forestry Decree (SK) No. P.07/2005 withdrew Ministry of Forestry Decree (SK) No. 317/1999 to ensure that small logging concessions no longer exist in national government law. To combat illegal logging, the Ministry of Forestry (MoF) and the Ministry of Industry and Trade (MoIT) banned the export of logs on 8 October 2001 (1132/ KPTS-II/2001 and 292/MPP/Kep/10/2001). But in the same year, the Governor of Papua issued a decree to export logs. Recently, the Ministry of Forestry and the Ministry of Industry and Trade also issued export bans for processing wood (sawn timber) (Act 350/Menhut-VI/2004 and 598/MPP/Kep/9/2004, dated 24 September 2004). While management authority over Natural Protected Areas (Kawasan Suaka Alam) and Production Forests (Hutan Produksi) has shifted back and forth from central (national) to regional governments since 1999, the Ministry of Forestry has continuously and consistently maintained legal decision making and management authority over Nature Conservation Areas (Kawasan Konservasi Alam). Law (UU) No. 22/1999, Article 7 and Government Regulation (PP) No. 25/2000, Article 2 retained Ministry of Forestry authority over Conservation Forests (Simorangkir and Sumantri 2002). Therefore logging inside Nature Conservation Areas (Kawa-

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san Konservasi Alam) is arguably the most unambiguous form of illegal logging in Indonesia today. Despite widespread legal confusion, the Indonesian legislation is clear on the illegality of several critical kinds of logging defined as illegal internationally. Cutting in all forest zoned as Nature Conservation Area (Kawasan Konservasi Alam), which includes National Parks (Taman Nasional), Grand Forest Parks (Taman Hutan Raya), Recreational Parks (Taman Wisata), and zoned as Natural Protected Area (Kawasan Suaka Alam), including Strict Nature Reserves (Cagar Alam) and Wildlife Sanctuaries (Suaka Margasatwa), and zoned as various kinds of Game Reserves, is illegal according to the Conservation of Living Resources and their Ecosystems Act (Law (UU) No. 5/1990). Cutting in all forest zoned as Natural Protected Areas (Kawasan Suaka Alam) is illegal according to the Forestry Act (Law (UU) No. 41/1999), which supersedes the original Basic Forestry Act (Law No. 6/1967). Cutting in Production Forest (Hutan Produksi) without concession rights and a license is illegal according to the Forestry Act (Law No. 41/1999). Loading and transporting logs or timber without a Certificate that Logs Were Legally Obtained (SKSHH: Surat Keterangan Sahnya Hasil Hutan) is illegal according to the Forestry Act (Law No. 41/1999). The penalties established in the laws and regulations governing forest conservation and management reflect the scale of the actual or potential environmental impact caused by different infractions. Infractions with the largest actual or potential environmental impact attract the largest maximum penalties (e.g., burning forest; logging inside Natural Protected Areas or Nature Conservation Areas, or near watercourses in Production Forests; and owning or trading protected species; see Tables 7.4.1 and 7.4.2 for lists of infractions and penalties). Infractions that may mask illegal activities that have large environmental impacts (e.g., transport-

Table 7.4.1. Types of crime and maximum penalties for wildlife trade

Type of crime

Legislation

Intentional Jail Fine (years) (Million Rp)

Hunting/cutting protected species

Act 5/1990

5

100

1

50

Transportation of protected species (flora-fauna)

Act 5/1990

5

100

1

50

Trade in protected species (flora-fauna)

Act 5/1990

5

100

1

50

Possession/rearing of protected species (flora-fauna) without a permit

Act 5/1990

5

100

1

50

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Negligence Jail Fine (years) (Million Rp)

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Table 7.4.2. Types of crime and maximum penalties for illegal logging

Type of Crime

Legislation

Intentional Jail Fine (years) (Million Rp)

Negligence Jail Fine (years) (Million Rp)

Logging in protected areas

Act 5/1990

10

200

1

100

Log protected species

Act 5/1990

5

100

1

50

Logging in protection forest

Act 41/1999

10

5,000

Logging in production forest without license

Act 41/1999

10

5,000

Illegal log trading

Act 41/1999

10

5,000

Illegal log transport

Act 41/1999

10

5,000

Illegal log loading

Act 41/1999

10

5,000

ing logs or timber without necessary permits) also attract large penalties. Infractions that undermine the quality of concession management or the sustainability of annual log harvests from concessions, but have low environmental impacts (e.g., violation of annual workplan requirements, or cutting of undersize trees), are not considered criminal acts and attract the smallest penalties. Types of illegal logging that break technical forestry regulations governing logging activities inside concessions are generally considered regulatory violations, and are addressed through administrative rather than criminal sanctions (Patlis 2002). The types of illegal logging and wildlife trade addressed in this report are all considered criminal acts by Indonesian forest law. As such, the enforcement system used to combat forest crimes is the regular Indonesian Code of Criminal Litigation (KUHAP: Kitab Undang-Undang Hukum Acara Pidana), defined in Law (UU) No. 8/1981. The one difference is the involvement of specialist Forest Civil Investigators (FCI), who lead or aid police in investigations of forest crimes because of the extra technical knowledge required to build effective cases for prosecution. The enforcement system under the Code of Criminal Litigation (KUHAP) is described in the Enforcement section, below.

Illegal Activities

illegal logging Indonesian tropical forest degradation destroys ecological and economic resources but also, especially in Papua, destroys traditional cultures, values, and social systems. Illegal logging is now extensive in several protected areas in Indonesia. The Ministry of Forestry (Press Release No. 51/II/PIK-1/2003) made the following observations. First, almost 43 million hectares of forest have been lost of the original

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120.35 million hectares, with forest degradation rates at about 2.1 million ha/year (2000). Second, illegal timber harvesting and illegal timber circulation reach about 50.7 million m3/year, causing financial losses of about Rp 30.42 billion/year. Third, timber smuggling from Papua, East Kalimantan, West Kalimantan, Central Kalimantan, Southeast Sulawesi, Riau, Nanggroe Aceh Darussalam, North Sulawesi, and Jambi reached about 10 million m3/year in 2003. The main destinations of this smuggled timber were Malaysia, China, Vietnam, and India. Fourth, some estimates place illegal logging in Papua as high as 600,000 m3/month, costing the state about Rp 600 billion/month, or Rp 7.2 trillion/year. Further information on the scale of the problem can be gleaned from comparing Indonesian export statistics with the import statistics of various importing countries. For instance, according to the International Tropical Timber Organization (ITTO), quoted by Cenderawasih Pos, 24 August 2002, Indonesian government records showed that in 2000 Indonesia did not export timber at all to Malaysia and only 336,000m3 to China, whereas data from Malaysia showed that Malaysia and China imported from Indonesia 632,000 m3 and 1,385,000 m3 timber, respectively. Commercial-scale illegal logging in Papua comes in various forms and is carried out by a variety of actors. The illegal activity of greatest concern occurs in areas where logging is not permitted (e.g., protected areas) and is linked to companies with licenses to log areas nearby. Those companies may carry out the logging themselves, subcontract to smaller local companies, or simply buy logs without obtaining appropriate papers to demonstrate legality. Logging by Community Cooperatives (Kopermas) is not treated as a forest crime in Papua because of the extensive legal uncertainty described in the previous section. No Kopermas cases were prosecuted in Papua, even after Kopermas were made illegal in 2002, and no cases were prosecuted for contravening national environmental laws, although Community Cooperative logging licenses were issued for areas official designated as Protection Forests. Cases related to Kopermas did not focus on the logging but on the illegal importation of heavy logging machinery into Papua.

illegal wildlife hunting, collecting, and trading Papua is a major source of wildlife trade because it has lots of attractive endemic and unique species such as birds of paradise, parrots, palm cockatoos, cassowaries, tree kangaroos, the Pig-nosed Turtle, and Boelan’s Python. The uniqueness creates demand for domestic and export purposes, no matter the level of protection of the species. As a result, suppliers find ways to meet the demand, and subsistence farmers and hunters participate in order to earn income to meet their basic needs. Profauna Indonesia (a nongovernmental organization focused on animal protection) reported that approximately 115,000 parrots are trapped each year in the wild in Papua and Maluku, including the highly endangered Palm Cockatoo (Probosciger aterrimus), Western Black-capped Lory (Lorius lory), and Sulphurcrested Cockatoo (Cacatua galerita); see www.profauna.or.id/English/animal-fact .html. The rate of hunting and collection is such that several species are likely to

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become critically threatened without immediate improvement in combating the illegal wildlife trade. Based on market surveys and monitoring conducted in Manokwari and Jayapura by Conservation International Indonesia and the Natural Resources Conservation Bureau (BKSDA: Balai Konservasi Sumber Daya Alam), in Papua in 2003, there is a need to continuously patrol wildlife trading hotspots. In Manokwari, the trading hotspots are Sanggeng, Wosi, and Borobudur markets, and in the vicinity of the offices of the Manokwari Regency and Makassar villages. Other priory locations outside of the town of Manokwari are Prafi and Masni. In Jayapura, critical locations of wildlife trade are the Abepura and Hamadi markets, and markets in Arso, Genyem, Koya/Tami, and Yoka. Wildlife traded in Manokwari in 2003 included protected species such as Western Black-capped Lory, Sulphur-crested Cockatoo, Green Turtle (Chelonia mydas), cassowary (Casuarius spp.), cuscus (Spilocuscus spp.), and Eclectus Parrot (Eclectus roratus), and the dried plumes of birds of paradise (Paradisaea spp.). Species were also found that were not protected, including Rainbow Lorikeet (Trichoglossus haematodus), beetles (Lamprima sp.), butterflies (Delias sp.), Irian Sandalwood (Aquilaria filaria), and orchids (Dendrobium spp.). In Jayapura, the species traded included Sulphur-crested Cockatoo, Western Black-capped Lory, several cassowary species, birds of paradise, Victoria Crowned Pigeon (Goura victoria), Rainbow Lorikeet, several cuscuses, and tree kangaroos (Dendrolagus spp.). In addition, in artifact shops were found the remains of birds of paradise and lories and cassowary feathers (Suryadi et al. 2004). The price of these traded animals varies according to the color of the plumage, the quality of the song (for the bird species), and the age. Prices vary from as low as Rp 500 up to Rp. 2,000,000 (see Tables 7.4.3, 7.4.4). The illegal wildlife trade is very lucrative, and hunters and smugglers have ample economic incentive to participate. The hunters and smugglers involved in the wildlife trade are known from the village level up to town level. They come from many walks of life, including from local communities and transmigration sites as well as from some government departments and enforcement agencies. They function as hunters, buyers, middlemen, and sellers. Not all are operating illegally; there are some officially licensed wildlife traders. Trade routes extend out from Manokwari and Jayapura. Wildlife leaves Papua for Sulawesi, Java, and Sumatra by ship (civil and military) and plane (civil and military). The few transport options out of Papua mean that clamping down on wildlife trafficking along major ship and plane routes could substantially reduce the trade in endangered and threatened species.

Enforcement Between 2001 and 2004 Conservation International (CI) carried out an investigation of the enforcement of conservation and forestry laws in Papua, identified weak spots, and implemented various training and technology transfer efforts to

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Table 7.4.3. Price of wildlife traded in and around Manokwari Price (Rp.) Sale

Location

Species

Buy

Manokwari Harbor

Orchids Lorius lory

5,000 100,000

10,000 200,000

Wosi Market

Lorius lory Cervus timorensis (antler) Cervus timorensis (venison) Chelonia mydas (egg) Casuarius spp Aquilaria fillaria

75,000–100,000 50,000 /kg 12,000 /kg — 100 20,000–200,000

150,000 65,000–70,000 20,000 500 400,000 50,000–500,000

Sanggeng Market

Lorius lory Chelonia mydas (eggs) Chelonia mydas (meat) Cacatua galerita Cervus timorensis

70,000–100,000 — — 200,000 200,000

150,000–200,000 600 10,000 300,000 400,000

Masni District (SP 6–11)

Lorius lory Cacatua galerita Trichoglossus haematodus Cervus timorensis (antler)

50,000–75,000 150,000 25,000–30,000 45,000

150,000 250,000 50,000 65,000

Prafi District

Beetles Butterflies

100–200 100–200

500 500

Area around regency offices

Cuscus sp. Cacatua galerita Eclectus roratus

— —

250,000–300,000 300,000 65,000–75,000

Arowi/Pasirido

Lorius lory Aquilaria fillaria

50,000 10,000–15,000

150,000 25,000–50,000

Borobudur Market

Cacatua galerita Cervus timorensis

300,000 250,000–350,000

450,000 500,000

Arkuki Sanggeng

Lorius lory Cervus timorensis (antler)

100,000 45,000–50,000

200,000 65,000–70,000

Makassar/Wosi

Aquilaria fillaria

15,000–50,000

50,000–100,000

Note: ‘‘Buy’’ price indicates the price that a shop owner paid to a hunter or trader to acquire an item; ‘‘Sale’’ price indicates the price that a shop owner charges for the item in the shop.

improve enforcement. The project started by explaining the purpose and approach to enforcement agencies such as the Natural Resources Conservation Bureau, forestry offices, police and military, prosecutors, and judges. CI then collected data from the field, based on official reports from various enforcement agencies involved in five regencies. Conservation International collaborated with the Natural Resources Conservation Bureau (BKSDA: Balai Konservasi Sumber Daya Alam) in Papua to conduct market surveys and monitoring of the wildlife trade in Manok-

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Table 7.4.4. Price of wildlife traded in Jayapura at Abepura Market, Hamadi, Yoka, and Waena. Sale price indicates the price that a shop owner charges for the item in the shop. Location

Species

Sale Price (Rp.)

Abepura market

Chalcopsitta duivenbodei Lorius lory Trichoglossus haematodus

95,000–100,000 140,000–175,000 20,000–30,000

Hamadi market

Lorius lory

150,000–175,000

Waena

Trichoglossus haematodus

25,000

Yoka

Paradisaea minor Goura victoria Cacatua galerita

1,000,000–2,000,000 2,000,000 200,000

wari and Jayapura. CI also developed and maintained a database to track the progress of illegal logging and wildlife trade cases through the justice system. The project profiled the enforcement system in operation in Papua and how it was performing in combating illegal logging and illegal wildlife trading. Wideranging consultations and further analyses were carried out to identify causes of weak enforcement and to prioritize where to focus on strengthening enforcement. The findings and results are summarized below.

the enforcement system Several institutions are involved in the enforcement system. These institutions and main organizational reporting lines are shown in Table 7.4.5. According to the Indonesian Code of Criminal Litigation (KUHAP), there are four enforcement agencies directly dealing with law enforcement: police investigators, prosecutors, judges, and prisons. Those four institutions are operated at the regency level, province level, and national level. Usually, each institution at the lower level should report or at least coordinate with its counterpart at the higher level. The enforcement agents at lower levels should follow the command of their higher-ranking officers. The roles of individuals and the processes that should be followed are outlined in KUHAP (see below for more details). Several institutions at the regency, provincial, and national level are directly or indirectly involved in governing forests or influencing forest governance. These are: supervisors (e.g., Ministry of Justice, Supreme Court, Attorney General, National Police, Indonesian Armed Forces), policy makers in forestry sectors (e.g., Ministry of Forestry, Natural Resources Conservation Bureau, local forestry departments), trade and industry institutions (e.g., Ministry of Industry and Trade, Customs), transportation institutions (e.g., Ministry of Transportation, Port Administration), budget and policy institutions (e.g., parliament, governor, regency heads), and institutions coordinating and directing technical matters at the national level (e.g., Coordinating Ministry of Politics and Security).

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Table 7.4.5. Enforcement institutions Institution

Reports to

Role

National Coordinating Ministry President of Politics and Security Ministry of Forestry

Coordinates ministries combating illegal logging and trade

President

Secures the forest by deploying rangers Enforces operations and actions of rangers and other staff Terminates logging licenses of law breakers Collaborates with other agencies in criminal cases

National Police

President

Implements strict administrative and criminal sanctions against police officers who aid loggers, investors, and buyers of illegal timber Assigns police protection and accompaniment for forest officers and forest rangers during patrols Assigns police to combat illegal logging and distribution Takes action against illegal loggers, including investors, directors of illegal logging operations, and buyers of illegal timber Apprehends, arrests, and investigates all persons involved in illegal logging

Attorney General

President

Implements strict administrative and criminal sanctions against those involved in illegal logging and distribution activities Accelerates litigation process of cases submitted by police investigators and the Civil Service Investigation Office (continued)

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Table 7.4.5. (Continued) Institution Armed Forces

Reports to

Role

President

Implements strict administrative and criminal sanctions against military officers involved in illegal logging, distribution and smuggling activities Indonesian Navy takes strict action to handle timber smuggling from Indonesia Coordinate and improve cooperation between Armed Forces combating illegal logging and distribution

Ministry of Trade and Industry

President

Implements strict administrative and criminal sanctions against officers of the Ministry of Trade and Industry who abuse Ministry’s power to license wood processing Terminates timber extraction license of those who buy illegal or unknown status timber Closes down unlicensed wood processing operations

Ministry of Transportation

President

Implements strict administrative and criminal sanctions against officers within the Ministry of Transportation who are involved in transporting illegal forest products Orders all Port Administrations not to provide service to ships that load illegal timber Takes legal action against any shipping company (and its staff) who transport illegal timber

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Conservation Laws, Regulations, and Legislation in Indonesia / 1291 Ministry of Justice

President

Implements strict administrative and criminal sanctions against judges who abuse their power during the litigation process Provides guidance to judges to apply maximum allowable penalties to defendant The guilty defendant should be punished in jail at Nusakambangan Prison

Local Provincial Forestry Department

Governor

Issues certificates that logs were legally obtained (SKSHH), patrols forests, checks validity of SKSHH, checks log loads, engages expert witnesses, records data on illegal logging

Natural Resources Conservation Bureau (BKSDA)

Ministry of Forestry

Confers CITES permits for the transport of endangered species, controls transport of unprotected wildlife, patrols conservation areas and wildlife markets, conducts investigations, gathers data

Provincial Police

National Police

Supports or conducts investigations

Provincial Attorney

Attorney General

Supports and conducts prosecutions

Provincial Court

Supreme Court

Receives cases, conducts trials, renders fair verdicts

Regency Forestry Department

Head of Regency (Bupati)

Issues certificates that logs were legally obtained (SKSHH), patrols forests, checks validity of SKSHH, checks log loads, engages expert witnesses, records data on illegal logging

Regency Police

Provincial and National Supports or conducts Police investigations

Regency Attorney

Provincial Attorney and Receives cases, conducts Attorney General prosecutions and additional investigations (continued)

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Table 7.4.5. (Continued) Institution

Reports to

Role

Regency Court

Provincial and Supreme Receives cases, conducts trials, Court renders fair verdicts

Port Administration

Province

Customs

Provincial and National Collects customs payments Police

Confers cruising licenses, controls ship liability

There are eleven institutions that influence enforcement performance in combating illegal logging. These are the Coordinating Ministry of Politics and Security, Ministry of Forestry, Indonesian Armed Forces, National Police, Ministry of Transportation, Ministry of Industry and Trade, Customs, Attorney General, courts, and local (province and regency) government. Coordination among these institutions is necessary because each has specific authority to reduce illegal logging. If they do not synchronize their authority, then no matter how serious and expensive their efforts are individually, their efforts may be neutralized by the poor performance of other institutions. Realizing how poorly coordinated these institutions were, the Ministry of Forestry took the initiative to propose Government Regulation in Lieu of Law (PERPU: Peraturan Pemerintah Pengganti Undang-Undang) to combat illegal logging in the forest. The draft defines illegal logging as cutting trees of more than 10 cm in diameter, but also includes the use, transport, storage, ownership, and distribution of this illegal timber. A new special force would be formed, named the National Body to Combat Forest Crime. The body consists of staff representatives from Ministry of Forestry, Indonesian Armed Forces, National Police, Attorney General, and related institutions. The body will have the special right to conduct inquiry, investigation, prosecution, apprehension, arrest, seizure and confiscation of the timber, and selling the impounded evidence through auction. Unfortunately, although this proposed Government Regulation in Lieu of Law (PERPU) is desperately needed, it has not yet been approved by the President’s Office. Despite decentralization, most of the agencies involved in enforcement still report to central government departments. The main exception is production forestry, where the Ministry of Forestry now has no direct reporting line to forestry departments at the provincial and regency levels. Instead, the regency forestry department reports to the Head of Regency (Bupati), with a coordination line to the provincial forestry department. The provincial forestry department reports to the Governor, with a coordination line to the national Ministry of Forestry. The criminal enforcement process has six main steps: inquiry, investigation, prosecution, trial, conviction, and imposition of penalty (see Figure 7.4.1). A case passes through the system, reaching either administrative sanction or the criminal

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Figure 7.4.1. The law enforcement process. FCI: Forest Civil Investigator; FR: Forest Ranger. justice system (see Figure 7.4.2 and Table 7.4.6). The following description of the process is based on the Code of Criminal Litigation (KUHAP), unpublished enforcement agency training manuals, and interviews with enforcement agency staff.

Step 1: Detection The first step in the enforcement process is detection, either by reports of illegal activity by members of the public to the Police or Forest Rangers, or by enforcement agents during forest patrols and inspections of trucks, planes, or ships. Patrols are generally carried out by Forest Rangers and police as part of routine operations, though special patrols by the Navy have been instigated since 2001 in response to the illegal logging crisis.

Step 2: Initial Investigation The purpose of the initial investigation is for investigators to ascertain whether an offense is a crime or not. If a Forest Civil Investigator (FCI) or Regency Police Investigator is part of a patrol that apprehends suspects carrying out an illegal act, then the initial investigation can be done on site and a decision made to arrest the suspect immediately. The investigators fill out an Event Report (Form LK, Laporan Kejadian) reporting on the case and requesting their supervisor to issue an Investigation Warrant (Surat Perintah Penyidikan, known as an SPP, SP2, or SPRINT) to carry out a full investigation (penyidikan). Suspects can be arrested by the police. The procedures the police use to apprehend and arrest suspects for illegal logging and wildlife trading are the same general procedures for enforcement laid out in the Code of Criminal Litigation (KUHAP).

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Figure 7.4.2. The enforcement process and agencies.

Step 3: Full Investigation The purpose of the full investigation (penyidikan) is to gather the evidence necessary for a successful prosecution. Once an Investigation Warrant (SP2) has been issued and an investigation (penyidikan) started, the case cannot be stopped without approval from senior supervisors. Police investigators, with approval from their supervisors, can stop a case by issuing a Letter of Termination of Investigation (Surat Perintah Penghentian Penyidikan, known as an SPPP or SP3) to the prosecutor and suspect explaining why the investigation was terminated, for example, because of insufficient evidence or witnesses, or, in cases of a missing permit, like a Certificate that Logs Were Legally Obtained (SKSHH), the suspect obtains the necessary permits.

Step 4: Police Review If the case does reach the supervisor of the Civil Service Investigation Office (Korwas PPNS: Koordinasi dan Pengawasan Penyidik Pengawai Negeri Sipil), then, depending on the strength of the case, the Korwas PPNS either requires the Forest Civil Investigator to carry out further investigation, or agrees that the case is strong enough and sends it to the prosecutor.

Step 5: Preparing the Case for Prosecution The prosecutor reviews the case, and, depending on the strength of the case, sends it back within 14 days to the forest or police investigators for further investigation (using Form P18 saying the prosecutor requires further investigation, and Form P19 describing the areas to be strengthened), asks investigators in the prosecutor’s office to take over the case and carry out further investigation, accepts the case for

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Table 7.4.6. Enforcement process and roles and responsibilities of enforcement agencies Process

Agencies

Role and Responsibilities

Collect Information

Forest Rangers, Police

Forest Ranger and Police: • Receive information from communities regarding suspicious illegal logging or wildlife trading activity Forest Ranger: • Check location of suspected illegal activity • Apprehend the suspect on location for further investigation • Report the illegal activity to their supervisor and to Forestry Civil Investigator for further action. Police: • Hand over the investigation to Forestry Civil Investigator (FCI) for further action, or check location of suspected activity • Apprehend the suspect on location for further investigation

Sweeping or Operation

Forest Ranger, Police, Military Police

Joint Task Force/Operation: • Conduct sweep in targeted area, such as market, airport, seaport, commercial ship, Navy ship • Check that documents are legal and valid • If the suspect is from the military, Police and Military Police will handle the case • If the suspect is a civilian, Police and Forest Civil Investigator will handle the case • Confiscate evidence, arrest the suspect for further investigation, and compile report • Move evidence to a secure place Routine Operation: • Conduct surveillance in targeted area, such as airport, seaport, Guard Post at protected areas • Confiscate evidence, arrest the suspect for further investigation, and compile report • Move evidence to a secure place (continued)

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Table 7.4.6. (Continued) Process

Agencies

Role and Responsibilities

Investigation (Penyelidikan)

Forest Ranger, Forest Civil Investigator, Police

Forest Ranger: • Check documents regarding harvesting and transporting forest product, verify cargo contents and volume • Search information and evidence of the crimes in forestry sectors. • Arrest suspect on site of crime, and hand the investigation over to Forest Civil Investigator or Police Forest Civil Investigator and Police: • Receive information from Forest Ranger or communities • Search for further information, witnesses, and evidence • Suspend activities, question suspect, check documents and identification • Confiscate evidence, and equipment; arrest suspect if necessary • Move evidence to a secure place

Further Investigation (Penyidikan)

Forest Civil Investigator, Police Investigator

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Forest Civil Investigator: • Check and verify the validity of the report or information on illegal activities • Interview the suspect and/or witnesses about the alleged illegal forestry activities • Search for and confiscate evidence, if that has not been done earlier in the investigation • Get information, legal advice, and supporting evidence from witnesses and technical experts • In coordination with the Police, apprehend and arrest the suspect • Compile and sign off investigation report, and submit report to Police • Suspend or terminate the investigation if evidence is not sufficient for prosecution

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Conservation Laws, Regulations, and Legislation in Indonesia / 1297 Police Investigator: • Receive investigation report from Forest Civil Investigator • Conduct necessary actions on site of illegal activities • Arrest suspects, search for and confiscate evidence, equipment, or documents; check validity of documents • Take fingerprints, photographs, and necessary documents for archiving the case • Invite suspects and witnesses to provide information about the case • Get information from technical experts regarding the case • Terminate the investigation Prosecution

Public Attorney

• Charge the accused • Implement judge’s sentence • Supervise the implementation of probation sentence • Conduct additional investigation (in coordination with investigators) for better accusation document before register and submit to the court • Speed up the process of forest crime which developed by investigators (FCI/Police) • Appeal to the higher court

Trial

Regency Court, Provincial Court, Supreme Court

• Judges receive the case, conduct the trial, and render a judgment based on the argumentation of the prosecution, defense, witnesses, and on available evidence

Sentence

Regency Court Provincial Court, Supreme Court

• Judges gives a jail sentence and fine to the accused person

Execution and Supervision

Attorney and Judge

• Attorney will execute the sentence were given by the judges and Supervisor Judge will supervise the execution • Attorney and defender have the right to appeal to the higher court

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prosecution, or closes the case according to the Code of Civil Litigation (KUHAP), Article 14h. The investigators in the prosecutor’s office can either complete the investigation to the prosecutor’s satisfaction or issue a Letter of Termination of Prosecution (SP3) stopping the prosecution.

Step 6: Trial, Verdict, and Sentencing Once the prosecutor registers the case with the court, the head of the court will assign judges to the case. The court secretary sets a date for the first hearing of the trial. Several hearings may be required to complete the trial, usually a week or so apart in illegal logging and wildlife trading cases (depending on the size of the crime, the complexity of the case, and the availability of witnesses).

enforcement of laws against illegal logging Getting reliable information on enforcement performance is difficult. Cases are often misreported, or not reported at all, and case information is often out of date. As part of the development of a case tracking system, cross-checking with the police, prosecutors, and courts was carried out to obtain an up-to-date status report. In a joint operation in 2001, the Ministry of Forestry and the Indonesian Navy arrested eight ships carrying 26,564 m3 of illegal logs in Papuan waters. This arrest alone provided Rp 63.6 billion of income for the country. In 2002 they arrested five ships carrying 2,500 m3 of processed wood and 11,300 m3 logs, resulting in Rp 447 billion of income. But in the field, collaboration between the Forestry Office and Papua Police was not well coordinated. Illegal logging in Papua has spread to almost all regencies. According to the case tracking database for Papua compiled by Conservation International Indonesia and the Ministry of Forestry, between 2000 and 2004 there were 58 illegal logging cases, of which 12 cases went to court. Of these 12, nine defendants were found guilty, two were found to have violated the law but were released (lepas), and one defendant went free (bebas). Convictions resulted in penalties, including time in jail (8–12 months, on average) and fines (between Rp 500,000 and Rp 30,000,000). Between 2000 and 2002, 40 cases occurred in Papua involving 44,532 m3 and 6,356 logs. And in 2003–2004, 18 cases occurred, involving 68,718 m3 and 14,656 logs. Although large-scale illegal logging is taking place, little evidence was found of enforcement efforts in the forest. This is largely because few Forest Rangers are available to carry out patrols. Instead, most enforcement efforts are targeted at interdicting large shipments carrying logs that are not accompanied by the correct paperwork (either because the logs were illegally cut, or because they were being smuggled out to avoid taxes and fees). Hence, the examples presented here generally concern the enforcement of shipping large quantities of illegal timber out of Papua on large ships. Although these illegal acts could have been considered either crimes or administrative violations, most were treated as criminal cases. We calculated the cost of enforcement (the risk of detection multiplied by the penalty) to illegal loggers and transporters. The probability of detection was ob-

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tained by dividing the quantity of illegal timber detected by the estimated total quantity of illegal logging. The quantity of illegal timber detected was obtained from the case information. The estimated total quantity of illegal logging was based on information from the Provincial Forestry Office (Kantor Wilayah Kehutanan). The results obtained by analyzing the available data are shown in Table 7.4.7. This analysis shows that although the profits to logging in this case are close to US$ 100,000, the value of the disincentive presented by the enforcement regime is under US$ 7 (see Table 7.4.7). When the value of confiscated timber is included the value of the disincentive is nearly US$ 1,000, but this is still far too small to create a significant deterrent (Akella and Cannon 2004). These results from Papua are similar to those seen across Indonesia. Information from the National Police showed that in 2002 there were 442 cases, of which 218 cases were investigated, 170 cases completed by the prosecutor, ready to judge (P21), and 12 cases terminated (SP3). In 2003 there were 546 cases, of which 282 cases were investigated, 268 cases ready to judge (P21), and 5 cases terminated (SP3). And in 2004 there were 463 cases, of which 267 cases were investigated, 174 cases ready to judge (P21), and 5 cases terminated (SP3).

enforcement of laws against illegal wildlife trading There are two types of illegal wildlife trade: first, trading in protected species, which is absolutely illegal; and second, being over quota for unprotected species or species under Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES) Appendix II. In this report, we focus on the first

Table 7.4.7. Logging enforcement performance results Cumulative Probability

Probability

Symbol

Value1

Probability of detection given illegal logging

Pd

0.032

Probability of investigation given detection

Pi兩d

0.68

0.022

Probability of police review given investigation

Pr兩i

0.84

0.018

Probability of prosecution given police review

Pp兩r

0.41

0.007

Probability of conviction given prosecution2

Pc兩p

0.85

Average value of penalty

$1,197

Average time elapsed (days)

t

269

Enforcement disincentive

ED

$6.474

Profits to illegal logging

0.006 3

$91,967.36

Probabilities translate into percentage likelihoods of 3.2%, 68%, 84%, 41%, and 85%. 2Probability of conviction is for cases heard in regency (local) courts. The probability of conviction presented here does not include the results of appeals to higher courts. 3The average value of the penalty and confiscated timber is US$164,706 (Rupiah to US$ exchange rate of 9,383 Rp./$ for the period 2001-2003; source: Pacific Exchange Rate Service at the Sauder School of Business, University of British Columbia). 4The enforcement disincentive including the value of confiscated timber is $890.43. 1

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type of trading which is regulated by Law No. 5/1990 and Government Regulations No. 7/1999 and No. 8/1999. Trading of protected wildlife for commercial purposes is illegal throughout Indonesia, including sale, transport, export, and keeping protected wildlife as pets. The data collected on the wildlife trade revealed that few cases went to court. Of the 160 cases that were investigated (1998–2004), only five went to court, resulting in four convictions and one acquittal. The average jail time was 3–12 months and fines were between Rp 150,000 and Rp 7,000,000. Clearly the economic disincentive imposed by the enforcement system is far smaller than the profits the smuggler can expect to make by participating in the wildlife trade. Most of the illegal wildlife trading cases were terminated without reason or by issuing a simple warning. This weak enforcement occurs because of inadequate equipment and resources, and the challenges of detection due to the relative ease of transporting wildlife. The illegal wildlife trade receives much less attention than illegal logging. There is less political will and financial reason to enforce wildlife laws, and in many cases it is understood that local people are hunting and collecting wildlife as an essential part of their livelihood. While the lack of will to enforce the laws under these circumstances is understandable, the end result of this unsustainable hunting and wildlife collection will be a loss of income and livelihoods. Blanket efforts to strengthen enforcement are not the answer, but no enforcement at all will end in disaster. A carefully balanced and integrated set of actions is required to protect endangered species, make the hunting and collection of more common species sustainable, and ultimately, to improve the livelihood options of local peoples to reduce the pressure to hunt and collect wildlife.

causes of poor enforcement Our qualitative analysis demonstrated that ineffective enforcement in Papua can be largely attributed to the following: overlapping and inconsistent laws governing logging; weak supervision of concession licenses; lack of coordination among agencies and among local, provincial, and central (national) offices of single agencies; inadequate budgetary resources (leading to insufficient staffing, poor equipment, etc.); insufficient numbers of trained forest investigators; lack of agents and controls in airports and harbors to combat smuggling of wildlife; cultural acceptance of keeping wildlife as pets; lack of incentives for effective performance within enforcement agencies; and corruption in the armed forces, enforcement agencies, and government in general. The probability of detecting illegal logging in Papua appears very low, but this assumes the estimates of the total quantity of illegal logging are correct. If they are, then improving detection is the topmost priority. Detection is low because of the lack of Forest Rangers, as noted above, but also because few people are willing to appear as witnesses or to inform enforcement authorities when they detect forest crimes. People are hesitant to come forward because enforcement agencies do not do enough to protect witnesses and informants from reprisal, and because

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people believe the enforcement agencies are corrupt or incompetent and will not use the information properly or effectively. Relatively speaking, investigation, police review, prosecution, and conviction are being done well, although there is still room for improvement. It is important to remember that the performance of other parts of the enforcement system also remains too low to create sufficient deterrence. Hence, it is not appropriate to focus all available resources on improving detection alone. Raising the probability of detection to 100% under these conditions would raise the enforcement disincentive (risk of detection times penalty) only up to roughly $200—larger than the current disincentive, but still an inadequate deterrent. Without larger penalties, confiscation of logs and equipment, and improvements in the other enforcement steps, illegal logging will continue to be lucrative. It is commonly suggested that raising fines can fix poor compliance, because it is thought that if the fine is higher, violators will be deterred from breaking the law. While it is true that higher penalties do present a greater deterrent, the effects of increasing penalty size are diminished by the low probability of actually being penalized. Three findings emerge from this observation. First, weaknesses in the enforcement system need to be fixed if large penalties are to translate into effective deterrents. Second, when the probability of actually being penalized is low, the size of the penalty required to create a deterrent becomes very large (in this case the penalty would need to be raised to many millions of dollars). Third, the required size of the penalty can quickly exceed what is politically, economically, or culturally viable, meaning other enforcement steps must be strengthened for realistically sized penalties to create an effective deterrent.

Recommendations for Strengthening Enforcement In 2003, the Governor of Papua issued Gubernatorial Decree (SK: Surat Keputusan) No. 50/2003 to formally create the Integrated Team to Combat Illegal Logging in Papua. This team consists of about 23 institutions, with the Papua Provincial Forestry Office (Kantor Wilayah Kehutanan Papua) acting as the team leader. Although the team was created, there has been insufficient follow-up action. Nongovernmental agencies have a critical role to play in strengthening enforcement. Conservation International Indonesia took the initiative to organize a meeting of the Papua Integrated Team to Combat Illegal Logging in July 2004. The meeting was supported by the Papua Provincial Forestry Office and the Institute for Civil Strengthening (a Jayapura-based NGO). The significant result from the meeting was a work plan for immediate action. While long-term systematic efforts are needed to strengthen enforcement and improve governance across Papua, urgent efforts are needed to address the problem in the worst-affected places. Refining the laws to clarify definitions of illegal logging are less pressing than defining a short list of critical illegal logging activities that can provide the main focus of any operational response. A working definition would facilitate building political support and political will from national and

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local government and support from related institutions and communities. This approach would encourage rapid action and help generate important momentum especially in order for the Papua Integrated Team to Combat Illegal Logging to really get full support and trust from public. Consumer countries are often also donor countries and should be doing more to help Indonesia and other producer countries improve compliance (e.g., aid for enforcement strengthening, adaptive management and monitoring), but also to encourage preventive measures (e.g., co-management of protected areas, support for alternative sustainable livelihoods). The illegal wildlife trade also requires the same kind of international attention as illegal logging. The illegal wildlife trade is a big threat to Indonesian biodiversity and enforcement must be undertaken to solve the problem. Specific actions that are required in Papua include the following, as described in the sections below.

improve case tracking and monitoring of enforcement performance The absence of a comprehensive case tracking system (CTS) is a fundamental barrier to promoting transparency in the enforcement system, as well as to monitoring and evaluating enforcement performance in order to identify key problem areas. Two case tracking systems should be created, one by the Natural Resources Conservation Bureau (BKSDA) of Papua and the other by the Forestry Office, reflecting the current responsibilities of each agency established by law to manage and protect Nature Conservation Areas and Production Forests, as defined in Law No. 5/1990 and Law No. 41/1999, respectively. The case tracking system used by each agency should be the same, to aid comparisons between the agencies, and ensure that staff moving between agencies are familiar with the case tracking system. Both the Natural Resources Conservation Bureau and the Forestry Office will need to draft Memoranda of Understanding (MoUs) covering data sharing and management arrangements with the police and the regency prosecutor’s offices. These institutions should also seek Memoranda of Understanding with nongovernmental organizations to share the information they obtain, and to provide independent verification of the accuracy of the information in the case tracking system. Training on how to generate and analyze enforcement statistics from case information is needed. Annual analyses of the data in the case tracking system can be used to evaluate the enforcement system, provided that case monitoring and data are accurate and up to date. This case tracking system has recently been launched in the Ministry of Forestry centrally in Jakarta, and installed in the Directorate of Forest Protection that leads enforcement efforts. Training in how to use the case tracking system is underway centrally and in numerous provinces.

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encourage public involvement Lack of trust by nongovernmental organizations and members of the public in the enforcement system undermines public involvement in the formal legal system. The resulting apathy among the public to report suspected illegal activity significantly reduces the ability of enforcement agencies to detect forest crimes. The public needs transparent and accountable information. Greater trust and belief in the fairness and efficacy of the enforcement system will result in greater public involvement in the enforcement system, further strengthening enforcement. When members of the public report suspected illegal activity to Forest Rangers, the enforcement procedures should require the Forest Rangers to fill in an official report and give the complainant a note of acknowledgement that a report has been filed. Such an acknowledgment should contain the number of the report, the date, and basic details of the suspected illegal activity. Reporting this data on complaints will help promote transparency. A supporting mechanism is an NGOrun illegal logging reporting line and crisis center, where members of the public can report suspected illegal activity, which would then be reported to Forest Rangers and police for investigation.

improve interagency cooperation Coordination and communication among enforcement agencies has not been effective. Each agency tends to be suspicious of the others, and ends up working alone to combat illegal logging. Increasing coordination and communication is desperately needed to combat illegal logging at the local and national level. One of the ways to increase coordination and communication between institutions is through related Memoranda of Understanding that lay out a common agenda and work plan to combat illegal logging.

create and strengthen positive incentives for enforcement agents Enforcement agents do have successes in catching illegal loggers and wildlife traders but seldom get any recognition or reward, even when taking great personal risk in the field. Promotions should be based on performance, making the Natural Resources Conservation Bureau (BKSDA), Papua Provincial Forestry Office, and police meritocracies. Annual bonuses, for instance, could be based on number of cases brought forward, and the proportion of those that are successfully prosecuted.

increase and focus training The consultation meeting on forest and conservation law enforcement held by Conservation International, the Natural Resources Conservation Bureau (BKSDA), and the Papua Provincial Forestry Office agreed on training recommendations for capacity building for each enforcement agency and group of staff, including Forest Rangers and Forest Civil Investigators, police, prosecutors, and

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judges. The training recommendations were: improve knowledge of biodiversity, species, and the ecosystem in general; provide refresher courses for Forest Civil Investigators in investigating forest and conservation cases; provide training in forest and conservation law enforcement for police, prosecutors, and judges; provide training in how to manage the case tracking system database monitoring forestry and conservation cases.

reform key enforcement policies and procedures The overlap of national and local forest regulations causes chaos in forest management. Illegal loggers and wildlife traders use the rift to exploit natural resources in Papua. Discrepancies and gaps between the various laws need to be filled to clarify legality. Under Special Autonomy, local government agencies now have the responsibility for enforcing timber cutting and trading. The procedures used by these agencies also need to be clarified and checked for consistency with national laws.

Discussion Strengthening enforcement is a means to an end, not an end in itself. The end goal of improving enforcement is to eliminate illegal activities or to reduce them to tolerable levels—in other words, to improve compliance. Enforcement contributes to that goal by directly suppressing criminal activity and by creating a deterrent effect. Strengthening enforcement is only one of a number of ways of contributing to the end goal of improving compliance. Other ways include preventive measures such as developing alternative legal sources of income, improving public awareness of and support for the laws, reducing opportunities to break the law, reducing the demand for illegal products, reducing the profits of illegal activities relative to legal ones, and reforming the law to change or clarify unjust and unworkable laws.

ways to improve compliance Considerable debate has taken place over the best ways to improve compliance with particular natural resource and environmental laws. The answer will vary according to the type and scale of crime, the market for the product, the identity of the perpetrators, and the reasons for illegal activity. For instance, traditional hunting of endangered and newly protected species by indigenous peoples requires a response different from that for professional wildlife poaching or commercialscale illegal logging by organized criminal gangs. Co-management approaches may often be the most just and efficient approach for the former type of illegal activity, but the assistance of government enforcement agencies is likely required for the latter. Co-management approaches involve local communities in enforcement activities in a way highly tailored to local issues. Whether illegal activities are practiced for cultural reasons or are key to livelihoods, co-management approaches

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often combine enforcement with developing alternative activities that are culturally acceptable. In Papua, the government, donors, and nongovernmental organizations should support reintroducing and improving existing traditional law (e.g., the sassi system in Raja Ampat, that allows and prohibits harvesting different natural resources at different times). The government enforcement agencies need to support these traditional systems, in part by allowing local people to govern small-scale resource exploitation on their own lands for local use. Thus local people will be able to use traditional laws to improve the protection and use of their natural resources. The best way to improve compliance will also vary depending on why enforcement against a particular crime is currently inadequate. Some internal causes, such as inadequate financial or human resources, or poor training, can be relatively easy to rectify. Others—particularly those where the solution may lie outside the control of enforcement agencies—are far harder to rectify. Problematic external causes include lack of support for laws widely viewed as unjust, political uncertainty, lack of political support for stronger enforcement, and ingrained corruption. Compliance is usually best improved by implementing a mix of preventive and enforcement-strengthening measures together. It is generally accepted that isolated efforts to strengthen enforcement are rarely the most effective way forward. However, this actuality does not mean enforcement strengthening can be ignored. A certain level of increased enforcement is necessary to improve compliance in most situations and will be a crucial element, for instance, when combating organized, commercial-scale illegal activities. Nongovernmental organizations and community groups can influence the efficiency and effectiveness of enforcement, particularly when the number of enforcement personnel is very limited, as is the case in Papua. Building trust between enforcement agencies and other groups will become more likely when enforcement agencies become more transparent, and when enforcement personnel engaged in illegal activities are seen to be punished appropriately.

addressing unjust laws and unfair enforcement A key concern about strengthening enforcement in isolation is the risk of enforcing unjust or counterproductive laws. For instance, traditional users may be unfairly criminalized when government forest policy and laws fail to respect the rights and concerns of indigenous peoples or local communities. The answer to unfair or counterproductive laws is legal reform and—depending on the circumstances—compensation for ‘‘regulatory takings’’ by government. The answer is not weak enforcement. In general, enforcement-strengthening efforts should not be applied to disputed laws that are undergoing reform, although such thinking has been exploited by illegal loggers in Papua for instance, who argue they are acting in community interests in order to escape enforcement efforts. Enforcing disputed laws will likely rouse opposition to strengthening enforcement, particularly from essentially pro-reform allies. This opposition would be

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unfortunate, because weak enforcement of other, undisputed laws is often identified as one of the reasons that local and indigenous groups are harmed by illegal activities. One way forward is to focus initial enforcement-strengthening efforts on noncontentious laws that are generally seen as legitimate. Ideally, those laws would be the ones governing crimes that have the largest economic, social, and environmental effect. Strengthening enforcement to combat crimes that are having direct negative effects on local populations could also help build local confidence in and support for law enforcement. Strengthening environmental and natural resource laws that protect the rights and livelihoods of the poor is also a key element of poverty-reduction strategies. In some cases, the laws may not be at issue, but their application is viewed as illegitimate by local stakeholders because the stakeholders are not adequately involved in decision making. In such situations, co-management approaches—for instance, of natural resources or protected areas—may be necessary to build local support. In other cases, the laws may be generally accepted as appropriate and fair, but enforcement may be applied unequally, with the rich and powerful—or the enforcement agency staff themselves—avoiding justice. Again, this circumstance is not a reason for weak enforcement but rather for increased efforts to ensure that laws are enforced more fairly. Unequal application of enforcement may simply indicate that the rich can afford better lawyers. In such cases, the main need may be for functioning legal assistance for those who are unable to afford lawyers themselves. However, unequal application of the law may also be a symptom of corruption.

ways to reduce corruption The different types of corruption—petty or grand, collusive or non-collusive— pose many challenges to the various approaches for improving compliance. Enforcement strengthening itself can help reduce corruption, both directly by detecting it and indirectly because better enforcement makes corruption more expensive and more difficult. Corruption itself undermines all parts of the enforcement system, and efforts to combat corruption must, therefore, be integral to enforcement-strengthening programs. Enforcement strengthening efforts should start by cleaning up the enforcement and licensing agencies themselves. Greater confidence and trust in government agencies is a necessary base for strengthening broader public respect for the law and its institutions. Anti-corruption efforts include supporting nongovernmental organizations (NGOs) or civil society watchdogs, as well as various efforts within the enforcement system, such as introducing appropriate checks and balances, ensuring pay and bonus structures to create appropriate incentives, revising staffing procedures, and making public the enforcement information needed to evaluate performance. Those efforts must include the judiciary. A clean and effective judiciary that hears cases fairly will encourage wellintentioned enforcement agents and prosecutors so they know there is a point to doing their jobs well. A clean and effective judiciary can also lead reform efforts

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within the enforcement system, thereby rooting out and punishing corrupt officials. In extreme cases, high levels of corruption can call into question whether investing in stronger enforcement is really worthwhile until corruption is brought under some semblance of control. While it is likely that corruption will reduce the effectiveness of investments in enforcement strengthening, it is unlikely that the benefits will be nullified completely. In fact, strengthening the rule of law is generally viewed as an essential part of anticorruption efforts. As a special case, the rigorous analyses of enforcement systems of the type described here promote transparency and understanding and are generally considered a useful component of efforts to reduce corruption. Another issue generally linked to corruption is military involvement in illegal activities. The commonly accepted long-term answer is to ‘‘get the soldiers back in the barracks’’ and create a ‘‘modern’’ army, meaning one that is fully funded by central government, is under fully under governmental control, and fulfills only regular military functions. In the short term, the answer may be to seek agreement that some questionable parts of the military’s business operations be legalized. However, the agreement would be conditional on not engaging in any illegal activities such as resource extraction in particular areas, particularly protected areas and indigenous lands. While some parts of the military may often be engaged in illegal activities, either directly or through corrupt practices, the military in general may need to be engaged directly in finding solutions. In many countries, for instance, the cooperation of the military is required to combat illegal logging carried out on a large scale by organized crime, particularly when elements in the military are involved in the illegal logging.

ways to raise public awareness In other cases, both the laws and their implementation may be fair, but public understanding and support for the laws may be low. Under those circumstances, awareness raising and education efforts are a priority. In other cases, a law may be fair in theory, but the absence of other livelihood options may make its application extremely punishing in practice. The answer is not weak enforcement, but additional assistance and support to develop alternative income opportunities, and possibly also legal reform. Raising awareness of the law and the reasons for it, and building support for the rule of law, could be achieved more effectively if a wide range of groups and institutions—not just enforcement agencies—were involved. Communities would more readily absorb the message about the laws and their rationale if it were explained in a nonthreatening way. Community acceptance would also be increased if the laws were seen to be applied more fairly, which could be helped by establishing a hotline to allow communities to report abuses of power anonymously, and protecting informers and whistle blowers. Legal and procedural reform, compensation, legal aid, public outreach, innovative co-management, anticorruption efforts, and support for alternative legal live-

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lihoods can improve compliance directly and can ease the enforcement challenge. Balanced compliance-improvement efforts that include such measures and enforcement strengthening are likely to be more cost-effective than efforts that overemphasize one measure at the expense of investments in the others. Efforts that recognize and address the legitimate concerns of various stakeholders are also likely to be most successful in building and maintaining political will for action. Opposition from elites defending their entrenched interests is often blamed for lack of political will for strengthening enforcement. However, unless laws are seen to be fair and to be applied fairly, then opposition from those supporting community rights and grassroots reform may undermine the political will for stronger enforcement. Building and maintaining the political will for improving compliance will generally require building and maintaining a broad pro-reform alliance.

enforcement: part of the solution For many good reasons, therefore, enforcement-strengthening efforts should not proceed as isolated activities. Rather, their success is augmented if they are carried out together with a package of activities that fall into two general classes: (1) preventive activities that directly improve compliance while reducing the enforcement challenge, and (2) activities designed to ensure that the laws and their enforcement are workable and fair. The most effective package of activities will vary from place to place and should be developed by local experts and stakeholders. In Papua it is critical also that civil and democratic oversight of the enforcement process is clearly established. Thus nongovernmental organizations, political parties, community groups, and other government institutions, not just enforcement agencies, must support enforcement-strengthening efforts. Finally, the government enforcement system must allow for and support traditional laws and enforcement approaches, and work to ensure these systems are consistent with national laws.

Conclusion The laws governing conservation of biodiversity and management of Papua’s natural resources remain in flux as the general national debate over decentralization and the discussions over special autonomy for Papua continue. Much needs to be done to clarify the situation and reform the legal system governing Papua’s living natural resources. Reform of the legal system is a necessary step, but it will not be enough on its own. Enforcement of the law is currently very weak, and significant strengthening is needed before the laws can have their intended effect. Biodiversity conservation is only one reason that such enforcement strengthening is needed. Other reasons include efficient management of these economically valuable resources, protection of indigenous rights, and improving governance overall. Forest resources play a critical role in indigenous and local community liveli-

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hoods. Unsustainable illegal use will ultimately undermine local livelihoods and exacerbate poverty. But for the moment a lot of money is made from illegal logging and in the illegal wildlife trade. In many parts of Papua illegal logging and wildlife trading are significant parts of the wider economy. For the governments in these areas, and for certain levels of government officials across Papua, illegal logging and wildlife trading are a major source of illicit income. Illegal use of living natural resources is therefore a significant cause of overall poor governance in Papua. Efforts to improve governance in Papua overall can be greatly strengthened by clarifying natural resources management and conservation laws, increasing the fairness and transparency of these laws in theory and in application, and strengthening enforcement in the sector. Systematic efforts are needed, but priorities should also be set to combat the most egregious cases that are doing the most damage, and establish some successes to build on. International NGOs, donors, and responsible forest sector companies should support these enforcement priorities, and contribute their respective skills and resources towards joint activities to improve detection, strengthen enforcement, reform laws, and direct assistance to affected communities.

Literature Cited Akella, A.S., and J.B. Cannon. 2004. Strengthening the weakest links: strategies for improving the enforcement of environmental law globally. Center for Conservation and Government report. Conservation International-Center for Conservation and Government, Washington, D.C. Barber, C.V., and K. Talbott. 2003. The chainsaw and the gun: the role of the military in deforesting Indonesian. Pp. 137–166 in Price, S.V. (ed.) War and Tropical Forests: Conservation in Areas of Armed Conflict. The Haworth Press. Dursin, K. 2004. Animal trade thrives amid crackdown. The Jakarta Post, March 5, 2004. Greenpeace. 2003. Partners in Crime. Greenpeace, London. Kurniawan, M.N. 2003a. Illegal logging costs $609M in environmental destruction. The Jakarta Post, January 17, 2003. Kurniawan, M.N. 2003b. Police, military told to curb animal smuggling. The Jakarta Post, March 29, 2003. Ministry of Forestry, Indonesia. 2003. Departemen Kehutanan Koordinasi Dengan Mabes TNI Dalam Pemberantasan Penebangan Liar. Press release Nomor: 51/II/PIK-1/2003. Indonesia Ministry of Forestry, Jakarta. Mittermeier, R., C.G. Mittermeier, P.R. Gil, J. Pilgrim, G. Fonseca, T. Brooks, and W.R. Konstant. 2002. Wilderness: Earth’s Last Wild Places. CEMEX, Mexico City. Nursahid, 2003. Law enforcement on Indonesia’s wildlife. Proceedings of Indonesia’s Wildlife Seminar. Pusat Informasi Lingkungan Indonesia, Jakarta. Palmer, C.E. 2000. The extent and causes of illegal logging: an analysis of a major cause of tropical deforestation in Indonesia. Centre for Social and Economic Research on the Global Environment working paper. University College London, and University of East Anglia, London. Patlis, J.M. 2002. Mapping Indonesia’s forest estate from the lawyer’s perspective: laws, legal fictions, illegal activities, and the gray area. Unpublished report to The World Bank–World Wildlife Fund Alliance

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1310 / s. s u r y a d i, a. wijayanto, and j. b . c a n n o n Poffenberger, M. 1994. Beyond the timber barons: rethinking Indonesian forest policy. Presented paper to OECD Development Center Experts’ Meeting on Trade, Environment and Sustainable Development, December 7–8, 1994. East-West Center, Washington, D.C. Repetto, R. 1988. The Forest for the Trees? Government Policies and the Misuse of Forest Resources. World Resources Institute, Washington, D.C. Scotland, N., and S. Ludwig. 2002. Deforestation, the timber trade and illegal logging. European Community Workshop on Forest Law Enforcement, Governance and Trade, Brussels. Sembiring, S., A. Akbar, B. Nasution, and C. Diah. 2003. Kajian Hukum Mengenai Illegal Logging dan Perdagangan Hidupan Liar. Institusi Hukum Sumber Daya Alam (IHSA) and Conservation International Indonesia, Jakarta. Simorangkir, D., and Sumantri, 2002. A Review of Legal, Regulatory and Institutional Aspects of Forest and Land Fires in Indonesia. International Union for the Conservation of Nature and Natural Resources and World Wildlife Fund Project FireFight South East Asia, Jakarta. Suryadi, S., Agustinus Wijayanto, and M. Wahyudi. 2004. Market Survey/Monitoring Perdagangan Hidupan Liar di Kabupaten Manokwari dan Jayapura, Papua. Enforcement Economic Report No. 5. Conservation International Indonesia and Seksi Konservasi Wilayah I Manokwari-Balai Konservasi Sumber Daya Alam Papua II, Jakarta.

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38. Neurothemis sp. dragonfly. (M. P. Moore)

39. A Papuan water strider. (M. P. Moore)

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40. A Papuan weevil. (M. P. Moore)

41. Mycalesis phidon, a forest-dwelling butterfly. (M. P. Moore)

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42. A terrestrial forest frog of the genus Callulops. (S. J. Richards)

43. Crocodile skink Tribolonotus sp. This unusual lizard is exported from Papua for the international pet trade. (S. J. Richards)

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44. A forest dragon of the genus Hypsilurus, a spectacular and possibly undescribed reptile from Papua’s Mamberamo Basin. (S. J. Richards)

45. The highly venomous small-eyed snake Micropechis ikaheka occurs throughout New Guinea, inhabiting lowland and mid-montane forests. (A. Allison)

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46. A tiny montane forest frog of the genus Albericus—a new species from the Foja Mountains. (S. J. Richards)

47. A new species of microhylid frog of the genus Choerophryne from the Foja Mountains. (S. J. Richards)

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48. The snake Tropidonophis multiscutellatus. (M. P. Moore)

49. Decorated terrestrial display bower of the Golden-fronted Bowerbird Amblyornis flavifrons, endemic to the Foja Mountains. (S. J. Richards)

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50. Golden-fronted Bowerbird Amblyornis flavifrons in display posture with blue fruit. (B. M. Beehler) 51. Papuan Boobook Owl Ninox theomacha, a common forestdwelling owl ranging through New Guinea’s hill forests and mountains. (B. M. Beehler)

52. This Cinnamon-browed Melidectes Melidectes ochromelas is an uncommon forest-dweller of selected mid-montane sites in New Guinea. (B. M. Beehler)

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53. This Wattled Smoky Honeyeater Melipotes sp. nov. was recently discovered in the Foja Mountains. It is a close relative of the more widespread Common Smoky Honeyeater. (S. J. Richards)

54. This Northern Cassowary Casuarius unappendiculatus is a shy but common inhabitant of lowland forests of the northern watershed of Papua. It is an important seed disperser in Papua’s forests. (M. P. Moore)

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55. Berlepsch’s Parotia Parotia berlepschi is a mid-montane endemic to the Foja Mountains. (B. M. Beehler)

56. Victoria Crowned Pigeon Goura victoria. This huge ground pigeon inhabits the lowland forests of the northern watershed of Papua. (B. M. Beehler)

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57. This young Grizzled Tree-Kangaroo Dendrolagus inustus is a widespread lowland forest-dweller on the northern side of Papua. (S. J. Richards)

58. Great Bare-backed Fruitbat Dobsonia moluccensis, an abundant fruit eater of Papua’s forests and important seed disperser. (B. M. Beehler)

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59. Painted Ringtail Pseudochirulus forbesi, a common hill and montane possum. (S. J. Richards)

60. Long-beaked Echidna Zaglossus cf. bartoni, one of Papua’s rarely seen monotromes. This is a species that has been heavily impacted by subsistence hunting with dogs. (S. J. Richards)

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61. A traditional fisherman in the Raja Ampat Islands, where fishing is an important part of the economy. (J. Jeffers)

62. As-Atat village, Asmat. Canoes and watercourses dominate the lifestyle of the people of the Asmat. (J. B. Burnett)

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63. Asmat village scene, 2003. (J. B. Burnett)

64. Asmat woodcarvers of southwestern Papua are among the most famous in the Pacific. (J. B. Burnett)

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65. A young Indonesian researcher, Burhan Tjaturadi, preparing herpetological field vouchers at a the Minarim camp at the base of the Foja Mountains. (S. J. Richards)

66. A Papuan woman of the western mountains in traditional dress. (M. P. Moore)

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67. Sago processing in Asmat, 2003. Sago is an important and ecologically sound food staple in the lowlands of Papua. (J. B. Burnett)

68. A highlands man of the western mountains of Papua, in traditional dress. (M. P. Moore)

69. Operations of the open-pit Freeport mine high in the Puncak Jaya Range. (M. P. Moore)

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70. Logging road in the Bomberai Peninsula, 2002. Industrial-scale selective logging is widespread in Papua, focusing on a range of valuable tropical hardwood species. (J. B. Burnett)

71. Oil palm monoculture. Oil palm is growing in importance in Papua. This is a particular threat to Papua’s remaining alluvial lowland rainforests. (B. M. Beehler)

72. A transmigration settlement near Timika. These government-sponsored settlement schemes sought to reduce population pressures in western Indonesia. (J. B. Burnett)

7.5. Opportunities and Challenges for Doing Conservation in Papua yance de fretes n d on e s i a i s o n e of the most biodiversity-rich countries in the world. Its placement across two biogeographic realms and the archipelagic nature of the country are two of the factors contributing to its high floral and faunal diversity in both marine and terrestrial ecosystems. With the additional consideration of cultural diversity, Indonesia may indeed be the most diverse country on earth. However, Indonesia’s biological diversity is not distributed evenly among its 17,000 islands. Papua Province (formerly known as Irian Jaya), which includes the western half of the island of New Guinea and numerous satellite islands, contains more than half of Indonesia’s biodiversity (Conservation International 1999). Papua also has a high occurrence of species endemism, particularly for plants, with up to 90% being endemic to the province (Myers 1988; Johns 1995; Chapter 3.1). Discoveries of species new to science continue to accumulate as field inventories are conducted (Chapters 3.1–3.6, 4.1–4.6). Recent Rapid Assessment Programs (RAP) led by Conservation International and its partners have contributed to the discovery of many new species (see Rapid Assessment Program, below) and confirmed that the richness of Papua’s biodiversity is poorly known (Mack and Alonso 2000). The rate of these discoveries shows no sign of decreasing, as a recent field visit to the Mamberamo Basin produced 15 species new to science from 24 frogs collected (S. Richards, pers. comm., 2004). Despite the alarming rate of deforestation worldwide, especially in Indonesia where the annual deforestation rate is estimated at 1.7% annually, approximately 80% of Papua’s primary forests remain intact (World Bank 2001). This is by far the largest remaining tract of pristine forest habitat in Asia and the Pacific, yet it is the very size and wealth of Papua’s forests that make them a compelling target for a range of logging companies from Indonesia and overseas. In turn, the rapid growth of these extractive practices leads to increasing population pressures, changing lifestyles, and government initiatives to boost economic development, which further intensify the threats to these unique forests. Conservation work has long existed in Papua; it started during the colonial administration with the declaration of the Lorentz (Snow) Mountains as a protected area in the 1919. During that time, many regulations were created to protect certain species. The most significant conservation work in Indonesia was initiated shortly after the United Nations ‘‘Conference on the Human Environment’’ in Stockholm in 1972. It began with the establishment of a Ministry of State Development Supervision and the Environment in 1978, which later became Ministry for the Environment in 1983. During that period, the Indonesian government, to-

I

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

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RAPID ASSESSMENT PROGRAM (RAP): PROCESSES AND PROGRESS Conservation International’s Rapid Assessment Program was created in 1990 to quickly provide biological information needed to catalyze conservation action and improve biodiversity protection. RAP assembles teams of host-country and international scientists to produce rapid, first-cut assessments of the biological value of poorly known areas that are not only potentially important biodiversity conservation sites, but also threatened by habitat conversion. The combined knowledge of these experts allows them to assess in three- to four-week surveys the uniqueness and conservation value of an area and to make recommendations about its management. This information is made quickly available to local decision makers through Internet reports, press releases, media coverage, and written reports. So far, biological information from 23 RAP expeditions to terrestrial and freshwater ecosystems throughout the world has resulted in the protection of thousands of hectares of tropical forest, including the creation of national parks in Bolivia and Peru, and in the identification of biodiversity priorities in numerous countries. Conservation International has held several RAPs in Indonesia in Togian and Banggai (2000). With close collaborations with Cenderawasih University, State University of Papua, the Indonesian Institute of Sciences (LIPI) and Papua Environmental Foundation (Yali: Yayasan Lingkungan Hidup Papua); several RAPs have been conducted in Papua. These include: Seiwa and Wapoga (Waropen) in April 1998. During this RAP, scientists discovered many species new to science, including two reptiles, two damselflies, three rainbow fishes, five plants (palms), 17 ants, 29 microhylids (frogs), and 36 heteropteras. RAP training in Yongsu Dosoyo (Jayapura) and Mamberamo (Tiri and Furu Rivers) in September 2001. During this RAP, one new species of freshwater fish, was discovered in Yongsu, and 24 new species (6 frogs, and 18 waterbugs) were discovered at Mamberamo. Marine RAP in Raja Ampat Islands (Sorong) from 27 March–10 May 2002. Four species of coral fish and 9 species of coral were discovered. The RAP team recorded 456 species of hard corals, more than half the world’s known corals!

gether with the United Nations Development Program/Food and Agriculture Organization (UNDP/FAO), launched a National Conservation Initiative, which led the production of a series of conservation plans for the whole country. In early 1980, numbers of protected areas were established in Indonesia, including Papua (Petocz 1989; Chapter 7.3). Species conservation, however, has witnessed relatively less progress than habitat conservation (mainly the establishment of protected areas). Many conservation documents or strategies only mention species that have commercial value or that are threatened from domestic consumption, such as

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parrots, birdwing butterflies, crocodiles, birds of paradise, and sea turtles (Food and Agriculture Organization 1981; Petocz 1989; Conservation International 1999). To date, much so-called species conservation still focuses on species inventory (Allen and Renyaan 1998, 2000; Mack and Alonso 2000; Richards and Suryadi 2002). Few attempts have been made to focus on species conservation (Kemp and Burnett 2003; Muskita 2002; Hitipeuw 2003), but in recent government strategies specific species conservation activities have been proposed (BAPPENAS 2003).

Papua: Global Conservation Importance Papua is becoming recognized as an important area for global conservation because of its extraordinary biodiversity, unique ecosystems (e.g., one of only three snowfields in the tropics) and the fact that its natural forest and some marine areas remain in prime condition (MacKenna et al. 2002). Papuan forest cover remains the highest in Indonesia, if not in the Asia Pacific region. In 1990, as a result of a number of conservation strategy exercises, Papua (with neighboring Papua New Guinea) has been recognized as significant for global conservation. In addition to global conservation assessment or strategies, regional conservation strategies were also developed for Papua. Many global and regional conservation strategies were based on mapping the broad-scale distribution of habitats, biological communities, and species, and their conservation or threatened status worldwide, and then delineating areas of conservation priority. In 1995, BirdLife International issued a report (Sujatnika et al. 1995) identifying about 140 Endemic Bird Areas (EBA) worldwide. Of these, about 24 EBAs are located in Indonesia, by far the highest for any country in the world, and 8 EBA sites are located in Papua. In 1997, Conservation International, together with leading government institutions on science, forestry, and planning in Indonesia (including the Indonesian Institute of Sciences, LIPI: Lembaga Ilmu Pengetahuan Indonesia, the Ministry of Forestry, and the National Development Planning Agency, BAPPENAS: Badan Perencanaan Pembangunan Nasional) and in Papua (Cenderawasih University and the Provincial Development Planning Bureau Papua, BAPPEDA: Badan Perencanaan Pembangunan Daerah Irian Jaya) held the Irian Jaya Biodiversity Prioritysetting Workshop (see Chapter 7.3). Important conclusions were that Papua contributes almost half of Indonesia’s biodiversity, and the current protected area system in Papua needs to be doubled to cover adequately all priority areas for biodiversity conservation in Papua (Conservation International 1999). In 2000, the World Wildlife Fund-U.S. issued its global conservation strategy ‘‘Global 200,’’ which identified about 238 important terrestrial ecosystems for protection. Ecoregions were based on the 26 major habitat types and biological communities, in each of the three major ecosystems (terrestrial, freshwater, and marine) across seven major biogeographic realms. Of the Global 200’s most outstanding Ecoregions of terrestrial, freshwater, and marine importance, eight were located in

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Papua (Table 7.5.1). In 2002, Conservation International identified the earth’s Wilderness Areas (Mittermeier et al. 2002), large tracts of intact natural habitat that harbored high species diversity and endemism. The Wilderness Areas were selected because more than 70% of their habitat remained in its original state, 0.5% of the earth’s vascular plant species are known to be endemic to the area, and human population density was less than 5 people per km2 (Mittermeier et al. 2003). New Guinea was identified as one of three Global Tropical Wilderness Areas (the others being Amazonia and the Congo Basin).

Opportunities and Challenges Declines of natural habitats worldwide, especially of tropical rain forests, have threatened countless species with extinction. Papua (along with Papua New Guinea) is among a handful of areas that still have sizeable tropical rainforest tracts (Mittermeier et al. 2003). As previously mentioned, various international organizations, through global analyses, have identified Papua as one of the most important conservation priorities (Sujatnika et al. 1995; World Wildlife Fund 2000; Mittermeier et al. 2003). Recent fieldwork in Papua has confirmed these findings. One significant advance has been the adoption of most components of the global conservation strategies for Papua by the Indonesian Biodiversity Strategy and Action Plan (IBSAP). This serves as a legal document for doing conservation in Indonesia (BAPPENAS 2003), and provides a roadmap for work in Papua. The enactment of Law No. 21/2000 was important for linking conservation to regional development, gave a greater impetus for local government and nongovernmental organizations to participate in the process, and provided more opportunity than ever before in the conservation sphere in Indonesia.

Table 7.5.1. Eight Ecoregions in Papua and neighboring countries Realm/ Major Habitat

Ecoregion

Terrestrial Realm Tropical and Subtropical Moist Broadleaf Forest Tropical and Subtropical Grassland, Savannas and Shrublands Montane Grasslands and Shrublands Mangroves

Southern New Guinea Montane Forest New Guinea Montane Forests Northern Australia and Trans-Fly savannas Central Range Subalpine Grassland New Guinea Mangroves

Freshwater Realm Small rivers and streams Small Lakes

New Guinea Rivers and Streams Lakes Kubutu and Sentani

Marine Realm Bismarck-Solomon Seas Source: WWF (2000).

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Another important issue is human population. Currently Papua’s population stands at about 2.2 million people, with average density of 6 people per km2, and an average growth rate around 3.18% per year (BP3D and BPS Papua 2003). Furthermore, Papua has already designated the largest proportion of natural habitat under conservation of any province in Indonesia. Still, a variety of major issues remain to be resolved. These include the considerable uncertainty about the current legal status of many of Papua’s delineated conservation areas, the implementation of Special Autonomy, and deficiencies of the conservation budget and staff allocation. Despite claims that the initial designation of conservation areas in Papua failed to apply a participatory approach, the existing protected area system shows promise (Chapter 7.3). The challenge now lies in making this system work effectively. As in many parts of Indonesia, Papua’s protected areas face a number of chronic management problems: unclear jurisdiction between the local and central governments and even among government departments, minimal management capacity and fiscal resources, a lack of political and public support, and frequent disputes between the government and traditional communities over land claims. This last issue remains a serious and unresolved issue in Papua and strongly influences the on-the-ground management of its protected areas. Local people often occupy conservation areas in defiance of government regulation in order to reinforce traditional land claims.

political and administrative changes Although the central government recently delegated a substantial amount of authority to the provincial and district governments through decentralization, conservation issues remain under the jurisdiction of the central government in Jakarta (Law No. 20/2000). The allocation of protected forests affects conservation issues and economic processes at a national scale, which may help to explain why the central government has chosen to maintain control over these areas despite the continuing emphasis on decentralization. Unfortunately, this arrangement is prone to management problems that stem from a lack of support from local governments and peoples. Rather than cede full control of Papua’s protected areas to a single level of government, a better arrangement may lie in the cooperative co-management of these areas by central, provincial, and district authorities. Each level of government is best suited to fulfill a particular role, but none is currently capable of handling the complete management process, from design and legalization to funding and on-the-ground management. The central and provincial governments are the most appropriate units for creating national policies, coordinating the planning of conservation areas, and acquiring funding. The district governments are best suited for management in the field. In the current environment of ambiguous and unregulated jurisdictions, district heads are rapidly demoting the status of conservation areas to Production Forests; this trend could escalate dramatically,

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with catastrophic results, if the district governments are given complete jurisdiction over conservation areas. The Special Autonomy for Papua Law No. 21/2001 dictates that conservation and sustainable development fall within the jurisdiction of provincial governments (see paragraph 58); while the national Forestry Act (No. 41/1999) dictates that the management of protected areas is still under the jurisdiction of the central government (i.e., the Department of Forestry; see Chapter 7.4). At this time, it is unclear how the two laws will be reconciled. As outlined in the Special Autonomy Law No. 21/2001, the local parliament must develop a number of provincial- and district-level regulations to handle natural resource management and environment issues within the space of a single year. This situation is widely viewed by the conservation community as an opportunity to influence local regulations; the cost of nonparticipation is too great to ignore. Despite the contradictions in the two Autonomy Laws (Law No. 20/1999 and Law No. 21/2001), both laws delegate a broader range of authority to manage natural resources—including protected areas—to local government agencies. The positive side of this development is that less bureaucracy will be involved in the management process than in previous decades. The negative aspect is that it carries significant risks in the short term. For decades, decisions impacting natural resources were made by the central government, yet now, under Law No. 20/1999 and Law No. 21/1999, authority is transferred to the district level, where there is little experience in balancing development demands with conservation requirements. The provincial and district authorities are burdened with a weighty legacy of unresolved policy conflicts incurred during the Jakarta government’s administration, inadequate and poor-quality information, weak management agency capacity, insufficient appreciation of the importance of biodiversity conservation within the context of socioeconomic development, and poor planning and management coordination among the various layers of government (i.e., district, provincial, and national), the local community, and nongovernmental agencies. Moreover, it is unclear which among the government agencies holds jurisdiction over the management of conservation areas and other forest types. In the past, the Ministry of Forestry held sole responsibility but with new political and governmental arrangements, this responsibility has been moved out to a variety of agencies. In particular, the Papua Provincial Forestry Office departments (which lie within the district governments) have been given greater authority to manage Papua’s forests and to address forestry issues. Unfortunately, these government agencies are also responsible for issuing logging permits. It is clear that without well-defined roles and responsibilities, as well as coordination mechanisms among government institutions, the management and protection of conservation areas will face many difficulties in the Special Autonomy era.

local capacity and budget allocations Effective management of protected areas incorporates many disciplines, including the natural sciences and conservation biology, as well as macro- and microeco-

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nomics, political science, and social anthropology. A protected area staffed entirely by postgraduate biologists or conservationists is not guaranteed to be well managed or protected. Nonetheless, individuals with a basic grasp of biology or other natural sciences and a postgraduate degree (i.e., individuals who have had significant development capacity) are likely to have the ability to navigate the complex issues inherent in the management of protected areas. In Papua and elsewhere in Indonesia, this basic set of skills is prohibitively rare. A report by the Directorate General of Conservation and Protected Areas (Dirjen PHKA 2003) indicates that about 76% of the current staff working at both the Natural Resources Conservation Bureaus (BKSDA: Balai Konservasi Sumber Daya Alam) Papua 1 and Papua 2, and at the two existing National Parks in Papua (Cenderawasih Bay and Wasur National Parks) have only a high school education. And even those with a university degree tend to be trained in nonforestry fields (Figure 7.5.1). Even though Papua has a large number of protected area staff, the ratio of staff to hectares of protected area is not encouraging (Table 7.5.2). This ratio in Indonesia averages one staffer for every 4,000 ha of protected area, but for Papua the ratio is 1:27,000 ha. A recent study on the effectiveness of protected areas in the tropics has shown that the number of park guards per hectare of protected area is directly and positively correlated with the park’s effectiveness in protecting biodiv-

Figure 7.5.1. Formal educational background of staff of BKSDAs and National Park Bureaus in Papua. Source: Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam (2003).

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Table 7.5.2. Ratio of number of staff at Natural Resources Conservation Bureaus (BKSDA) to total area of existing conservation areas (excluding National Parks) Province

Number of Staff

Area vs. Staff (ha/person)

DKI Jakarta

105

233

2

Jogyakarta

83

418

5

Area (ha)

Central Java

137

3,504

26

Bali

107

18,982

177

West Java

363

76,889

212

East Java

250

54,274

217

Lampung

97

35,979

371

Bengkulu

109

41,700

383

North Sulawesi

74

48,765

659

North Sumatra

202

161,475

799

West Sumatra

194

163,957

845

82

83,246

1,015

West Nusa Tenggara

111

125,715

1,133

South Sulawesi

193

267,285

1,385

East Nusa Tenggara

Jambi

183

337,216

1,843

Riau

91

202,530

2,226

Maluku

73

174,453

2,390

South Sumatra

126

461,104

3,660

Central Kalimantan

73

325,559

4,460

Central Sulawesi

75

354,326

4,724

NAD (Aceh)

85

428,500

5,041

West Kalimantan

66

395,543

5,993

Southeast Sulawesi

92

1,654,643

17,985

East Kalimantan

82

1,536,651

18,740

Papua

249

6,633,900

26,642

South Kalimantan

107

na

na

Gorotanlo

na

76,950

na

North Maluku

na

25,965

na

Banten

na

6,254

na

Source: Dirjen PKHA (2003).

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ersity. In fact, the study shows that 3 guards per 100 km2 is the most effective ratio for achieving biodiversity conservation (Bruner et al. 2001). Unfortunately, there are no recent data available for the numbers and educational background of nongovernmental organization staff in Papua. But an assessment from 1997 (Ohee and Wakum 1997) indicates that the majority of NGO staff had similar formal educational backgrounds to the staff of the Natural Resources Conservation Bureau (BKSDA) in Papua, in which only 65% had a high school education. There is no reason to believe that this situation has improved. The recent major change in priority focus by bi- and multilateral agencies and donors in support of poverty alleviation has resulted in a reduction in funding and staffing for the region’s biodiversity programs. A list of 115 NGOs in Papua indicates that many NGOs are working in multisectoral projects, ranging from agriculture development to human rights and gender issues (Anon 2002). Figure 7.5.2 shows that the major issues addressed by NGOs in Papua include improvement of community economic welfare; health care, including HIV prevention; and gender and children’s issues. Only a few NGOs work on conservation and or environmental issues (Appendix 8.10). Of these 115 NGOs in Papua, only one local NGO works exclusively on environmental issues (education and awareness), while a majority NGOs invest only 10–25% of their project components in environmental or conservation aspects. As for their geographic distribution, the majority of NGOs are based in the big towns in Papua: Jayapura (30%), Manokwari (17%), and Sorong (17%; Figure 7.5.3).

Figure 7.5.2. Major activities undertaken by NGOs in Papua. Source: Anon (2002)

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Figure 7.5.3. NGO concentration in Papua. The majority are based in large towns like Jayapura, Manokwari, Merauke, and Sorong Source: Anon (2002).

The role of NGOs in the management and protection of the environment in Papua is important as it has been delineated under the Law No. 21/2001, article 64/3. Nevertheless, as our rudimentary analysis indicates, this role is far from being fulfilled. The present condition is a fundamental challenge that must be sufficiently addressed if conservation efforts are to be sustainable and meaningful in Papua. If we can assume that government budget allocation is equal to government investment, then the central government has made an insufficient investment in conservation in Papua. If Papua contributes up to 50% of Indonesia’s biodiversity, this should be reflected in the budget allocation for conservation. The reality is different. Only a small proportion (6%) of the total national conservation budget supports the Natural Resources Conservation Bureau’s (BKSDA’s) work in Papua (Figure 7.5.4). Of the total budget allocation for field conservation for the 29 Natural Resources Conservation Bureaus (BKSDAs) in Indonesia, two BKSDAs in Papua received a considerable amount. However, if one considers the size of the conservation areas under the jurisdiction of these two BKSDAs, then their budget allocation is the smallest in the country (Table 7.5.3).

Further Challenges

gaps in the protected area system The designation, legalization, and management of protected areas in Papua is an enormous challenge, but one that must be addressed to ensure protection of a

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Opportunities and Challenges for Conservation in Papua / 1321

Figure 7.5.4. Budget allocation for field conservation in Indonesia for 2000. Source: Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam (2003).

significant and sustainable portion of Papua’s unique biota. The preceding analysis indicates that the current protected area system must be expanded to encompass additional priority areas that have been identified as priority locations for biological conservation. These expansions, of course, should be supported with strong science and proper delineation with respect to both resource and human needs. Any new conservation areas should preferably be designated as wildlife sanctuaries, and must be larger than the 25,000 ha minimum size (the majority of existing wildlife sanctuaries are smaller than 25,000 ha; see Chapter 7.3). In addition to expanding the existing protected areas, all proposed conservation areas should be brought under legal status (i.e., formally gazetted). Given the constraints in managing protected areas, adding lands to the network of conservation areas is likely to intensify current management problems. A strategy that may alleviate this problem would be to develop biodiversity-friendly laws to regulate development in the areas already delineated as critical for biodiversity.

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Table 7.5.3. Budget allocation for Natural Resources Conservation Bureaus (BKSDA) in Indonesia, 2002 Budget (x 1000 rupiah)

Budget vs Area (rupiah/ha)

Province

Area (ha)

Banten

6253.5

na

na

North Maluku

25964.53

na

na

Gorotanlo

76950

na

na

South Kalimantan

na

2,250,359

na

DKI Jakarta

232.84

2,528,986

10,861,476

Jogyakarta

417.83

1,485,309

3,554,817

Central Java

3503.9

4,276,001

1,220,355

Bali

18981.8

2,742,578

144,485

East Java

542744,7

45,388

87,434

West Java

76889.04

6,186,091

80,455

Lampung

35979.4

1,840,231

51,147

Bengkulu

41699.68

1,541,749

36,973

North Sulawesi

48764.5

1,795,105

36,812

Jambi

83246.11

2,343,379

28,150

West Nusa Tenggara

125715.48

3,220,220

25,615

North Sumatra

161475.36

3,865,117

23,936

South Sulawesi

267285.22

5,628,295

21,057

Maluku

174453.08

2,965,694

17,000

East Nusa Tenggara

337216.41

5,465,711

16,208

Riau

202529.57

3,018,016

14,902

West Sumatra

163956.93

2,219,063

13,534

South Sumatra

461104

5,333,935

11,568

Central Kalimantan

325559

2,860,332

8,786

Central Sulawesi

354326.12

2,450,553

6,916

NAD (Aceh)

428500

2,444,589

5,705

West Kalimantan

395543

1,793,672

4,535

Southeast Sulawesi

1654642.66

2,631,235

1,590

East Kalimantan

1536650.7

2,184,466

1,422

Papua

6633900.46

4,602,319

694

Source: Dirjen PHKA (2003). Note: Area is total size of conservation areas within the province (excluding National Parks). This table indicates inappropriate budget allocation, in that areas with greater biodiversity and sizeable conservation areas received less funds.

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It would also be useful to create new protected area categories more relevant to local conditions in Papua. In particular, a ‘‘Traditional Community Reserve’’ designation would be most appropriate. This could be designed in ways that promote biodiversity and forest conservation without threatening the livelihoods of local inhabitants.

constraints on conservation field research Without a doubt, Papua has recently attracted much conservation interest and investment from conservation organizations, international donors, and the private sector. Biodiversity field surveys may be the reason for this renewed attention. For instance, based on the results of the Rapid Assessment Plan (RAP) of Wapoga (Mack and Alonso 2000), a major multilateral agency has dedicated Papua as important part of its biodiversity program in Indonesia. The marine RAP in Raja Ampat (McKenna et al. 2002) has also generated much national and international interest for Raja Ampat’s marine biodiversity and natural beauty. The findings have attracted many tourists and divers from Europe and the United States. However, field research in Papua is a never-ending struggle for permits and clearances from authorities. The government makes doing research very difficult. The need for additional focused research grows by leaps and bounds, but the barriers to conducting this research seem to grow as well. As indicated before, we hardly know a thing about Papua’s biodiversity. Without a strategic long-term research program, this ignorance will continue to hinder appropriate planning and conservation efforts. The government at all levels must step up and support actively a coordinated multi-institutional program of study of Papua’s biodiversity and conservation needs. The authorities need to actively encourage researchers from around the world to focus their work on Papua. A range of nongovernmental institutions should be pressing for this broad change. The change will not happen without strong pressure. Any research program, however, should be constructed in such as way that it can be used to strengthen the most desperately needed capacity of local scientists and institutions. Results and information should be immediately passed back to local communities and authorities, in language that they can understand, and in a way that will raise their awareness and willingness to protect natural resources and biodiversity. Given the lax governmental policies that allow foreign ecotourists to visit remote and ecologically sensitive areas, the unfavorable treatment experienced by most researchers is surprising. Foreign research not only generates knowledge of biodiversity and conservation needs, but also increases the capacity of local partners and generates significant economic benefits, during and often long after the research activities. Raja Ampat is a good example, as discussed above.

Conclusions and Recommendations In conclusion, there is no active species conservation work in Papua except for the recent international field inventories. Past conservation work has focused on

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strategic development and, to lesser extent, protected area management. It is interesting to note that no plan or strategy for species conservation exists (except for crocodile farming, which was developed by Whittaker et al. 1985). There is an urgent need to develop a holistic species conservation strategy, especially to focus on species listed under threatened categories by International Union for the Conservation of Nature and Natural Resources (IUCN 2004), as has been strongly recommended under the current Indonesian Biodiversity Strategy and Action Plans (BAPPENAS 2003). Although existing protected areas constitute the largest provincial network in the country, a significant proportion remain ungazetted, so their future legal status still uncertain. The current coverage of protected areas (proposed and gazetted) should be expanded if all species are to be included in the protected area systems, and there is urgent need to bring all the existing protected area under effective management. There is no easy fix to Papua’s management constraints. A way forward is to strengthen the capacity for better program development (including budgeting and planning) by building the capacity of the Natural Resources Conservation Bureau (BKSDA) staff. Another way forward is to increase long-term sustainability of existing conservation areas by actively involving and influencing Papua socioeconomic development plans at a strategic level. Some may argue that there is already sufficient information and documentation on conservation strategies for Papua. What is lacking is real conservation work on the ground. Future conservation efforts should aim to increase more meaningful and long-lasting field-based conservation, built upon a strong knowledge basis, which also must be generated from on-the-ground field studies.

Literature Cited Agency for Planning and Coordination of Regional Development and Central Bureau of Statistics Propinsi Papua. 2003. Papua dalam angka 2002 [Papua in Figures 2002] BP3D and BPS Propinsi Papua, Jayapura Allen, G.R., and S.J. Renyaan. 1998. Survey of the freshwater fishes of Irian Jaya Phase II (B): 1998 Fishes of the Raja Ampat Islands. Interim research report. Allen, G.R., and S.J. Renyaan. 2000. Survey of the freshwater fishes of Irian Jaya: fishes of the Raja Ampat Islands, Misool Island, and South-Central Vogelkop Peninsula. Interim research report. Anon. 2002. Daftar Lembaga Swadaya Masyarakat (LSM) yang Berkiprah di Papua Menurut Kabupaten [List of NGOs in Papua by kabupaten]. National Development Planning Agency 2003. Indonesian Biodiversity Action Plan. BAPPENAS, Jakarta. Bruner, A.G., R.E. Gullison, R.E. Rice, and G.A.B. da Fonseca. 2001. Effectiveness of parks in protecting tropical biodiversity. Science 291: 125–128. Conservation International. 1999. Lokakarya penentuan prioritas konservasi keanekargaman hayati Irian Jaya. Laporan Akhir [Irian Jaya’s Biodiversity Priority Setting Workshop, final report]. Conservation International, Jakarta. Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam. 2003. Statistik perlin-

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Opportunities and Challenges for Conservation in Papua / 1325 dungan dan konservasi alam Tahun 2002 [Statistics of nature conservation and protection 2002]. Departemen Kehutanan, Jakarta. Food and Agriculture Organization. 1981. National conservation plan for Indonesia Vol VII: Maluku and Irian Jaya. Field Report of UNDP/FAO National Park Development Project INS/78/061. Food and Agriculture Organization, Jayapura. Government of Indonesia. 2000. Undang-undang No. 20 Tahun 2000: tentang otonomi daerah (Decree No. 20/2000: regional autonomy). Government of Republic Indonesia, Jakarta. Government of Indonesia. 2001. Undang-undang No. 21 Tahun 2001: tentang otonomi khusus bagi Provinsi Papua (Decree No. 21: special autonomy for Papua province). Government of Republic Indonesia, Jakarta. Hitipeuw, C. 2003. Status of sea turtle populations in Raja Ampat. In Donelly, 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 Area. Bali, Indonesia. International Union for the Conservation of Nature and Natural Resources (IUCN). 2004. 2004 IUCN Red List of Threatened Species, www.redlist.org. February 2005. Johns, R.J. 1995. Malesia—an introduction. Curtis’s Botanical Magazine 12 (2): 52–62. Kemp, N.J., and J.B. Burnett. 2003. Laporan: Kera ekor panjang (Macaca fascicularis) di Pulau Nugini: penilaian dan penatalaksanaan resiko terhadap keanekaragaman hayati. Indo-Pacific Conservation Alliance and Cenderawasih University, Jayapura. Mack, A.L., and L.A. 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. McKenna, S.A., G.R. Allen, and S. Suryadi (eds.). 2002. A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, D.C. Mittermeier, R.A., C.G. Mittermeier, P.R. Gil, J. Pilgrim, G. Fonseca, T. Brooks, and W.R. Konstant. 2002. Wilderness: Earth’s Last Wild Places. CEMEX, Mexico City. Mittermeier, R.A., C.G. Mittermeier, T. M. Brooks, J.D. Pilgrim, W.R. Konstant, and G.A.B. da Fonseca. 2003. Wilderness and biodiversity conservation. Ecology 100 (18): 10309–10313. Muskita, Y. 2002. Peredaran dan kepemelikan satwa asal Papua di Kota Jayapura: Laporan Survey. World Wildlife Fund, Jayapura. Myers, N. 1988. Threatened biotas: ‘hotspots’ in tropical forests. Environmentalists 8: 187–208. Ohee, H.L., and T. Wakum. 1997. Kapasitas lembaga swadaya masyarakat (LSM) di Irian Jaya: suatu analisa untuk penyusunan program pelatihan. Conservation International–Indonesia Program/Yayasan Pengembangan Masyarakat Desa Irian Jaya, Jayapura. Petocz, R. 1989. Conservation and Development in Irian Jaya: A Strategy for Rational Resource Utilization. E.J. Brill, Leiden. Richards, S.J., and S. Suryadi (eds.). 2002. A Biodiversity assessment of Yongsu: Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, D.C. Sujatnika, P. Jepson, T. Soehartono, R. Crosby, J. Mike, and A. Mardiastuti. 1995. Conserving Indonesia Biodiversity: the Endemic Bird Area approach. Directorate of Forest Protection and Nature Conservation/BirdLife International–Indonesia Program, Bogor. Whittaker, R., P. Sukran, Hartono, and Chadis. 1985. The crocodile resources in Irian

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1326 / yance d e fretes Jaya. World Wildlife Fund–Indonesia/International Union for the Conservation of Nature and Natural Resources, Bogor. World Bank. 2001. Indonesia: environment and natural resources in a time of transition. World Bank, Washington, D.C. World Wildlife Fund. 2000. The Global 200: a representation approach to conserving the Earth’s distinctive ecoregions. World Wildlife Fund, Washington. D.C.

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7.6. Community-Based Conservation in the Trans-Fly Region michele bowe Biological Importance of the Trans-Fly Landscape h e la n d s ca p e o f t h i s ch a p t er is the Trans-Fly region, the southern portion of the island of New Guinea. The region includes the dry lowlands south of the Digul River in Papua and the Fly River in Papua New Guinea (PNG), including a small section bordering the north bank of the lower Fly River. The Trans-Fly supports the most distinct regional avifauna in New Guinea and is listed by BirdLife International as an Endemic Bird Area (Stattersfield et al. 1998). Significant expanses of savanna, grasslands, wetlands, reeds, and monsoon forest support a unique mixture of New Guinea and north Australian flora and fauna. In contrast with the lush tropical areas of most of New Guinea, the Trans-Fly has a distinctly monsoonal climate. Almost 75% of the annual 1,875 mm of rain falls in the December to May wet season; the remainder falls in a dry season from June to November (Paijmans et al. 1971). This results in predominately savannas and monsoon forest vegetation. With a maximum elevation of some 55 m, the landscape is unique in New Guinea for its open flatness as distinct from the rugged mountainous interior of the island, and it strongly resembles the coastal and adjacent areas of northern Australia. Land systems and flora are reasonably well described at least on the Papua New Guinea side, following systematic surveys by the Commonwealth Scientific and Industrial Research Organisation (CSIRO) in the Morehead-Kiunga region. Four broad geographic regions are described: ‘‘the coastal plain; the floodplains of the Fly River and its tributaries; the Oriomo plateau, a low undissected flat to slightly undulating shelf; and the intricately dissected plateau of the Fly-Digoel [Digul] Shelf’’ (Paijmans et al. 1971). The land forms relevant to this chapter are the coastal back plain and the Oriomo Plateau. The Oriomo Plateau, running eastwest across the region (see Figure 7.6.1) is transected by a number of smallish rivers with narrow catchments that drain southwards into the coastal plain. The plateau itself is low-lying, with a maximum elevation of about 55 m above sea level. On poorly drained soils that are seasonally inundated, the vegetation consists of Melaleuca savanna, while on undulating, better drained soils, the vegetation consists of tall mixed savanna and the critically important monsoon forest (Paijmans et al. 1971; Chapter 5.12). The coastal plain consists of flats and low beach ridges and swales near the coast and a back plain, which is crossed by numerous small- to medium-sized rivers. From west to east these include the Bian, Kumbe, and Maro (in Papua), and the

T

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

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Figure 7.6.1. Landforms of the Trans-Fly. Bensbach/Torassi, Morehead, Wassi Kussa, and Mai Kussa (in PNG). Most of the flats are flooded or inundated by freshwater in the wet season, giving rise to some of the region’s most extensive wetlands. Lying roughly between the Mai Kussa River in PNG (14213⬘E 0910⬘S) and the Bian River in Papua (13955⬘E 0807⬘S), the savannas and coastal back plain wetlands are important for wintering Australian birds, large concentrations of waterfowl and migratory paleoarctic waders. A significant proportion of the world population of Little Curlew (Numenius minutus) passes through this area on its annual migration (N. Stronach, pers. comm.). During the wet season, tens of thousands of waterfowl visit the region and of these, significant numbers of Magpie Geese, White Ibis, Black-necked Stork, and Brolga Crane breed. The avifauna is unusually diverse, reflecting the variety of habitats and the conjunction of the Australian and New Guinean subregions. In Wasur National Park, 335 bird species have been recorded (Craven and Bowe 1992), but the full species list is likely to be likely to be much longer. Red-listed species known to be present in viable populations are Scheepmaker’s Crowned Pigeon (Goura scheepmakeri) and New Guinea Harpy Eagle (Harpyopsis novaeguineae) (Stattersfield et al. 1998). Three Trans-Fly endemic bird species have been recorded, including the Fly River Grassbird (Megalurus albolimbatus) and the Grey-crowned Munia (Lonchura nevermanni) (Beehler et al. 1986). Mammals, reptiles, amphibians, and fish of the region have not been surveyed in detail, but are likely to be diverse. Some 63 species of fish are reported from the Bensbach

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Community-Based Conservation in the Trans-Fly Region / 1329

River alone (Allen 1991). The vegetation consists of reeds and tall sedges in permanent swamp; low swamp grassland and Melaleuca swamp forest in semi-permanent to seasonal swamp; and, on slightly higher ground, Melaleuca savanna and Imperata grassland (Paijmans et al. 1971; Chapter 5.12). The vegetation has a long history of manipulation by humans through traditional subsistence activities, notably through the use of fire in grasslands and savanna, and through small scale and local shifting cultivation.

threats The Trans-Fly monsoon forests have a particularly restricted distribution and contain high numbers of endemic plant and animal species. The monsoon forests of the Oriomo River district in the eastern part of the Trans-Fly contain five endemic tree species listed on the International Union for the Conservation of Nature (IUCN) Threatened Trees Register. Large-scale forestry operations in PNG and clearance for agriculture and settlement in Papua threaten almost the entire extent of this forest type in the Trans-Fly (see Figure 7.6.2). The majority of the savannas and wetland areas are also under threat, particularly in Papua. Large areas of the southern lowlands are rapidly being settled by people and converted to agriculture, originally as part of a government-sponsored transmigration program, through which migrants from overpopulated islands largely in western Indonesia were settled in sparsely populated regions in Papua.

Figure 7.6.2. Agriculture, forestry, and settlements in the Trans-Fly.

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The grasslands and wetlands are threatened by exotic weed infestations, including Water Hyacinth (Eichhornia crassipes) and the Giant Sensitive Plant (Mimosa pigra). These weeds pose a critical threat to the integrity of the region’s natural ecosystems. Additional grave threats are the impacts of feral and introduced animals including cattle, horses, dogs, and wild pig, but particularly vast numbers of introduced Rusa Deer (Cervus timorensis). Numerous introduced fish species are also beginning to pose a threat to the region’s native fish species (Kitchener 1997). Other management concerns include non-indigenous fire regimes and large-scale vegetation changes relating to invasion of woody scrub onto the floodplain grasslands (Kitchener 1997; Stronach 2000).

Conservation Areas in the Trans-Fly Eight conservation areas are listed in the Trans-Fly. Of these, only one (Bupul Nature Reserve in Papua) is intended to protect the critically important monsoon forest. Two conservation areas protect the savannas and coastal plain wetlands: Wasur National Park in Papua and Tonda Wildlife Management Area (WMA) in Papua New Guinea. Wasur and Tonda are contiguous across the international border and it is the management of these two areas that formed the nucleus of World Wide Fund for Nature (WWF) activities in the Trans-Fly for many years. Wasur National Park, situated in Merauke Regency, is the management responsibility of the Indonesian Ministry of Forestry’s Directorate of Conservation and Protected Areas (PHKA). The Ministry of Forestry originally declared the Wasur area to be a National Park with a total area of 308,000 ha under a Decree dated 6 March 1990. The park was finally gazetted in 1997 with a total area of 413,810 ha (SK Menteri Kehutanan No. 282/Kpts-VI/1997). Nine major habitat types have been identified, from coastal mudflats, beaches, and mangroves, through flood plains, savanna grasslands, and savanna woodlands inland. Wasur was nominated as Indonesia’s fourth Ramsar Site by the Convention on Wetlands of International Importance Especially as Waterfowl Habitat in 1996. Contiguous with Wasur to the east lies Tonda Wildlife Management Area (WMA). Tonda is situated in the southwest corner of Papua New Guinea’s Western Province. The closest major town is Daru, on the coast some 200 kilometers to the east. In Papua New Guinea, Wildlife Management Areas are owned by traditional communities in which the landowners form a management committee to decide on rules for protection, management, and exploitation of the fauna. Establishment of Wildlife Management Areas involves the consultation and participation of landholding communities before being established by the Department of Environment and Conservation. Tonda WMA was first gazetted on 20 January 1975, with the first set of rules decided upon by the management committee in February 1975. Tonda is the largest of the Wildlife Management Areas in PNG, covering some 590,000 ha with habitats similar to those described for Wasur National Park. Tonda was designated as PNG’s first Ramsar Site in 1993.

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communities and the development context in the trans-fly Language The cultural and linguistic diversity of the island of New Guinea is well documented and the Trans-Fly region is no exception, with approximately 60 documented languages (Wurm and Hattori 1981). The languages and dialects spoken in the region are non-Austronesian languages belonging largely to three language families classified as the Marind; Marori; and Morehead and Upper Maro River families (Wurm and Hattori 1981). The Marind and Marori language families are only found in Papua, whereas the Morehead and Upper Maro River language family straddles the international border with PNG. Within these broad language families are a proliferation of dialects and subdialects. When World Wide Fund for Nature was working with the Tonda Wildlife Management Area Committee through a series of workshops in 2000 to identify how the WMA Committee could be more effective, the Committee members proposed a series of subcommittees based partly along language and dialect divisions. On the Papua side, language is also an important identifier for resource management issues. During the process of trying to incorporate more community input into the management of the park, WWF assisted the formation of various Traditional Community Associations (LMA: Lembaga Masyarakat Adat), by which customary owners grouped themselves into cultural units corresponding to the main language groups.

Socioeconomic Situation During the Dutch and Indonesian administrations of Papua, for ease of schooling, health care, and administrative expediency, villagers were encouraged to settle into permanent settlements along roads or in areas that were easily accessible to administrators (Figure 7.6.3). There are currently 13 settlements within the boundaries of Wasur National Park, including four that are predominantly inhabited by new settlers. In 1992 Marind, Yei, Marori-Mengey, and Kanum people numbered around 2,000 out of a total population in the Park of around 2,500 (Rumawak 1992). More recently in the late 1990s the government settled approximately 500 more people in the village of Sota, the traditional lands of the Kanum people, on the border with PNG. The Indonesian government provides some services to communities, including road access, although often access is possible only in the dry season. Education and health service infrastructure is provided in most communities but service provision is chronically underfunded and, as a result, schools and clinics in remoter villages are nonfunctional. Opportunities have been provided for villagers to participate in the cash economy through the provision of agricultural extension to develop cash crops and small business enterprises, but these have largely been unsuccessful. During the early part of the World Wide Fund for Nature project in Wasur National Park, much emphasis was placed on developing economic opportunities that are based at least in part on natural resources. The government strat-

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Figure 7.6.3. Roads and settlements in Wasur and Tonda. egy has been to replicate the standard local level administration in each village, although frequently villages are too small to provide staff for all the positions required, and people often lack the literacy and other skills to fulfill the duties expected of them. Despite government attempts to provide services and the fact that local people live close to town, most local people still rely predominantly on natural resources, and hunt and gather forest products to augment their subsistence agriculture. The Marind, Yei, and Marori-Mengey people primarily practice sago cultivation and the Kanum are predominantly yam gardeners. Despite numerous outside influences, peoples’ use of local language and their cultural beliefs and practices generally remain strong. In Tonda Wildlife Management Area the development situation is much the same, although without the presence of a nearby major town and consequent pressures due to settlement of outsiders. The area is remote and geographical factors have so far inhibited the development of services, the provision of which is currently poor. The few roads in the district are rough in the dry season and impassable in the wet. There are no banking or credit facilities in the district, only a few trading stores, and schools and health clinics suffer from chronic shortage of resources. Access is primarily via barge to the government station of Morehead, and to Bensbach. There is currently no direct air access. Some 1,500 people are settled in 28 villages throughout the Wildlife Management Area. Most households live largely by traditional subsistence means, exchanging produce and goods locally. Lifestyles revolve around hunting and yam cultivation. The introduction of cash cropping has been stifled by lack of marketing opportunities and by a 32

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kilometer border quarantine zone to prevent the spread of pests and livestock diseases. Despite a history of projects aimed to provide income for local people, such as deer and crocodile farms, these projects have all failed and development activities in the area are virtually nonexistent. Local communities are keen to increase trade in wildlife and forest products and there are many local species with a potentially high economic value, including deer meat, crocodile skins, candlenut (Aleurites sp.), saratoga fingerlings (Sclerophages sp.), Barramundi (Lates calcarifer) and Asteromyrtus symphocarpa—a local tree used for the distillation of essential oil. Local industry based on these resources is underdeveloped and economic activity is constrained by severe difficulties with accessing markets for local produce. Daru, the provincial center for Western Province and the nearest town in PNG is 200 km from Tonda Wildlife Management Area’s western boundary. Local people therefore see more marketing opportunities in freeing up cross border trade into Papua via the border trade post at Sota in Indonesia and thence to the much nearer Merauke.

History of WWF Involvement in the Trans-Fly Region WWF’s involvement in the Trans-Fly region began in Wasur National Park in 1991. The reserve had been identified as one of seven priority areas for management planning, following work by WWF/IUCN in the province during the 1980s (Petocz 1989). At that time (as it is now) the reserve was the most accessible protected area in the entire terrestrial reserve system in the province. It suffered greatly from commercial poaching of Rusa Deer (Cervus timorensis), which resulted in the development of a vast network of bush tracks causing incidental damage to the area, hunting of other native species such as Agile Wallaby (Macropus agilis) when deer were not available, and large-scale burning of grasslands to entice deer to graze. Other threats included extraction of trees for timber and firewood, the quarrying of sand for road construction, and the development of major settlements along the western boundary of Wasur National Park for transmigration schemes, including two places within the reserve. Petocz considered the threats to the reserve to be so great that ‘‘within another three years a considerable portion of habitat and wildlife will be wiped out if an intensive management and protection program is not implemented now’’ (Petocz 1989). At that time the park was the responsibility of a local branch office of the Natural Resources Conservation Department (KSDA: Konservasi Sumber Daya Alam), which had very few staff or funds to have much of an impact in terms of enforcement within the National Park. Prompted by the results of the Petocz study, the Ministry of Forestry’s Directorate of Forest Protection and Nature Conservation (PHPA: Perlindungan Hutan dan Perlindungan Alam,), now the Directorate of Conservation and Protected Areas (PHKA: Perlindungan Hutan dan Konservasi Alam), decided to prioritize Wasur, and WWF was asked to help design and implement a management strategy for the park. The Directorate of Forest Protection and Nature Conservation (PHPA) recognized the importance of integrating local community interests into the management strategy of the National

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Park and, in contrast with other protected areas in Indonesia, no effort was directed at relocating Park residents outside the area. The project started in March 1991 and for the first two years looked mainly at the social aspects of park management to design strategies for involving the National Park’s 13 communities in park planning and implementation. An initial focus was on developing culturally compatible economic incentives for the indigenous people, based, at least in part, on the sustainable use of the National Park’s resources, particularly deer. Concurrent with community development activities, the project was active in seeking solutions to some of the Wasur National Park’s most pressing problems, including poaching, sand mining, and the ad hoc development of roads and settlements. In the course of working on these issues, other problems became apparent, including weed infestations and the change of traditional fire management to less structured practices, leading to large-scale wildfires in the late dry season. It also became evident that within living memory, landscape scale changes to the swamps and grasslands had taken place, where Melaleuca scrub was invading at the expense of grassland. These latter problems did not respect the international border and, as a result of working on community mapping that included landowners from neighboring PNG, it became clear that these problems were also affecting Tonda Wildlife Management Area. The need to link management actions and strategies across the border was obvious. For example, weed control work in Wasur National Park will not be effective unless communities in Tonda WMA have a similar awareness of the problem and have the resources and capacity to act as well. Wildfires also affect both conservation areas. Customary owners of both Wasur National Park and Tonda WMA have practiced fire management extensively for generations, with an awareness of the impacts on landscape, wildlife, important food resource areas, and gardens. Traditionally people practiced burning early in the dry season to protect important resources from late hot season fires, by progressively burning on a small-scale to create a mosaic patchwork effect of vegetation (Craven and Bowe 1992; Stronach 2000). In more recent years, hot fires burn large tracts of land later in the season, with the result that people’s gardens are destroyed, important areas of monsoon forest are damaged, and fires burn uncontrolled until the next rains. Finally, an obvious example of the need to work collaboratively to achieve successful management of Wasur National Park and Tonda WMA is the large number of feral animals, especially Rusa Deer. The deer are implicated in the conversion of grassland to Melaleuca scrub. It is thought that large numbers of grazing deer have reduced the formerly extensive swamp grasses, notably Phragmites karka and Eleocharis spp. The consequent drying out of the swamps and grassland areas has favored the spread of Melaleuca spp. (Stronach 1995, 2000; Kitchener 1997). Although World Wide Fund for Nature had been raising the issues of weed control, feral animals, and fire for several years, the park managers had been reluctant to act on any of these issues. Several reasons may explain their reluctance, including the protected species status of Rusa Deer throughout Indonesia, because of its scarcity in its original range. Any attempt to control or cull the deer popula-

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tion ran contrary to national legislation, even though in New Guinea the species is a destructive exotic. In addition, management of parks in Indonesia relies largely on enforcement rather than proactive management and most park managers had a tropical forestry background, with limited understanding of savannas and the need for active management to maintain largely anthropogenic environments. In 1993, after three years of working in Wasur National Park, WWF Indonesia brought a group of park managers and customary land owners from Wasur National Park to visit Kakadu National Park, a well-managed park in northern Australia with similar environments facing similar threats. The visit was an unqualified success. Managers and communities could see for the first time a wellfunded national park that was actively managed, and where fire management and control of feral animals and weeds were integral components of a park management plan. Interest in tackling similar issues in Wasur National Park grew as a result of the visit, and the possibility of further technical assistance, especially from another park management authority, meant that the managers in Wasur could promote seemingly alien management concepts to head offices in Jakarta with greater confidence. It was within this framework of similarity of environments and threats, management needs, and the desire to promote information exchange that WWF began to formulate its Tri-National Wetlands Program. In 1995 WWF Indonesia sought funding for this program, which would facilitate collaboration between the three conservation areas of Wasur, Tonda, and Kakadu. The initial funding contained a component directed at the WWF South Pacific Programme, to enable them to initiate a conservation effort in Tonda WMA similar to that in Wasur National Park, and to investigate community interest in participating in a cross-border conservation project linking Tonda with Wasur. In 1996 WWF South Pacific Programme and PNG’s Department of Environment and Conservation joined forces to conduct a ten-month project initiation exercise in Tonda WMA. The aim of this exercise was to assess the declining management of Tonda and Tonda’s conservation and development needs, and how these needs could be addressed in a framework of collaboration with Wasur National Park (Chatterton et al. 1997). The report resulting from this initiation exercise—‘‘Caring for Country’’ (Lukautim Graun)—contained a proposal for a five-year program of activities. The report outlined how the landowners of Tonda, acting as its primary managers, could be supported to undertake integrated conservation and development activities in collaboration with a range of stakeholders in the region. There was to be a twoyear hiatus in activity in Tonda following the publication of the report while WWF sought funding to implement the project. Work finally began in Tonda to support the Wildlife Management Area Committee in 1998. The Tri-National Wetlands Program has also continued to expand and develop. In 1998 further funds were sought for the program from Environment Australia. The broad aim of the program is to ‘‘facilitate cooperative and integrated wetlands management between Tonda, Wasur and Kakadu’’ through sharing technical and management expertise; training workshops; collaborative research projects; exchange

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visits for communities, reserve managers, rangers, and government officials; and development of a trilateral intergovernmental Memorandum of Understanding. The program not only looks at technical assistance to help solve common threats and undertake joint training and research programs, but also attempts to share information on different systems of community-based natural resource management, particularly within the context of conservation areas. Wasur, Kakadu, and Tonda have much in common in terms of environments and threats, but have three very different models of management, as a result of the legislative contexts of the three countries. The three management models have their strengths and weaknesses and the Tri-National Wetlands Program looks at ways in which the three areas can learn from each other and, hopefully, adapt the best elements of each system to achieve strong models of community-based conservation area management. The three government conservation agencies from Indonesia, Papua New Guinea, and Australia signed the Memorandum of Understanding for the Tri-National Wetlands Program in June 2002.

Land and Resource Rights and Conservation Legislation Throughout the Trans-Fly, kinship groups built through marriage and blood relationships own collectively all local natural resources, including coastal and land areas. The holders of land and sea country rights, allocations of access, and boundaries of customary-owned land are generally known by those directly involved, although this information is rarely registered in contracts, leases, or other formal agreements. The customary system and traditional methods of managing the use of different areas, habitats, and species, founded on local ecological knowledge, language, and culture, have been the basis for the reasonable allocation of resources for as long as the area has been settled. Custom and traditional laws are provided for in the Fifth National Goal and Directive Principle and in Schedule 2 of the National Constitution of PNG (Callister et al. 1997). Thus the right of communities to own their customary lands is clearly respected and is of prime relevance to the management of the country’s biodiversity and natural resources. The Fauna (Protection and Control) Act of 1966 allows for the establishment of the customary-owned Wildlife Management Areas and also provides for controls on the taking, possession, and trade of species from land of any tenure. Wildlife Management Areas are managed by a committee of locally elected customary landowners. The committee’s function is to develop rules that govern management and exploitation of wildlife, as well as decide on WMA boundaries. Committee members, WMA rules, and boundaries have legal status following publication in the National Gazette and therefore the traditional land and resource rights of local communities are fully recognized nationally and locally reflected in the management rules. The State agency responsible for supporting the management of the conservation area system is the PNG Department of Environment and Conservation (DEC). The PNG DEC provides support to Wildlife Management Areas’ management committees and, in the case of Tonda,

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has one wildlife officer located in the conservation area. In practice, due to budgetary constraints the amount of technical support that DEC can provide to the WMA committee is limited. Although the Fauna Act enables the establishment of Wildlife Management Areas, in the case of Tonda it does not adequately provide a legislative framework for its management. Integrated conservation-and-development-style projects are not addressed within WMA management prescriptions, which are related solely to the management of wildlife and the establishment of associated licenses, fees, and royalties. In 1978, the PNG Conservation Areas Act was formulated to provide for the establishment of Conservation Areas which are intended to be managed by a committee comprising local landowners and local level government. Responsibilities of this committee would include the formulation of a management plan for any development projects and activities. This would allow a broader range of management activities for conservation areas, that could include weed control, fire management, feral animal management, and the development of local income generation activities based on natural resources. World Wide Fund for Nature is currently considering how the Conservation Areas Act might be applied to Tonda. In Indonesia, the rights of indigenous communities to their customary lands and their role in the conservation and management of protected areas are less clearly stated in national legislation than they are in PNG. The management authority for Indonesia’s protected area system rests with the Minister of Forestry’s Directorate of Conservation and Protected Areas (PHKA; formerly PHPA). A National Park Bureau (BTN: Balai Taman Nasional) is set up to manage an established National Park. This unit reports directly to the central government department, although it has a mandate to liaise with local government agencies. Responsibilities of such a unit include the development and implementation of a park management plan. Zonation is a critical element of the management plan, which provides for different levels of use within the park. One of the critical successes of the early WWF work in Wasur National Park was to lobby for a ‘‘Traditional Use Zone’’ that covered the entire Park area. This enabled traditional communities to carry out traditional management activities and customs relating to their land, including hunting and gathering of forest products and traditional fire management. This was the first park in Indonesia with this high level of recognition of local communities’ rights and needs. Since the collapse of the Indonesian New Order government in 1998, government agencies are increasingly cognizant of traditional or customary rights. The WWF project in Wasur has worked closely with the National Park Bureau (BTN) to investigate ways of increasing community participation in management. This has included facilitation of the development of Traditional Community Associations (Lembaga Masyarakat Adat) that would provide a channel for community input to park management. In addition, WWF co-founded, with the National Park Bureau (BTN), a local nongovernmental association to assist with community development work and capacity building for the Traditional Community Associations (LMAs; see below). Despite these very significant advances in recognition of the role and rights of the National Park’s

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traditional communities, there is no legal recognition yet that they are the outright owners of the land.

world wide fund for nature’s conservation approaches in the trans-fly region World Wide Fund for Nature’s approaches to assist community-based conservation and sustainable use of natural resources in the Trans-Fly region have by necessity been different on each side of the international border. Although the prevailing ecological problems and the development needs of communities on both sides are similar, differences of approach have been dictated by a variety of factors. These include: the differences of recognition of resource and tenure rights afforded local people by government systems; national frameworks, structures, and legislation for conservation area management; regional and local level development planning frameworks, including devolution of authority to regional agencies; staffing and resources both of governments and the WWF entities involved; the potential for development of partnerships with other organizations in program development; the presence of local community structures and leadership to link with; levels of community interest and capacity; the political relationship between the two countries and existing treaty arrangements; and practical problems relating to ease of access, costs of fieldwork, and language In Wasur National Park, World Wide Fund for Nature approaches initially focused on working with government institutions and continually pushing to obtain a greater role for local communities in the management of the park. This included encouraging the government to acknowledge that communities should be able to carry out their traditional management activities and lifestyles throughout the park. A substantial amount of effort in the early years of the project involved working with communities to understand their traditional land ownership systems and management practices and their hunting and gardening activities, and promoting the value of maintaining these with the National Park Bureau (BTN). While this was a significant advance in terms of recognizing community rights within the park, it did not address community concerns that they should be able to have a more direct role in determining the management of the park itself on a daily basis. Indeed the areas classified as ‘‘Traditional Use Zones’’ in some ways relegated the national park’s traditional inhabitants to a status of no greater importance than key wildlife sites that were protected within a core zone—another type of zone accommodated within the park’s management plan. Traditional communities themselves were not keen merely to see their lifestyles maintained or quarantined within a Traditional Use Zone, and wanted to be part of a development process that would see them derive income from their natural resources and become part of the modern economy. World Wide Fund for Nature looked at many ways to assist in developing income generating schemes based at least in part on natural resources. A number of successful schemes were developed, including the hunting of deer and pig for fresh and dried meat. Other schemes included the production of salt fish and an essential oil from the leaves of Astero-

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myrtus symphocarpa, a locally abundant tree species in the park. Although WWF was skilled at looking at the environmental sustainability of such ventures, it was not necessarily the most skilled agency at long-term community development ventures. Neither was the National Park Bureau (BTN), whose staff was largely qualified in forest management. Existing local nongovernmental organizations (NGOs) skilled at community development were busy with their own work areas and none was operating within the park area at that time. So WWF, together with the National Park Bureau, investigated opportunities to found a community development NGO that would work within Wasur National Park’s boundaries. It took several years and a few false starts before an NGO, Wasur Conservation Foundation (Yayasan Wasur Lestari), was fully operational with funds of its own and a business plan that would enable it to be self-sustaining in the longer term to support the development needs of park communities. The long-term goal of the Wasur Conservation Foundation is to assist local communities in developing and running their own businesses. Concurrent with this process in the mid- to late 1990s, World Wide Fund for Nature and the NGO Wasur Conservation Foundation (Yayasan Wasur Lestari) were looking at options that would enable local communities to participate more fully in the management of Wasur National Park. Although the Indonesian government remained fully supportive of the need to involve local communities in park management and to develop sustainable industry, this support was aimed more at ensuring park communities were integrated with park management, not necessarily actively involving them in decision making processes. In 1997 and 1998 WWF was part of a Papua-wide consortium of NGOs, The Consortium for Strengthening Traditional Communities of Irian. The aim of the consortium was ‘‘to review the model of natural resource management in Papua to ensure that Papuan people had access and control over natural resource management in order to enable social, cultural and economic benefits to communities.’’ In a workshop held in Merauke in May 1998 the consortium resolved to assist the formation and development of Traditional Community Associations (LMA: Lembaga Masyarakat Adat), particularly in areas where large investment interests were exploiting local natural resources on customary land. A number of Traditional Community Associations (LMAs) were formed around this time, and the four that represented tribal groups within Wasur National Park were included within the network. These were LMA Marind, LMA Kanum, LMA Yei, and LMA Marori-Mengey. The Traditional Community Associations (LMAs) are supported by the NGO Wasur Conservation Foundation (Yayasan Wasur Lestari) to strengthen their role in the management of the park. Ways of doing this are being explored through the WWF-facilitated Tri-National Wetlands Program (TNWP), one of the aims of which is to share knowledge of different models of community management of conservation areas. A number of cross-visits have taken place as part of the TNWP and this has included visits of Wasur National Park staff and communities to Kakadu National Park and Tonda Wildlife Management Area. As a result of these visits, attempts are being made in Wasur to investigate how the four Wasur Tradi-

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tional Community Associations (LMAs) could become an integral part of a Wasur National Park Advisory Board (Dewan Penasehat Taman Nasional Wasur). This Advisory Board would also include representatives of local government, nongovernmental organizations, and scientists (Warta 1998; Rahawarin, pers com). Although it is not yet clear whether the National Park Bureau (BTN) envisages that this Advisory Board can play a decision making role in park management, it is a great step forward from the model that existed before. It could be that the Advisory Board might provide a stepping stone to a fully participatory Wasur National Park Board of Management. The combination of a fully involved community in decision making processes, and a skilled National Park Bureau (BTN) that provides infrastructure and assistance to tackle regional threats such as weeds and feral animals is potentially a powerful one. The National Park Bureau (BTN) already recognizes the importance of traditional management practices such as fire management and the role of community associations that benefit financially from the park is important in assisting in the general enforcement effort of keeping outsider entrepreneurs under control. In reality there is still a long way to go. Ideally the National Park ranger service should be able to provide more benefit to communities as a possible career path for young Papuans living in the park. Currently very few local people have jobs within the park’s management and administrative structure. In addition, although well funded in comparison to Tonda WMA, current funding in Wasur National Park is far from adequate to tackle the plethora of threats. While approaches in Wasur have focused on ensuring greater provision for communities in park management decision making processes, in Tonda the situation is somewhat reversed. The role of local indigenous communities in Tonda WMA seemingly represents the ideal in terms of locally owned and driven natural resource management policy. The twin realities of increasingly slim government support for Wildlife Management Areas and the strong connection of land ownership and royalties from tourism and hunting as a source of revenue mean that there is often conflict in the community in terms of land disputes, leading to disillusionment and occasional unrest. Over the past ten years, financial and technical assistance from government agencies to support the management of Tonda WMA have dwindled to almost nothing. Committee funds derived from royalties relating to hunting and fishing had been mismanaged by the appointed agent. In addition, a land conflict relating to the airstrip at the Bensbach Tourist Lodge has resulted in closure of the lodge to many visitors and thus a subsequent drying up of any further royalties for the Committee and local landowners from this operation. In the face of this dwindling support and conflicts that the communities have not been able to resolve, the WMA had become virtually moribund. Since 1998 WWF has focused its work in Tonda on revitalizing the WMA committee and reaffirming its vital role as the key institution responsible for management of Tonda. Working directly with the Committee, WWF has been trying to encourage a process of management planning for the Wildlife Management Area through a series of practical workshops designed to: review committee structure to improve land-

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owner representation, develop better financial management of royalties, assist the Committee to understand prevailing WMA and other resource management legislation, review the rules for the WMA, and assist the Committee to understand the roles and responsibilities of committee members, rangers, and agents. Reviving the role of the Committee has enabled WWF to also work on raising awareness about the range of ‘‘new’’ threats to the Wildlife Management Area, including weeds, feral animals, and vegetation change. Through awareness talks and slide shows that involved people from Wasur National Park and Kakadu National Park, the Committee has become involved in discussions about management planning for the Tonda WMA and the potential links with the other two protected areas. As WWF has become familiar with the range of issues that the WMA faces and the Committee has become aware of the work that has been carried out across the border in Wasur National Park, WWF and the Committee have begun discussions on a management planning process for the Wildlife Management Area. This process has included: capacity building to manage environmental threats; learning about the importance of the WMA from an international perspective; strategic goal setting for management planning; local level planning to decide on actions related to natural resource management; participating in exchanges relating to research into vegetation change; and examining aspects of traditional fire management that could be revitalized to help mitigate the vegetation change problem in conjunction with Wasur. Identifying mechanisms for long-term funding of the Wildlife Management Area Committee and assisting it to respond to regional issues has had to be a major focus for WWF. One of the key identified needs for the traditional communities of Tonda has been the generation of revenue from natural resources. Tonda WMA is located in one of the poorest parts of PNG—there is little government investment in the province as a whole and community ability to participate in economic opportunities is limited by the scarcity of local markets to raise revenue from natural resources. People generally see great potential in developing market opportunities across the border in fast-growing Merauke. At this stage the potential of trans-border collaboration is seen primarily as one which could facilitate links with the NGO Wasur Conservation Foundation (Yayasan Wasur Lestari) and the development of a community-run trade post at Sota—the closest border post—and thus promote the development of community-based small industry in Tonda. If the sale of local produce across the border post was operated as part of a cooperative community venture this could potentially assist the Committee to fund itself. In reality, unless the Committee can develop a system of self-funding to tackle the range of threats to Tonda and the services required to do this in the absence of government funding, Tonda is unlikely to function as a conservation area in more than name. Without a strong locally-funded community institution to manage even local level management, there is never going to be an effective means of operating more broadly at the cross-border level. Another key component of WWF’s work in Tonda is trying to re-awaken government interest in and commitment to the Wildlife Management Area. Although

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local officers remain committed to the conservation area, in reality they are constrained in providing effective assistance due to the budgetary constraints of the Department of Environment and Conservation. Participation of Tonda WMA in a high profile cross-border program has done much to stimulate interest in Tonda on the part of government officers. WWF is currently investigating funding opportunities that would enable a longer-term project presence in Tonda. In comparison with work in Wasur National Park, the WWF work in Tonda is still in its early stages. In addition to finding long-term locally driven funding for the area, much work needs to be done in looking at capacity building for the local community and government institutions. Other proposed work includes investigating mechanisms for aligning resource management planning to provincial level development planning processes in order to operationalize the organic law reform and better integrate local level plans to regional planning and funding processes. Seeking political endorsement of the cross-border conservation program is a key element of WWF’s work in the region. In the early stages of the development of the Tri-National Wetlands Program it was evident that there was a need for some formal government arrangement through which collaboration between Indonesia, Papua New Guinea, and Australia could take place. The linkages between Wasur and Tonda required special attention because of the political dimension of the shared international border, across which it was hoped that activities could take place. Therefore before looking at mechanisms for collaboration between the three governments, WWF commissioned a report that identified mechanisms for legal and institutional coordination for a proposed cross-border conservation area between Wasur and Tonda. Conclusions of the report were for PNG and Indonesia to proceed with cross-border collaboration through the inclusion of the program within the framework of an existing bilateral treaty between the two governments. The ‘‘Basic Agreement between the Government of Papua New Guinea and the Government of the Republic of Indonesia on Border Arrangements’’ includes articles relevant to the cross-border work particularly and the Tri-National Wetlands Program generally. This includes Article 18 on protection of flora and fauna in the vicinity of the border. Pursuing collaboration through this high level forum has been time consuming and complex. However, the Basic Agreement on Border Arrangements provides the framework in which to address a broader range of issues relevant to the cross-border conservation area, such as cross-visits, joint research and training activities, security issues, border crossings, and trade. Support at this level has particularly helped to ease administrative problems relating to securing visas and border passes for Indonesian and PNG nationals to cross the border at Sota, which is not currently a formal port of entry for either country. Both the Indonesian and PNG government agencies have tabled the Tri-National Wetlands Program at Border Liaison meetings that are held regularly under the Basic Agreement on Border Arrangements, to seek overall endorsement of the program as well as regularly provide updates on program activities. This has provided a structure for long term continuity of program activity even if government officers change, as well provide political endorsement of the various activities that

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take place. In addition to aligning the program to the Basic Agreement on Border Arrangements, WWF has also facilitated a Tri-National Memorandum of Understanding that provides an environment agency framework for the program and hopefully will stimulate continued interest and funding for the program. The Memorandum of Understanding was signed in Indonesia in June 2002.

Extending the Approach: Ecoregion Conservation at the Broad Scale In the late 1990s WWF began to change the way it thinks about, and seeks to implement, conservation globally. Recognizing that conservation efforts must be strategically focused in order to conserve the full range of biodiversity, WWF undertook a global analysis of the world’s ecosystems and highlighted the most urgent conservation priorities as the Global 200. The Global 200 is a comparative analysis of biodiversity to cover every major habitat type, spanning five continents and all the world’s oceans. It uses Ecoregions, which are large units of land or water that contain geographically distinct species, habitats, and processes, as the unit of scale for comparison and analysis. The Global 200 are the most outstanding examples of each of the world’s diverse terrestrial, freshwater, and marine Ecoregions. The Trans-Fly covers parts of four terrestrial Ecoregions, three of which are Global 200, and one freshwater Global 200 Ecoregion. The region then, is clearly of global significance. WWF adheres to the principle that the intrinsic value of biodiversity and its critical importance to human welfare means that it must aim for zero loss of species due to human intervention. In practice this means planning and implementing conservation and development programs on a larger scale than has been attempted so far. It also means broadening the focus to encompass landscape and regional scales, working closely with key stakeholders and using the best scientific information to inform planning and management. Moving to these larger scales will help to address more effectively the broader social, economic, and policy factors that are critical to sustainable livelihoods and ecosystems. Much of the work that has been undertaken in the Trans-Fly over the last ten years will be an important foundation for extending the scale of work even further to the rest of the Ecoregion. WWF is now developing an Ecoregion Conservation Program for the Trans-Fly. Beginning in 2002, WWF has been preparing to develop a biodiversity vision for whole of this unique area, through collecting data and mobilizing partnerships for long-term conservation effort in the region. Development of the conservation program for the Trans-Fly revolves around a number of key elements. First, reconnaissance: a multidisciplinary rapid assessment to help frame the development of an Ecoregion Conservation Plan and identify any urgent needs that require immediate action. Second, a clear biodiversity vision, which will set out long-term (50 year) goals for conservation of the TransFly’s biodiversity, identifying key sites, populations, and ecological processes. The vision will be the touchstone for WWF’s conservation efforts—guiding the development of the Ecoregion Conservation Plan, and strategic decisions as circum-

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stances and opportunities change. Third, the Ecoregion Conservation Plan will set goals over a 10–15 year time frame for conservation of the Trans-Fly’s biodiversity, and identify the actions needed to achieve these goals. Serving as a guide for everyone concerned with biodiversity conservation in the Trans-Fly, the plan will be a comprehensive blueprint for conservation action and identifies first steps on the road to achieving the biodiversity vision. Fourth, the WWF Ecoregion Conservation Plan outlines the contribution that WWF will make towards the Ecoregion Conservation Plan over the next five years. It outlines what WWF needs to do, directly and by enlisting others, to help contribute to the achievement of goals in the Conservation Plan. Coalitions (among NGOs, government agencies, private sector, and customary resource owners) committed to the Ecoregion conservation process have developed. These coalitions bring together resources, knowledge, and partnerships that reflect the full spectrum of stakeholder interests and needs over the long term. Developing an Ecoregion conservation program for the Trans-Fly that includes the means to sustain the multitude of species, habitats, and ecosystems is a formidable task, due to the complexity, large area, and different political contexts of the region. In addition, current knowledge of biodiversity, including species and biological and evolutionary processes, which is vital for the formulation of the vision, is nonexistent over much of the area. Given time, money, and expert assistance however, collecting this information and formulating a vision or biodiversity plan is not an impossible task. However, it will not be sufficient to focus on strategic spatial planning based on biodiversity values at the landscape scale. All the lessons of working in Tonda Wildlife Management Area and Wasur National Park over the years have shown the critical importance of thinking regionally but engaging locally, especially given systems of land tenure and traditional lifestyles. The key to long-term conservation success therefore will be developing and implementing the biodiversity vision in a way that can engage customary landowners in the region. Recognizing this, WWF has been developing a rapid ethnoecological assessment methodology for developing a community vision that is proceeding concurrently with the collection of biological information to inform the biodiversity vision. Essentially this involves identifying how communities value their landscapes and species, as well identifying how communities are important in maintaining biodiversity values (e.g., fire to maintain grasslands, protect monsoon forest). This process has included exploring the values attached to totem species within the clan structures and or specific species that have spiritual, story, hunting, or other significance. The biodiversity vision is based upon representation of all habitats and the needs of focal species and ecological processes as the basis of planning. One of the things the ethnoecology work has focused on is to identify overlap between clan totem species with other focal species that might be used in the vision process (e.g., endemics, migrating and congregating species, species requiring special ranges). Even if no overlap exists, WWF is planning to use clan totems as ‘‘flagship

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species’’ that can provide a layer in the biodiversity vision and be used as a way of increasing community identification with the biodiversity vision. These are very early days in the study and it will be a long task to identify community values over the entire Trans-Fly landscapes, which include around 60 different language groups. It is vital to combine community and biological values if the biodiversity vision and conservation plan are to be relevant to communities and therefore meaningful as tools to achieve long term conservation success.

Summary While in theory Wasur National Park lies far to the opposite end of the continuum of indigenous community input and legal involvement in natural resource management from Tonda Wildlife Management Area, each model can valuably inform the other. Over the past three years WWF has placed considerable emphasis on cross-border collaboration as a means of solving a number of intractable environmental problems, including issues of weed invasion, exotic and feral animals, and fire management. These issues do not respect political boundaries and require concerted efforts on both sides to tackle them successfully. WWF’s Tri-National Wetlands Program is emphasizing reciprocal visits for community members and government officials to raise awareness of environmental threats, conducting joint training workshops and some collaborative research, and sharing knowledge of different models of community management of conservation areas. At one end of the management authority and responsibility continuum, Tonda Wildlife Management Area is struggling for resources and government assistance to implement the ideal of fully recognized and supported land and resource rights. At the other end of the continuum, the customary owners of Wasur National Park, with ten years of funding and support, still strive for autonomy and legal recognition. In a meeting in Merauke, Papua, in February 2000, representatives of Tonda WMA committee showed admiration and envy for the combined resources and collaborative efforts of WWF, the Wasur National Park Authority, the Traditional Community Associations (LMAs), and the NGO Wasur Conservation Foundation (Yayasan Wasur Lestari) and wished for a similar level of assistance. Meanwhile members of the four Traditional Community Associations (LMAs) in Wasur considered that a level of autonomy and recognition of rights to manage their resources in a manner similar to Tonda’s was preferable to the current support. A desirable solution would seem to contain elements of both models. As WWF moves towards a more extensive approach towards conservation in the Trans-Fly, with the development of the Trans-Fly Ecoregion Conservation Program, the lessons learned through cross-boundary community-based conservation in Wasur and Tonda will be vital. The added value that a biodiversity vision and conservation plan for the Trans-Fly brings to previous initiatives in the two countries will be powerful. By engaging stakeholders throughout the vision process and developing strong coalitions, the Trans-Fly Vision and Conservation Plan will reflect the needs, interests, aspirations, and values of the people of Papua and

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Papua New Guinea within a single, coherent framework. It will provide an important reference for national and local projects and can significantly enhance their implementation through the sharing of lessons and identification of new opportunities.

Acknowledgments This chapter discusses the results of many years of collaborative work in southern New Guinea. Although the opinions expressed are my own, it would not have been possible to write this chapter if it were not for the combined efforts over the years of WWF staff, communities and government officers in Indonesia, Australia, and Papua New Guinea. In particular I would like to acknowledge the work of Junne Cosmas, Toby Yakumani, and Biatus Bito of WWF South Pacific Programme; Linke Rahawarin, Zulfira Warta, and Benja Mambai of WWF Indonesia; Paul Mitchell of WWF Australia; Bronwen Fyfe-Golder of WWF International; Adam Tomasek and John Morrison of WWF U.S.; and Marc Wohling.

Literature Cited Allen, G.R. 1991. Field Guide to the Freshwater Fishes of New Guinea. Christensen Research Institute, Madang. Ayres, M. 1983. This side, that side: locality and exogamous group definition in the Morehead area, southwestern Papua. Ph.D. diss., Dept. of Anthropology, University of Chicago, Chicago. Beehler, B.M., T.K. Pratt, and D.A. Zimmerman. 1986. Birds of New Guinea. Princeton University Press, Princeton, New Jersey. Bowe, M. 1997. Turning a threat into an asset: an income generating scheme for community development and exotic species control in Wasur National Park, Irian Jaya, Indonesia. Case study no. 8 in Claridge, G., and B. O’Callaghan (eds.) Community Involvement in Wetland Management: Lessons from the Field. Wetlands International, Kuala Lumpur. Callister, D., A. Casson, and J. Genolagani. 1997. Mechanisms for legal and institutional coordination in proposed south New Guinea (Tonda/Wasur) Cross-Border Conservation Area. In the project initiation report Lukautim Graun: community land care in the south New Guinea savanna region. World Wide Fund for Nature South Pacific Program, Suva. Chatterton, P., J. Cosmas, J. Opu, B. Tapari, and the communities of Tonda Wildlife Management Area. 1997. Project initiation report Lukautim Graun: community land care in the south New Guinea savanna region. World Wide Fund for Nature South Pacific Program, Suva. Craven, I., and M. Bowe. 1992. Wasur National Park Plan of Management final draft, WWF Indonesia Project ID 0105. World Wide Fund for Nature, Jakarta. Eaton, P. 1986. Grassroots conservation: Wildlife Management Areas in Papua New Guinea. University of PNG Land Studies Centre Report 86/1. University of Papua New Guinea, Port Moresby. Fithriadi, R. June 1998. Laporan Kegiatan Fasilitasi Pertemuan Konsorsium Penguatan Masyarakat Adat Irian, Merauke, 5–6 Mei 1998 [Facilitator’s report of the meeting of

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Community-Based Conservation in the Trans-Fly Region / 1347 ‘‘The Consortium for Strenghtening Traditional Communities of Irian’’ held in Merauke 5–6 May]. Workshop report—Biodiversity Support Program Kemala. Kakadu Board of Management and Parks Australia. 1998. Kakadu National Park Plan of Management. Commonwealth of Australia Kitchener, D. (ed.). 1997. Wasur National Park—Tonda Wildlife Management Area biodiversity research planning workshop, Jakarta, 8–10 December 1997. Report of workshop proceedings. World Wide Fund for Nature Indonesia Programme. Paijmans, K., D.H. Blake, P. Bleeker, and J.R. McAlpine. 1971. Land resources of the Morehead-Kiunga area, Territory of Papua and New Guinea. Land Research Series No. 29. Commonwealth Scientific and Industrial Research Organisation, Melbourne. Petocz, R.G. 1989. Conservation and Development in Irian Jaya: A Strategy for Rational Resource Utilization. E.J. Brill, Leiden. Rumawak, A. 1992. Sensus penduduk tahun 1992 pada desa dan kampung di kawasan Taman Nasional Wasur [1992 population census of villages and hamlets in Wasur National Park. WWF project ID 0105. World Wide Fund for Nature. Stattersfield, A.J., M.J. Crosby, A.J. Long, and D.C. Wedge. 1998. Endemic Bird Areas of the World: Priorities for Biodiversity Conservation. BirdLife International Conservation Series 7, Cambridge. Stronach, N. 1995. The rusa deer of Wasur National Park, Irian Jaya. WWF Project ID 0105. World Wildlife Fund. Stronach, N. 2000. Fire in the TransFly Savanna, Irian Jaya/PNG. In Russell-Smith, J., G. Hill, S. Djoeroemana, and B. Myers (eds.) Fire and sustainable agricultural and forestry development in Eastern Indonesia and Northern Australia. Proceedings No. 91. Australian Centre for International Agriculture, Canberra. Warta, Z. 1998. Dewan Penasehat Taman Nasional Wasur [Wasur National Park Advisory Board]. Discussion concept: Wasur National Park Management Plan Workshop. Project report. World Wildlife Fund. Williams, F.E. 1936/1969. Papuans of the TransFly. Clarendon Press, Oxford. World Wildlife Fund. 2003. Ecoregion Conservation: A Portfolio of Stories. World Wildlife Fund, Washington, D.C. Wurm, S.A., and S. Hattori (eds.). 1981. Language Atlas of the Pacific Area: Part 1: New Guinea Area, Oceania, Australia. Canberra: Australian National University.

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7.7. A Non-native Primate (Macaca fascicularis) in Papua: Implications for Biodiversity neville j. kemp and john burke burnett o m et i m e in t h e rec e n t p a s t , Macaca fascicularis, a monkey species also known as the Long-tailed Macaque or Crab-eating Macaque that is native to mainland Asia and western Indonesia, was accidentally or deliberately introduced into the Jayapura area of northeastern Papua. A population of approximately 60 individuals in six troops has become established in and around Jayapura. At present, Jayapura macaque populations are still small and geographically confined. However, there is no physical or ecological barrier to prevent macaque colonization out of the forest fragments it now inhabits into the greater forest hinterland of Papua and Papua New Guinea. The relatively small size and confined area of Jayapura macaque populations makes them amenable to several possible effective population control strategies, but unless such an effort is undertaken in the very near future, macaque colonization of additional New Guinea forests is likely. The presence of macaques in Papua has raised concern both because New Guinea contains a very high degree of endemic and restricted range species that evolved in the absence of nonhuman primates, and because M. fascicularis is highly ecologically adaptable, able to both prey upon and outcompete native fauna for food resources (Baskin 2002; Mapes 2001). Many conservationists and conservation nongovernmental organizations (NGOs), as well as the governments of PNG and Indonesia, are troubled by the possibility that M. fascicularis may become invasive, which would seriously threaten the island’s ecosystems and native and endemic species. On the other hand, some animal welfare groups have voiced concern for the macaques, suggesting either that the animals should be trapped and relocated abroad, or asserting that the threats and risks posed by the macaques are not significant and that New Guinea ecosystems and species will be able to adapt to the presence of M. fascicularis. From a global perspective, invasive exotic species are increasingly recognized as major long-term threats to biological diversity (Baskin 2002; McNeeley et al. 2001; Sherley 2000; Quammen 1998; Veitch 2002). Increasing interest and effort are being spent on mitigating negative impacts of invasive alien species and on preventing additional exotic species from being inadvertently introduced or established. Macaca fascicularis has become an invasive species outside of its natural range on at least two other islands, Mauritius and Angaur (Republic of Palau). The species is thought to have been a factor in the long-term alteration of Mauritian habitats (Mungroo and Tezoo 1999). In a memorable turn of phrase, Quammen

S

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

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(1996: 269), commenting on M. fascicularis in Mauritius, said that the species is ‘‘capable of spreading . . . like a tribe of Visigoths’’ and are a ‘‘much underestimated . . . contributing factor in the extinction of the dodo.’’ In this chapter we provide an abridged, summary report on a study of Macaca fascicularis in Papua; the full report is available at www.indopacific.org/macaca .asp.

Historical Origins of Macaca fascicularis in Papua The first unconfirmed report by a conservation organization of Macaca fascicularis in the Jayapura area was in the 1980s. We conclude that the earliest likely time for an introduction was with the arrival of the Dutch administration in the Hollandia (now Jayapura) area in 1910, but it is more likely to have occurred sometime between 1941 and the 1970s. Suitable habitat for macaques is abundant in Papua, yet macaque distribution remains restricted. From circumstantial evidence we believe macaque introduction into Papua occurred in a single event and not through repeated introductions and subsequent escapes to the wild over a period of time. Local communities in the Kotaraja-Vim area consistently claim that the macaques arrived before the Indonesian administration. But because the Jayapura macaques are resident only in a relatively small area, it seems likely that the introduction occurred more recently. The longer the time since the macaques’ arrival, the further they are likely to have spread, especially because the fragmented forests they now inhabit were part of a continuously forested area not long ago. It may be impossible to determine conclusively when and how macaques arrived in Jayapura, but even if the historical vector were known for certain, the fact would be of limited importance. In the end, the original vector is relevant only to understanding what steps are necessary to prevent future (re)introductions or releases. The essential point is how easy it is for a destructive alien species to take hold in Papua. The prevailing belief seems to be that all local species are ‘‘Indonesian,’’ rather than from Java, Papua, Bali, Sulawesi, and so on; knowledge of basic biogeography or concepts of endemic and alien species is quite poor. Certainly there is very little awareness among local residents, the military, or policy makers of the possible negative consequences that released or escaped exotic pets might have on local ecosystems. Given other pressing political and economic priorities, this is in some sense understandable but no less alarming to anyone concerned with the preservation of New Guinea’s biodiversity.

Macaca fascicularis’s Potential as an Invasive Species M. fascicularis is one of the world’s most numerous and widespread nonhuman primates (Wheatley 1999), second only to the Rhesus Macaque (M. mulatta) of southern to eastern Asia. The natural range of M. fascicularis extends from main-

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land Southeast Asia (including southern Myanmar, southern and eastern Thailand, Cambodia, southern Laos, Vietnam, and Malaysia), through the islands of the Philippines, into western Indonesia (Figure 7.7.1). Within Indonesia, its natural range includes the islands of Sumatra, Borneo, Java, Bali, and eastern Lesser Sunda Islands (Nusa Tenggara) (Poirier and Smith 1974; Supriatna et al. 1996; Wheatley et al. 1996; Wheatley 1999). Macaca fascicularis is neither rare nor endangered because of its abundance and very large geographic range. The species is currently listed under Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES) Appendix II which allows regulated commercial trading and export of animals (UNEP-WCMC 2003) and in the IUCN Red List as ‘‘Lower Risk–Near Threatened’’ (IUCN 2002) which indicates that the species is not threatened with extinction in the medium- to long-term. It is listed as one of world’s worst 100 invasive species by the Invasive Species Specialist Group of the IUCN Species Survival Commission (Lowe et al. 2000). Macaca fascicularis is an alien or exotic species in at least four locations: Mauritius (Sussman and Tattersall 1986), Angaur, Palau (Poirier and Smith 1974), Hong Kong (Walker 2002), and Papua. The species has become invasive in at least two of these locations, Mauritius and Angaur (Cheke 1987; Office of Environmental Response and Coordination 2002), both of which are tropical islands. Angaur is located relatively near to New Guinea and the two share many ecological and climatic characteristics.

Figure 7.7.1. Native range of Macaca fascicularis (shaded).

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Several criteria can help predict the potential invasiveness of M. fascicularis in Papua which should be considered in managing the invasive risk of the species. Low (1999) suggests there are four main factors that cause an alien species to become invasive. These are: the species is an ecological generalist that can adapt to new conditions or habitat; the presence of vacant ecological niches in the new habitat; the absence of threats by predators or diseases that control populations of the alien species in its native range; and the species is favorably adapted to or shows a preference for disturbed habitat. In the case of Macaca fascicularis and New Guinea, all of these factors obtain. Based on the general ecological characteristics of Macaca fascicularis, and evidence from other islands where it is present as an alien species (Mauritius, Palau, etc.), macaques are a potentially invasive species for the entire island of New Guinea.

Ecological Characteristics of M. fascicularis Macaca fascicularis displays robust adaptability that allows it to thrive in a diverse array of tropical habitats. It inhabits mangrove and nipa swamp, coastal forest, and riverine, secondary, and primary forests. M. fascicularis can thrive from sea level to 1,800 meters elevation (B. P. Wheatley, pers. comm.). Primarily arboreal, it is an extremely agile species, able to climb nearly vertical rock cliffs. Unlike its close relative, Macaca nemestrina, which prefers primary forest habitats, M. fascicularis is most successful in and shows a preference for disturbed habitats—secondary forest and the forest periphery, riverine forest, and even urban settings (Wheatley 1999; Sarawak Forest Department 2001; Bonadio, no date). However, it can and does inhabit primary forest as well. M. fascicularis exhibits a tendency to roost in trees along rivers, a behavior called ‘‘riverine refuging’’ (van Schaik et al. 1996). M. fascicularis is highly adaptive to new, dynamic, and often harsh environments. In Angaur, populations of the species persisted despite intense bombing of its habitat during World War II, as well as through several severe cyclones that depleted its food supplies (Wheatley et al. 1999; Poirier and Smith 1974). All of the habitats from the native range of M. fascicularis are present throughout New Guinea. There are many floristic elements common to western Indonesia, mainland Southeast Asia, and New Guinea. In terms of availability of suitable habitat, M. fascicularis could presumably thrive throughout almost all of lowland and parts of upland New Guinea. Because there are many mountain passes in New Guinea below 1,800 meters, the species could cross the Central Mountain Range into the southern half of the New Guinea island. Macaca fascicularis has adapted to several types of lowland habitat in Papua. The original groups of macaques were found in the sago-dominated swamps of Kotaraja. Although little of this habitat now remains due to urban development, one small group is still found there. Breeding troops of M. fascicularis were found in primary forest over limestone (with evidence of very light selective logging), secondary regrowth lowland forest over limestone, and freshwater swamp forest

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dominated by sago (Metroxylon sagu). In Papua, M. fascicularis does not show a preference for secondary forests over primary forests. Three troops are found in primary (or very lightly disturbed) forest, while three troops are in secondary or disturbed forest. One of the troops, Tanah Hitam, is found in very disturbed gardens and sago swamps that have heavy human activity. However, troops in primary forest do spend a large amount of time on the forest edge—which can be considered a more disturbed habitat—probably because many of its favored foods grow more abundantly there. Other troops are found in degraded secondary forest planted with fruit trees, selectively logged forest (tall secondary forest), and some areas of primary forest. Jayapura macaques are also adapted to living near busy roads, near human habitation, and are commonly seen traveling through alang-alang (i.e., anthropogenic Imperata grassland) close to their home range forest. The ever-increasing degree of anthropogenic forest disturbance (mainly through logging or agricultural conversion) within and around remaining tracts of primary forest in Papua is increasing the range of potential habitat for M. fascicularis, and increases its invasive potential both within Papua and PNG as well. Although predominantly arboreal, all Jayapura troops spend significant amounts of time on the ground. The forest floor shows clear signs of pathways regularly used by M. fascicularis to move quickly through its home range. Macaques typically drop to the ground when fleeing after spotting a human observer. From direct field observations, surveys, and community interviews, we concluded with a very high degree of probability that all breeding populations of M. fascicularis were restricted to the area immediately around Jayapura at the end of 2003. Despite repeated and extensive surveys and community interviews in the areas around Sentani, Arso, the PNG border, Mulia, and Manokwari, we found no evidence that macaque populations have spread beyond the Kotaraja area. We therefore also conclude that macaques have not yet entered Papua New Guinea. Macaca fascicularis is not yet an invasive species in Papua, but it has high potential to become invasive in the future. Approximately 60 individual macaques, in six different troops, live in the same general vicinity south of Jayapura city. Data for these troops are displayed in Table 7.7.1, and their locations shown in Figures 7.7.2 and 7.7.3. A single macaque individual was located in the Maunou and Tiaknou hills. Information received at the very end of this study indicated that there may be an additional troop near Entrop that we did not document, but if so, its proximity to Jayapura and its general habitat are not significantly different from the other troops. There are reliable reports of pet macaques being kept in Manokwari, Merauke, and possibly other major towns and cities in Papua. Because these animals seem to be all or mainly individuals kept in cages, pets probably do not represent a major risk of immediate spread. It is alarming nonetheless, given the potential for accidental release/escape of a male-female pair or of a pregnant female. Indeed, pets do escape from time to time, as apparently occurred in 2004 in the Wasur

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Table 7.7.1. Macaca fascicularis populations in Papua Troop Size (No. of Individuals) Geographic Coordinates

Elevation (m a.s.l)

Troop

Location

Kapal

Eastern slopes of Meermokh hill

5

S 235.950⬘ E 14041.350⬘– S 236.110⬘ E 14041.500⬘

15–200

Vihara

Western slopes of 14 Meermokh hill

S 236.050⬘ E 14041.090⬘– S 236.220⬘ E 14041.310⬘

35–185

Meer

Southern and western slopes of Meermokh hill

13

S 236.200⬘ E 14041.080⬘– S 236.240⬘ E 14041.550⬘

20–225

Mangga

Waymhoruk hill

12

S 236.330⬘ E 14041.250⬘– S 236.650⬘ E 14041.280⬘

50–90

Pantai

South end of Waymhoruk hill

5

S 236.650⬘ E 14041.210⬘– S 236.820⬘ E 14041.180⬘

30–60

Tanah Hitam Tanah Hitam sago swamps

10

S 0236.904⬘ E 14040.680⬘– 10–15 S 0236.959⬘ E 14040.979⬘

area (S. J. Richards, pers. comm.). Clearly the issue of pet macaques must be addressed in any comprehensive macaque control strategy in Papua.

feeding behavior and diet preference Macaca fascicularis is a diurnal species, periodically active from dawn to dusk. Approximately 60–90% of the macaque diet consists of fruits, but it ingests a wide variety of other foods including leaves, bark, buds, flowers, seeds, and insects (Bercovitch and Huffman 1999), so they are more accurately characterized as ‘‘opportunistic omnivores’’ (Poirier and Smith 1974). M. fascicularis hunts and consumes small- and medium-sized vertebrates on an opportunistic basis (R. Gibson, pers. comm.; Nowak 1995). M. fascicularis is also a nest predator, taking bird eggs and sometime nestlings (Sarawak Forest Department 2001; Carter and Bright 2002; R. Gibson, pers. comm.). A related species, Macaca nigra, was recorded capturing and devouring a nesting parrot in Sulawesi (R. Sinclair, pers. comm.). In Mauritius, Mungroo and Tezoo (1999) believe macaques to have been an important factor in the extinction of forest birds such as the Scops Owl (Scops commersoni). Macaques continue to be a major pest in Mauritius, inflicting considerable predation damage on native birds, including eating eggs and nestlings and possibly adult birds (R. Gibson, pers. comm.). Remote trigger cameras have provided photographic documentation of macaque predation on bird nests (Carter and Bright 2002). Gibson (pers. comm.) also suspects opportunistic predation by macaques on endemic geckos (Phelsuma spp.).

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Figure 7.7.2. Map of Jayapura, Sentani, and Arso areas showing the distribution of Macaca fascicularis in Papua. In Sarawak, eastern Malaysia, M. fascicularis has been observed to take insects, frog eggs, crabs, and other coastal invertebrates (Sarawak Forest Department 2001). According to Nowak (1995), all macaque species will eat insects and other small invertebrates when they are available and occasionally take eggs and small vertebrates. Macaca fascicularis has also been recorded eating crabs, crustaceans, shellfish, and other littoral animals exposed to the tide (Lekagul and McNeely 1977). Macaca fascicularis also regularly raids agricultural plots, taking cassava, maize, beans and many other domesticated vegetables and fruits, and is therefore consid-

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Figure 7.7.3. Home ranges of five Macaca fascicularis troops at Vim, Kotaraja.

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ered a major pest by many rural communities. With the expansion of agricultural areas and human population, this particular behavior of M. fascicularis may be responsible for the increase in the species’ range (Wheatley et al. 1996). Unlike New Guinea’s native mammal fauna, M. fascicularis has opposable thumbs, which allows it to open fruit casings and nuts and thus to exploit a wide range of food items. Macaques are consequently a strong competitor with native species for food resources. This generalist behavior is extremely advantageous to a species colonizing new areas because they are not limited by specific food requirements. This generalism allows macaques to move into other species’ niches, thus increasing M. fascicularis’ invasive potential. The diet of M. fascicularis in Papua is consistent with its diet in its native range and other colonized habitats. We documented that its diet consists largely of fruit, especially figs (Ficus spp.), and fruits of planted fruit trees. Macaques also eat leaves and young shoots, especially Intsia bijuga and Pandanus spp.; flowers; roots (cassava); bark (of several spp. of liana, Aglaia, and Pometia pinnata); sago pith (when available after processing by humans); and invertebrates, including grasshoppers and sago grubs (larvae of Rhynchophorus ferrungineus or R. bilineatus). Although we documented the commonly eaten items in its diet, our list cannot be considered complete. Importantly, most of the items used as food are widespread throughout lowland forest in Papua, and so it is highly probable that M. fascicularis could easily survive in areas of New Guinea outside its present range. We were not able to confirm through direct observation whether macaques in Papua are nest predators, but because nest predation is documented in Mauritius, and based on other evidence we collected, it is reasonable to infer that macaques are nest predators in Papua. Macaques take small lizards in Mauritius, and so are likely to take lizards in Papua as well. Further evidence for these claims comes from our comparative plot-based studies of bird and lizard diversity and abundance inside and outside macaque ranges in Papua.

competitors, predators, parasites, and pathogens Throughout its native range, M. fascicularis lives sympatrically with a number of other diurnal nonhuman primates. Individual home ranges of M. fascicularis in Sumatra and Borneo may overlap with the home ranges of two or three species of langur, two or three species of Sumatran or Bornean langur, two or three species of ape, and another macaque, M. nemestrina. This co-existence of various primate species undoubtedly creates competitive pressure for resources, which will in turn limit the carrying capacity (or the maximum population size with a home range) of M. fascicularis. In its native range, M. fascicularis also has a number of predators and competitors, including panthers and sun bears, which may control its population (Hoogerwerf 1970; MacKinnon et al. 1996; Wheatley 1996). Predation by large carnivores and snakes (and possibly large raptors) is a likely source of mortality for macaques in its native range; this pressure is largely absent on New Guinea. With the exception of hunting dogs and (perhaps) large snakes of the family

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Boidae (boas and pythons), M. fascicularis has no predators or competitors in New Guinea. Various poisonous snakes may possibly envenom M. fascicularis. But the relative absence of predation and competition gives macaques on New Guinea an ecological opportunity for a high potential population growth rate, and thus increases its invasive potential. Hunting by humans is a significant source of mortality to M. fascicularis in Papua. One hunter, Mr. Ruatakurei from Kotaraja, reported that he caught fourteen individuals within two years, which represents up to 20% of the total macaque population in Papua. Human hunting of these populations may be the major factor that has limited macaque population expansion to date. Smaller troop sizes may be a reaction to hunting (Wheatley et al. 1999). Smaller troops are harder for humans to detect than larger troops, so human hunting would be a selective pressure towards smaller troops. It is notable that group sizes in Papua are smaller than those seen in Sumatra or Borneo. Another reason that the range of Jayapura macaques has not yet expanded more broadly is that populations may have been suppressed by a disease endemic to Papua to which they had not been exposed before their arrival, such as a different, more virulent strain of malaria or arbovirus (D. A. Polhemus, pers. comm.). If so, the pathogen-driven suppression of the population may or may not continue, since host immunological resistance and adaptation of the infective agent are dynamic evolutionary processes.

Impact of Macaca fascicularis on Biodiversity in Papua We carried out field plot studies in areas of primary and secondary forest, both with and without M. fascicularis, to census native fauna and test for discernable impact of macaques on these taxa. Bird, small reptile, and large mammal fauna all have lower diversities and numbers in areas that support M. fascicularis, compared to similar habitats that do not support M. fascicularis. This is strong evidence that M. fascicularis disturbs or displaces native Papuan species. For this reason alone, macaque populations in Papua should be managed to reduce or eliminate this risk. Results for birds in our field plot surveys using point census are shown in Table 7.7.2. Significantly higher numbers of individual birds and bird species were recorded at sites without M. fascicularis. Bird species discovery curves are steeper and higher for sites without M. fascicularis (Figure 7.7.4), demonstrating higher diversity for sites where M. fascicularis is absent. Secondary sites were also more diverse than the paired primary sites, which we attribute to edge effects. The difference in average diversity (number of bird species recorded) between sites with M. fascicularis and sites without M. fascicularis is highly significant (F1,76  19.804, n  80, p ⬍ 0.001). The difference in average abundance (number of individuals recorded) between sites with M. fascicularis present and sites without M. fascicularis is also highly significant (F1,76  31.644, n  80, p ⬍ 0.001). The differences in average number of bird species and individuals recorded in point

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Table 7.7.2. Bird surveys in primary and secondary forests with and without Macaca fascicularis MacKinnon list method No. of bird species recorded

Primary with M. fascicularis (A)

No. of species recorded

Point census method Mean Mean no. of no. of species individuals

Shannon diversity index

22

19

5.05

7.45

2.54

Primary without M. fascicularis (C)

33

27

6.55

11.06

2.72

Secondary with M. fascicularis (B)

26

22

4.85

7.13

2.55

Secondary without M. fascicularis (D)

36

31

7.35

14.33

2.82

Forest type

Figure 7.7.4. Bird species discovery curves generated using the MacKinnon list method.

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A Non-native Primate (Macaca fascicularis) in Papua / 1359

censuses between habitats (with or without M. fascicularis) were not statistically significant. Thus the presence of M. fascicularis is associated with lower numbers and diversity of birds. Qualitative patterns may also be important. For example, sites with M. fascicularis have noticeably fewer species and lower abundance of pigeons (Columbidae), parrots (Psittacidae), birds of paradise (Paradisaeidae), and frogmouths (Podarigidae). These species (many of which are endemic to New Guinea) may be more susceptible than other birds to any nest predation or competition for food resources imposed by M. fascicularis. The design of our surveys did not allow us to test this hypothesis. It is suggestive that we were unable to locate any bird nests at all within M. fascicularis’ home ranges (even though more survey time was spent in those areas), while several nests were recorded in the comparison sites. In our field plot study, the difference between sites with and without M. fascicularis was dramatic for lizards, both in terms of abundance and species diversity. Many more species and individuals (F1,44  33.88, n  48, p ⬍ 0.001) and individuals (F1,44  32.035, n  48, p ⬍ 0.001) were found in the sites without M. fascicularis. The primary forest site without M. fascicularis was the richest site for lizards. For mammals and large reptiles, our study found small but potentially ecologically significant differences between sites. No large arboreal mammals were found in the secondary site where M. fascicularis was present. Visibility was the best at this site, yet only the small nectar feeding Sugar Glider (Petaurus breviceps) was found. The large, slow-moving Common Spotted Cuscus (Spilocuscus maculatus) was found in both sites without M. fascicularis, whereas the Northern Common Cuscus (Phalanger orientalis orientalis) was observed in the primary site with M. fascicularis where Pometia pinnata (a food plant) is dominant. Only one species of large snake, Morelia amethistina amethistina, was recorded in our surveys, and only at sites without M. fascicularis. Smaller ground-dwelling and arboreal snakes were recorded from sites that supported M. fascicularis. No arboreal Macropodidae (tree kangaroos) were found in the area, as they were probably hunted out long ago; no conclusions can therefore be drawn for that family. Although census data are insufficient to draw definitive conclusions, the data collected do strongly support the hypothesis that M. fascicularis negatively impacts large mammals (especially Phalangeridae) and reptiles, presumably through increased competition over food resources. Increased competition with these mammals may reduce successful reproduction and therefore reduce food supplies for larger predators such as Morelia amethistina.

Impacts of Macaca fascicularis on Humans in Papua

crop raiding The role of macaques as crop raiders has been amply documented (Chalise 2001; Hill 1998; Wheatley 1999; Wheatley et al. 1999). As opportunistic foragers, macaques feed not only on wild items but also cultivated crops and fruits. Wheatley

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et al. (1999) noted that macaques in Angaur (Palau) were considered by local residents to be a major pest because of their garden raiding. The state government of Angaur described the macaques as a ‘‘serious pest and a major threat to Angaur’s agriculture . . . causing severe damage to Angaur’s fruits and crops’’ (Office of Environmental Response and Coordination 2002). Antipathy among Angauran villagers towards macaques for damage to their crops has given rise to hunting of macaques by the local community, as well as an Angaur state government-sponsored eradication program established in June 2001. A report on this eradication effort notes that ‘‘in this campaign over 500 monkeys were eliminated from the island. This has not completely solved the problem, so the question remains how to control this pest species in Palau’’ (Office of Environmental Response and Coordination 2002). Macaca fascicularis troops in Papua also frequently raid crops, taking a variety of vegetables and fruits (cassava, maize, beans, peanut, papaya, bananas, sweet potato, mango, young coconut, pineapple, breadfruit, and jackfruit). Local farmers in the Jayapura area traditionally plant a variety of fruits and vegetables in gardens, and fruit trees in secondary forest. In many areas, the main agricultural system is swidden, with gardens located on the forest edge. This anthropogenic disturbance of native ecosystems creates new potential food sources, as well as a suitable habitat, for the possible future expansion of M. fascicularis within Papua and Papua New Guinea. M. fascicularis causes significant economic damage to poor rural farmers in Papua through crop raiding. Our estimate of the economic damage to farmers in the Kotaraja area alone caused by macaque crop raiding is on the order of Rp 30,000,000 per year (US$ 3,500/year). If the range and population of M. fascicularis increases, this species will cause far more widespread and serious losses to the regional economies in Papua and PNG.

health M. fascicularis is also the carrier of a number of diseases that can potentially spread to humans (Baskin 1999; Brown 1997). Within its native range in Indonesia, the Philippines, and Malaysia, macaque individuals have been documented as having a high positive response for many viral antibodies, indicating that diseases are highly prevalent within wild populations (Matsubayashi et al. 1992). These diseases, for which macaques may be either reservoirs or potential vectors to humans, include B-virus (Cercopithecine herpesvirus 1 [CHV-1] or Herpesvirus simiae), rabies, Simian Varicella Virus, Cytomegalovirus, Rhesus Rhadinovirus, Ebola (Reston type), Hepatitis E, and other viruses (see Baskin 1999). Of particular concern to humans is B-virus, since it is the most common and potentially pathogenic virus in wild populations of Macaca fascicularis. The estimated prevalence of B-virus in individual adult wild macaques is 73–90% (UNEPWCMC 2003; Baskin 1999; Ostrowski et al. 1998). While B-virus is generally benign in macaques, all carriers shed viruses and can transmit the disease to humans and other primate species. In humans, B-virus is highly pathogenic, causing a

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A Non-native Primate (Macaca fascicularis) in Papua / 1361

rapidly ascending encephalomyelitis which is fatal in about 55% to 80% of cases (Laboratory Primate Newsletter 1995; Baskin 1999; Centers for Disease Control 2003). Those that survive the disease suffer severe brain and nervous system damage (Laboratory Primate Newsletter 1995; B Virus Working Group 1987; Baskin 1999; Brown 1997; Centers for Disease Control 2003). Macaques are also potential carriers of rabies. All wild macaques from rabiesinfected areas are assumed to be at risk. Papua is currently rabies-free, but Indonesian authorities are aware of the potential for macaques to act as a vector for the disease. We tested Jayapura macaques for rabies and Poxvirus, with negative results. Pathological tests on brain and major organ tissue were normal. B-virus, hepatitis, and other viruses could not be tested because there are inadequate testing facilities in Indonesia, and because of a ban on export permits for blood specimens. Given the extreme virulence of B-virus in particular, a precautionary approach should be taken. B-virus is a rare disease in humans and generally only those who work with macaques in laboratories or zoos are at high risk. However, this is simply a function of exposure—any humans that come into direct contact with wild macaques are potentially at risk. Unless proven otherwise, it should be assumed that zoonotic viruses endemic to M. fascicularis are present in Papuan populations. Papuan macaques thus pose a potential heath threat to humans and livestock, though the risk is small to anyone except those in direct contact with animals, such as hunters, whose chance of exposure to bites or blood is high.

Summary Current populations of Macaca fascicularis in Papua consist of six separate troops that inhabit primary and secondary forest ‘‘islands’’ south of Jayapura city. The current forest habitat of Jayapura macaques does not prevent its colonization of new areas, because the troops can and do move across unforested areas. Macaques have a negative impact on native New Guinea species, ecosystems, and rural economies, and also pose a significant public health risk. Unless population control measures are taken in the very near future, Macaca fascicularis populations in Papua are highly likely to become invasive. If this occurs, there is a high probability of additional and more severe negative impacts on the environment and rural livelihoods. Given the probable consequences of macaques becoming invasive, immediate implementation of a risk management program, involving threat removal (i.e., population control), community awareness and dialogue efforts, and capacity and policy-level enhancements, is therefore of the highest priority.

Acknowledgments Material in this chapter was previously published as part of a project undertaken in 2002–2003 by Indo-Pacific Conservation Alliance (IPCA) in partnership with

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Cenderawasih University (UNCEN). The authors wish to thank our partners at UNCEN, particularly S. J. Renyaan, F. Wospakrik, E. Holle, and V. Agustini. Extraordinary thanks to Pieter M. I. Torobi for assistance and invaluable expertise. Special thanks also go to the dedicated team of biology students from Cenderawasih University: H. M. Buiney, A. R. Yoku, H. Manufandu, L. Burame, H. Wambrauw, K. Seh, E. Awundi, T. Buiney, A. S. Tipawael, Y. D. B. Maro, R. H. Mambrasar, I. Panjaitan, and M. Simbiak. Thanks also to BAPPEDALDA Papua Province, BAPPEDALDA Kotamadya Jayapura, Nature Conservation Authority, Department of Forestry, and the Police of Papua for supporting the project and approving necessary documents; Wayne Takeuchi and University of Papua Herbarium for identifying plants specimens; YPLHC and Roy Rindo-Rindo (WWF) for assistance on awareness campaigns; and Mapping and Inventory Section, Department of Forestry for their cooperation in providing digital maps. We also wish to thank many others who provided helpful guidance and comments during project research and writing of the original report, as well as others who facilitated the project in critical ways; space limitations prevent their full listing here but are in the Final Report. Finally, a special thanks to Prof. Jared Diamond for his interest in and support of this research, and whose attention to this problem was an early catalyst and a continuing motivation.

Literature Cited Baskin, G.B. 1999. Pathology of Nonhuman Primates. Tulane Regional Primate Research Center, Tulane University, Covington, Louisiana. Available at www.primate.wisc.edu/ pin/pola6-99.html. Baskin, Y. 2002. A Plague of Rats and Rubber-Vines: The Growing Threat of Species Invasions. Island Press, Washington, D.C. Bercovitch, F.B., and M.A. Huffman. 1999. The macaques. In Dolhinow, P., and A. Fuentes (eds.) The Nonhuman Primates. Mayfield Publishing, California. Bonadio. no date. Macaca fascicularis. Museum of Zoology, University of Michigan, Ann Arbor. Available at http://animaldiversity.ummz.umich.edu/accounts/macaca/ m._fascicularis.html. Brown, D.W.G. 1997. Threats to humans from virus infections of non-human primates. Reviews of Medical Virology 7: 239–246. Carter, S.P., and P.W. Bright. 2002. Habitat refuges as alternatives to predator control for the conservation of endangered Mauritian birds. Pp. 71–78 in Veitch, C.R., and M.N. Clout (eds.) Turning the Tide: The Eradication of Invasive Species. IUCN/SSC Invasive Species Specialist Group. International Union for the Conservation of Nature and Natural Resources, Gland, Switzerland, and Cambridge, UK. Centers for Disease Control. 2003. National Center for Infectious Diseases. B Virus (Cercopithecine herpesvirus 1) Infection. Center for Disease Control, Atlanta. Available at www.cdc.gov/ncidod/diseases/bvirus.htm. Accessed 19 July 2003. Centers for Disease Control B-Virus Working Group. 1987. Guidelines for prevention of Herpesvirus Simiae (B Virus) infection in monkey handlers. CDC MMWR Weekly 36 (41): 680–682, 687–689.

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A Non-native Primate (Macaca fascicularis) in Papua / 1363 Chalise, M.K. 2001. Crop raiding by wildlife, especially primates, and indigenous practices for crop protection in Lakuwa Area, East Nepal. Asian Primates 7 (3–4): 4–9. Cheke, A.S. 1987. An ecological history of the Mascarene Islands, with particular reference to extinctions and introductions of land vertebrates. In Diamond, A.W., A.S. Cheke, and Sir H.F.I. Elliot (eds.) Studies of Mascarene Island Birds, for the British Ornithologists’ Union. Cambridge University Press, Cambridge. Hill, D., and K. Hill. 1998. Primates as pests: conflict and conservation. Abstracts of the Congress of the International Primatological Society. Hoogerwerf, A. 1970. Udjung Kulon. E.J. Brill, Leiden. International Union for the Conservation of Nature and Natural Resources (IUCN). 2002. 2002 IUCN Red List of Threatened Species. Available at www.redlist.org/search/ details.php?species12551. Accessed 20 October 2003. 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. Available at www.indopacific.org/macaca.html. Laboratory Primate Newsletter. 1995. New B Virus Guidelines Available. Laboratory Primate Newsletter 34 (2): 5. Lekagul, B., and J.A. McNeely. 1977. Mammals of Thailand. Sahakarnbhat, Bangkok. Low, T. 1999. Feral Future: The Untold Story of Australia’s Exotic Invaders. Penguin Books Australia. Lowe, S., M. Browne, and S. Boudjelas. 2000. 100 of the world’s worst invasive species: a selection from the Global Invasive Species Database. Aliens 12. Lift-out booklet. Invasive Species Specialist Group of the IUCN Species Survival Commission. MacKinnon, K., G. Hatta, H. Halim, and A. Mangalik. 1996. The Ecology of Kalimantan–Indonesian Borneo. Periplus Editions (HK) Ltd., Hong Kong. Mapes, T. 2001. Monkeys threaten New Guinea’s wildlife: greedy macaques have bad reputation among locals. The Asian Wall Street Journal, 2 October 2001. Matsubayashi, K., S. Gotoh, Y. Kawamoto, T. Watanabe, K. Nozawa, M. Takasaka, T. Narita, O. Griffiths, and M.A. Stanley. 1992. Clinical examinations on Crab-eating Macaques in Mauritius. Primates 33 (2): 281–288. McNeeley, J.A. (ed.). 2001. The Great Reshuffling: Human Dimensions of Invasive Alien Species. IUCN, Gland, Switzerland, and Cambridge, UK. Mungroo, Y., and V. Tezoo. 1999. Control of Invasive Species in Mauritius. In Lyons, E.E., and S.E. Miller (eds.). Invasive Species in Eastern Africa: Proceedings of a Workshop Held at ICIPE, July 5–6, 1999. Nowak, R.M. 1995. Walker’s Mammals of the World. Johns Hopkins University Press, Baltimore. Available at www.press.jhu.edu/books/walker/primates.cercopithecidae .macaca.html. Office of Environmental Response and Coordination. 2002. National Report to the United Nations Convention to Combat Desertification, Office of Environmental Response and Coordination, Office of the President of the Republic of Palau, April 2002. Poirier, F.E., and E.O. Smith. 1974. The Crab-eating Macaque (Macaca fascicularis) of Angaur Island, Palau, Micronesia. Folia Primatologica 22: 258–306. Quammen, D. 1996. Song of the Dodo: Island Biogeography in an Age of Extinctions. Scribner, New York. Quammen, D. 1998. Planet of weeds: tallying the losses of earth’s animals and plants. Harper’s Magazine October 1998: 57–69. Sarawak Forest Department. 2001. Macaca fascicularis. Forest Department Sarawak. www.forestry.sarawak.gov.my/forweb/wildlife/fauna/mammal/itmac.htm.

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1364 / n e v i l l e j. k e m p & jo h n b u r k e b u r n e t t Sherley, G. (technical ed.). 2000. Invasive species in the Pacific: a technical review and draft regional strategy. SPREP, Apia, Samoa. Supriatna, J.A., Yanuar, Martarinza, H.T. Wibisono, R. Asinaga, I. Sidik, and S. Iskandar. 1996. A preliminary survey of Long-tailed and Pig-tailed Macaques (Macaca fascicularis and Macaca nemestrina) in Lampung, Bengkulu and Jambi Provinces, Southern Sumatra, Indonesia. Tropical Biodiversity 3 (2): 131–139. Sussman, R.W., and I. Tattersall. 1986. Distribution, abundance and putative ecological strategy of Macaca fascicularis on the island of Mauritius, southwestern Indian Ocean. Folia Primatologica 46: 28–43. United Nations Environment Programme–World Conservation Monitoring Centre. 2003. UNEP-WCMC Species Database: CITES-Listed Species. Available at http://sea.unep wcmc.org/isdb/CITES/Taxonomy/tax-species-result.cfm?displaylan guageeng& GenusMacaca&Speciesfascicularis&sourceanimals&Country& tabname legal. van Schaik C.P., A. van Amerongen, and M.A. van Noordwijk. 1996. Riverine refuging by wild Sumatran Long-tailed Macaques (Macaca fascicularis). In Fa, J.E., and D.G. Lindberg (eds.) Evolution and Ecology of Macaque Societies. Cambridge University Press, Cambridge. Veitch, C.R., and M.N. Clout (eds.). 2002. Turning the Tide: The Eradication of Invasive Species. IUCN SSC Invasive Species Specialist Group. International Union for the Conservation of Nature and Natural Resources, Gland, Switzerland, and Cambridge, UK. Walker, B. 2002. Hong Kong determined to deal with monkey problem. Reuters Press Release, April 3, 2002. Available at www.enn.com/news/wirestories/2002/04/04032002/ reu_46836.asp. Wheatley, B.P. 1999. The Sacred Monkeys of Bali. Waveland Press, Long Grove, Illinois. Wheatley, B.P., D.K. Harya Putra, and M.K. Gonder. 1996. A comparison of wild and food-enhanced long-tailed macaques (Macaca fascicularis). In Fa, J.E., and D.G. Lindberg (eds.) Evolution and Ecology of Macaque Societies. Cambridge University Press, Cambridge. Wheatley, B.P., R. Stephenson, and H. Kurashina. 1999. The effects of hunting on the Longtailed Macaques of Ngeaur Island, Palau. In P. Dolhinow, and A. Fuentes (eds.) The Nonhuman Primates. Mayfield Publishing, California.

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7.8. Exotic Herpetofauna: A New Threat to New Guinea’s Biodiversity? burhan tjaturadi, stephen richards, and keliopas krey n v as i v e ex o t i c s p e c ie s are a major threat to native biotas, and their effects on natural ecosystems may be particularly severe on islands (O’Dowd et al. 2003). The large tropical island of New Guinea is one of the last great wilderness areas and has extraordinary levels of endemism and biotic diversity (Mittermeier et al. 2002, 2003). An assessment of conservation priorities for the western half of New Guinea (Papua Province of Indonesia, formerly Irian Jaya) identified exotic fauna as a serious threat to New Guinea’s biodiversity (Conservation International 1999). Of particular concern have been feral populations of macaques in northeastern Papua (Chapter 7.7) and exotic freshwater fishes (Allen et al. 2000, 2003; Chapter 5.5). The introduction of exotic herpetofauna into Papua, primarily through the pet trade, is also a serious concern. Here we document a total of eight exotic species of herpetofauna that have been introduced to the island of New Guinea, seven of which are known exclusively from Papua. The potential for these species to spread into Papua New Guinea is extremely high. All species records are substantiated by voucher specimens or photographs. Three species of exotic herpetofauna have previously been documented from the island of New Guinea. Bufo marinus: The Cane Toad (Bufo marinus), was introduced to eastern New Guinea to control sweet potato moths and now occurs in isolated populations on New Ireland, New Britain, and in both southern and northern mainland Papua New Guinea (Lever 2001). It also occurs widely in the Solomon Islands but has not been reported from Papua to date. Remarkably, B. marinus has been present on the small island of Daru in southwestern Papua New Guinea for many years (Lever 2001) but appears not to have reached the nearby mainland where the vast tropical savannas of the Trans-Fly provide a perfect environment for this tropical toad (S. Richards, pers. obs.). Bufo melanostictus: The widespread Asiatic Toad (B. melanostictus) was reported from the vicinity of Manokwari in western Papua (Menzies and Tapilatu 2000) and a second population has now become established in Sorong, Papua (K. Krey, B. Tjaturadi, and S. Richards, pers. obs.). Limnonectes cancrivora: Menzies (1996) reported the large southeast Asian frog Limnonectes cancrivora from the urban areas of Sorong, Sentani, and Jayapura, and in 2003 and 2004 an additional population was found at Manokwari (K. Krey, pers. obs.). Menzies (1992) also reported the related L. verruculosa from western

I

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

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Papua based on call data only, but no confirmation of this species’ occurrence in the province has been forthcoming and we exclude this species from the list of exotic herpetofauna in Papua pending conclusive evidence. Five additional species of herpetofauna were recently documented in Papua. Trachemys scripta: The Red-eared Slider (Trachemys scripta), is a freshwater turtle with a native distribution extending from mainland North America to Brazil (Ernst and Barbour 1989). It is commonly sold in pet shops in Indonesia; during 2003 and 2004 young animals were available for sale in Jayapura, the capital of Papua. A large specimen in the collection of the Biology Department, State University of Papua (UNIPA: Universitas Negeri Papua) was collected from the wild in the vicinity of Manokwari. Iguana iguana: The Green Iguana (Iguana iguana) is a herbivorous lizard native to the neotropics. In August 2004 a single specimen of the Green Iguana was found in the suburbs of Jayapura. This species is common in the Indonesian pet trade, but there is no evidence that populations have become established in New Guinea. Lycodon aulicus: Two specimens of this southeast Asian colubrid snake were collected from the suburbs of Sentani in early 2005. Mabuya multifasciata: This skink species has a natural distribution encompassing much of lowland Southeast Asia. It has recently become established in gardens around Sentani. Previous reports from far-western Papua probably also represent human introductions. Cuora amboinensis: This freshwater turtle is commonly sold in the pet trade in Indonesia and at least one specimen is known to have escaped in Papua, where the species is occasionally kept as a pet. Table 7.8.1 provides details of the exotic herpetofauna that have been documented from New Guinea. The impacts of exotic herpetofauna on native New Guinean biota have not been assessed and studies to redress this lack of information are urgently required. Some species, such as the Green Iguana, are unlikely to establish viable populations or may establish populations that are highly localized. Other species with high fecundity and rapid dispersal abilities are likely to have serious impacts on native biota. Potential impacts include direct predation by introduced species on native fauna, the introduction of diseases, and competitive interactions with local species. Of particular concern is the introduction and rapid establishment and spread of Bufo melanostictus in Papua. Toads of the genus Bufo have powerful toxins that are present in eggs, larvae, and adults. Native Australopapuan predators of anurans have no evolutionary history of exposure to these toxins and may be particularly vulnerable to their effects (Crossland and Alford 1998). The diet of Bufo melanostictus in Papua is currently being examined to determine whether native frogs are eaten, and whether they are competing with local species for food. In Manokwari high densities of the native ranid frog Platymantis papuensis persist in microsympatry with extremely high densities of Bufo melanostictus, suggesting that at least some species are not affected by its introduction (K. Krey, B. Tjatur-

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Table 7.8.1. Exotic herpetofauna1 documented from New Guinea Species

Distribution in New Guinea

Reference

Bufo marinus

PNG, widespread below ⬃1,500 m, not yet documented from Papua

Lever 2001

Bufo melanostictus

Populations established in Manokwari and Sorong areas, Papua Province

Menzies and Tapilatu 2000; Krey, Tjaturadi, and Richards, pers. obs. June 2005.

Limnonectes cancrivora Populations established in Jayapura, Menzies 1996; Manokwari, Sorong, and Sentani K. Krey, pers. obs. 2004 areas, Papua Province Cuora amboinensis

Escapees but probably no established populations, Jayapura, Papua Province

Tjaturadi, pers. obs.

Trachemys scripta

Individuals regularly encountered in wild, Jayapura and Manokwari, Papua Province

Tjaturadi, Krey, and Richards, pers. obs. August 2004, June 2005.

Lycodon aulicus

Population probably established at Sentani (Jayapura), Papua Province

Tjaturadi, pers. obs. 2005

Mabuya multifasciata

Population established at Sentani (Jayapura), Papua Province

Tjaturadi, pers. obs. 2005

Iguana iguana

Single escapee, no population established, Jayapura, Papua Province

Tjaturadi and Richards, pers. obs. August 2004

Excluding Limnonectes verruculosa (Menzies 1992).

1

adi, and S. Richards, pers. obs.). The diet of the large introduced frog Limnonectes cancrivora in Papua should be examined to determine whether it is eating native frogs. The active movement of exotic herpetofauna into Papua increases the risk of introducing virulent diseases to the island. For example, the American Bullfrog (Rana catesbeiana), is traded across Southeast Asia and is likely to be imported to Papua in the future. This species is a potential carrier of Chytridiomycosis, a fungal disease that has been implicated in widespread amphibian population declines (Daszak et al 2004). Its introduction to Papua would be catastrophic for local amphibian populations. It is clear that additional species of herpetofauna will be documented from Papua. For example, unconfirmed reports in 2005 indicate that the large and voracious Tokay Gecko (Gekko gecko), has recently been introduced to Sorong. Other species such as the House Gecko (Hemidactylus frenatus), that are not strictly

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considered ‘‘exotic’’ are spreading as human commensals to remote sites across Papua. Although some of these introductions are accidental, resulting from mass movement of cargo across the Indonesian archipelago, of equal concern is an active pet trade that imports exotic species and exports native frogs and reptiles to markets in western Indonesia.

Acknowledgments Fieldwork in Papua and preparation of this manuscript were supported by Conservation International.

Literature Cited Allen, G.R., K.G. Hortle, and S.J. Renyaan. 2000. Freshwater Fishes of the Timika Region, New Guinea. PT Freeport, Timika. Allen, G.R., H. Ohee, P. Boli, R. Bawole, and M. Warpur. 2002. Fishes of the Yongsu and Dabra areas, Papua, Indonesia. In Richards, S.J., and S. Suryadi (eds) A Biodiversity Assessment of Yongsu-Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, D.C. Conservation International 1999. The Irian Jaya Biodiversity Conservation PrioritySetting Workshop: Final Report. Conservation International, Washington, D.C. Crossland, M.R., and R.A. Alford. 1998. Evaluation of the toxicity of eggs, hatchlings and tadpoles of the introduced toad Bufo marinus (Anura: Bufonidae) to native Australian aquatic predators. Australian Journal of Ecology 23: 129–137. Daszak, P., A. Strieby, A.A. Cunningham, J.E. Longcore, C.C. Brown, and D. Porter. 2004. Experimental evidence that the bullfrog (Rana catesbeiana) is a potential carrier of Chytridiomycosis, an emerging fungal disease of amphibians. Herpetological Journal 14: 201–207. Ernst, C.H., and R.W. Barbour. 1989. Turtles of the World. Smithsonian Institution Press, Washingon, D.C. Lever, C. 2001. The Cane Toad: The History and Ecology of a Successful Colonist. Westbury Academic and Scientific Publishing, Otley. Menzies, J.I. 1992. Ecological and taxonomic notes on ranid frogs (Amphibia: Ranidae) from far western New Guinea. Science in New Guinea 18: 115–122. Menzies, J.I. 1996. 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. 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: 70–73. Mittermeier, R.A., C.G. Mittermeier, T.M Brooks, J.D. Pilgrim, W.R. Konstant, G.A.B. da Fonseca, and C. Kormos 2003. Wilderness and biodiversity conservation. Proceedings of the National Academy of Sciences 100: 10309–10313. Mittermeier, R.A., C.G. Mittermeier, J. Pilgrim, G. Fonseca, W.R. Konstant, and T.M. Brooks. 2002. Wilderness: Earth’s Last Wild Places. CEMEX, Mexico City. O’Dowd, D., P.T. Green, and P.S. Lake. 2003. Invasional ‘meltdown’ on an oceanic island. Ecology Letters 6: 812–817.

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Appendix 8.1. Glossary of botanical terms (compiled by Andrew J. Marshall) b e lo w a r e l i s t ed definitions of botanical terms and related ecological concepts used in the Flora section of this volume which may be unfamiliar to some readers. Terms are listed alphabetically. abaxial: describing the side of leaves, petals, etc. that face away from the stem or the main axis, i.e., the ventral (lower) side accrescent: gaining in girth or length with age or following fertilization, or growing together, as the calyx of some plants after flowering achene: a small, dry, hard, indehiscent, one-seeded fruit formed from a single carpel actinomorphic: describing the arrangement of parts in an organ or organism such that any cut taken through the center results in similar halves, i.e., radially symmetrical; this term is usually used in reference to flowers adaxial: describing the side of leaves, petals, etc. that face towards the stem or main axis, i.e., the dorsal (upper) surface adnate: united, fused adventitious root: a root that grows from somewhere other than the primary root, e.g., roots that arise from stems or leaves albumen: starchy and other nutritive material in a seed, stored as endosperm inside the embryo sac or as perisperm in the surrounding nucellar cells; any deposit of nutritive material accompanying the embryo alpha diversity: the diversity within a particular area or ecosystem, usually expressed as the number of species (i.e., species richness); the diversity at a single point alternate: not opposite or whorled, but placed singly at different heights on the stem; used to describe the arrangement of leaves or other parts alternipetalous: describing stamens borne on alternating radii with respect to petals anamorph: the asexual form of a fungus, characterized by the absence of sexual spores anatropous: an ovule that is inverted and straight, with the micropyle next to the hilum and the radicle consequently inferior androdioecious: a plant breeding system in which a species, to produce seeds, must have a male plant with flowers having only stamens and a bisexual plant with flowers having both stamens and pistils androecium: refers collectively to the stamens of one particular flower androphore: a stalk bearing the androecium angiosperm: a group of plants that produce seeds enclosed within an ovary, which may mature into a fruit; i.e., the flowering plants annular: in the form of a ring annulate: ringed or banded anther: the male part of a flower that produces pollen, usually on a filament; the anther and filament together are described as the stamen 1371

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anthesis: the action of opening a flower; the period of time during which a flower is open aperturate: containing one or more openings or apertures apex: the top part of a plant or structure, the growing point apical: 1. borne at the tip of an organ, farthest from the point of attachment, e.g., a bud which is located at the end of a stem; 2. describing the cells composing the apex of the leaf which are often broader and shorter than the cells of the middle of the leaf apiculum: a short, pointed, flexible tip apocarpous: with carpels separate rather than united apotropous: located under the funiculus arborescent: tree-like in size and growth habit aril: the outer covering, or sometimes merely an appendage, of a seed formed from the funicle; may be hard, waxy, or pulpy arillate: bearing an aril arillode: a false aril; an aril originating from the micropyle instead of from the funicle or chalaza of the ovule auricles: inrolled margins autotrophic: nutritionally independent of other organisms axil: the angle formed between a leaf stalk and the stem to which it is attached; in flowering plants, buds develop in the axils of leaves axile: belonging to, or found in, the axil axillary: growing from an axil ballistic: dispersal method originating from parent plant or diaspore; explosive dispersal mechanism barochory: unassisted dispersal, seeds drop to the ground close to or beneath the parent plant basal: 1. at or referring to the base of any structure; 2. describing cells at the base or insertion of the leaf, often of different shapes and colors from those of the main part of the leaf basifixed: attached by the base berry: a pulpy or fleshy fruit with numerous seeds embedded in the pulp beta diversity: the change in species diversity between ecosystems; the rate at which species accumulate as one moves away from a single point bilateral symmetry: the arrangement of parts in an organ or organism such that it can only be split into similar halves along a given plane, e.g., most leaves are bilaterally symmetrical along the midrib biseriate: arranged in two rows boreotropical: describing a mixed assemblage of plants, containing both temperate and tropical elements brachyblast: a short branch or shoot bract: any reduced or modified leaf associated with a cone or flower, often found at the base of a flower or flower cluster and sometimes forming a cup around the flower

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bracteole: possessing bracts bryophyte: a group of non-flowering plants generally lacking conducting tissues in which the gametophyte generation is the larger, persistent phase; bryophytes include the Hepaticophyta (liverworts), Anthocerotophyta (hornworts), and Bryophyta (mosses) buttresses: a flange of tissue protruding from the trunk of a tree, tapering outward at the base caduceus: falling off early calyptra: a lid or hood; in mosses, the thin veil or hood covering the mouth of the capsule calyptrate: having a calyptra calyx: a small whorl of modified leaves (sepals) at the base of a flower and, where present, enclosing the other parts of the flower in bud campanulate: bell shaped, cup-shaped with a broad base: applied to the shape of a flower campylotropous: describes an ovule or seed curved so as to bring the apex and base nearly together capitate: 1. shaped like a head; collected into a head or dense cluster; 2. terminated by a bulbous, swollen area carnose: fleshy carpel: female reproductive part of flower including ovary, style, and stigma, which may be solitary, grouped, or fused carpellate: bearing or consisting of carpels caruncle: a horny outgrowth near the hilum of a seed cataphylls: in cycads, a scale-like modified leaf which protects the developing true leaves; an outgrowth on a plant or animal such as a fowl’s wattle or a protuberance near the hilum of certain seeds caudate: having a slender tail-like appendage caudicle: the thread-like or strap-shaped stalk of a pollinium cauliflorous: flowering from the branches or trunk cauline: relating to or growing on a stem chamaechory: wind dispersal, where diaspore is rolled along the ground surface by wind ciliate: fringed with very fine hair-like filaments cincinnus: a monochasial cyme on which flowers appear in an order along a spiral circinate: 1. curved into a circle so that the apex is nearly or quite bent around to the leaf base; 2. coiled in a spiral, with the apex at the center; 3. coiled from the top downward, as the young frond of a fern clade: a group of organisms with common ancestors cladistics: a method of classification of organisms based on their common ancestry; exploring the evolutionary relationships between organisms or groups of organisms cladogram: a graphical representation of the evolutionary divergence of a clade from its common ancestor

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colleter: a group or tuft of secretory hairs, often found near the base of the leaf lamina and on the calyx in the families Apocynaceae and Asclepiadaceae coma: a tuft of leaves at the tip of a stem or branch compound: describing leaves with two or more leaflets attached to a single leaf stem conduplicate: folded together lengthwise connate: united; used to describe similar structures joined from the start connivent: coming close together or touching without joining; converging, but not fused copal: a resinous substance exuded from some species of tropical trees and hardening in air into a semi-transparent or transparent solid ranging in color from red to yellow or brown corolla: the collective group of petals which may occur separately or fused into a cup, tube, or other structure corolline corona: fleshy ridges or outgrowths of tissue attached to the corolla tube coronas: a crown of appendages between the corolla and stamens, or on the corolla or stamens corpusculum: the central part of a pollinarium, characteristic of the families Orchidaceae and Asclepiadaceae corymb: a flat-topped or convex open flower cluster, in which the outer flowers open first corymbose: in corymbs, or corymb-like cosmopolitan: having a world-wide distribution cotyledons: the first embryonic leaf or leaves produced by the embryo of a seed plant that serve to absorb nutrients packaged in the seed until the seedling is able to produce its first true leaves and begin photosynthesis; the number of cotyledons is a key feature for the identification of the two major groups of flowering plants cotyliform: shaped like a cotyle or a cup crenate: with notched edges; dentate with the teeth much rounded crenolate: with shallow-toothed edges crenulate: finely crenate crustaceous: hard and brittle in texture cryptocotylar: with the cotyledons remaining inside the seed; seed usually remaining below ground cryptogam: lower plants reproducing by spores rather than seeds, e.g., ferns, mosses, and fungi cupuliform: cup-shaped cyanobacteria: a division of microorganisms that are related to bacteria but are capable of photosynthesis cycad: any plant of the order Cycadales, consisting of palm-like, cone-bearing, evergreen tropical plants that reproduce by means of spermatozoids and have large pinnately compound, usually fan-shaped leaves

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cyme: an inflorescence with the oldest flower in the center of the cluster, i.e., with its central or terminal flowers blooming earliest cymose: bearing cymes, or cyme-like cystoliths: a nodule within a cell cavity consisting of calcium carbonate and occurring in plants such as figs deciduous: shedding periodically or annually or at maturity; often applied to the seasonal shedding of leaves decumbent: lying on the substrate with an ascending apex; reclining, but with the tip ascending decussate: alternating in pairs at right angles dehiscent: splitting or bursting open at maturity along defined lines, e.g., of a fruit depressed: flattened from above dextrorse: turned to the right diaspore: the reproductive portion of a plant (e.g., a seed or bud) that is dispersed and may give rise to a new plant dichogamy: the differing times of maturation of stamens and pistils in a flower dicotyledon: a large group of flowering plants with two cotyledons that initially emerge from the seed digitate: finger-like; compound, with the members arising together at the apex of the support dioecious: unisexual, with male and female flowers on separate plants discoid: resembling a disk distichous: in two vertical ranks domatia: a small structure located in the axils of the primary veins on the lower surface of leaves in some woody dicotyledons, usually consisting of depressions and being partly enclosed by leaf tissue or hairs dorsifixed: attached at the back drupe: a fleshy or pulpy fruit containing a single seed with the inner portion of the pericarp hard or stony duodichogamy: having three stages of flowers, which open directly after each other, with a certain degree of overlapping dyszoochory: animal dispersal, specifically where diaspore is eaten intentionally endemic: originating from and confined to a specific, usually small, geographic area endocarp: the inner layer of a pericarp endosperm: the mass of nutrient tissue formed within the embryo sac endozoochory: animal dispersal, where diaspore is eaten intentionally or unintentionally entire: with a continuous margin without teeth, lobes, or indentations entomophilous: pollinated by insects epicarp: the outer layer of the pericarp or matured ovary epigeal: germinating with cotyledons above the ground; of or relating to the emergence of cotyledons above the surface of the ground

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epimatium: expanded, fleshy, seed-bearing bract scale in certain conifers, especially some Podocarpaceae epipetalous: borne on or attached to the petals epiphyte: a plant which grows upon another plant, using the host plant for structural support epitropous: describing the adaxial curvature of an ovule with respect to the ovary axis epizoochory: adhesive dispersal by animals where diaspore is carried accidentally, such as by being caught in fur essential oil: any volatile plant oil used in perfume or flavorings exfoliate: to peel off or shed, as the thin layers of bark exocarp: the outer layer or skin of a pericarp exserted: projecting beyond an envelope, e.g., stamens from a corolla exstipulate: having no stipules extrastaminal: located outside the stamens extrorse: facing outward facies: the top to bottom appearance of a plant, including foliage, flowers, fruit, roots, etc. fascicle: a close bundle or cluster fasciculate: in close bundles or clusters frond: the leaves of ferns and some other cryptogams, usually having many divisions; frequently used to designate any fern-like or feather-like foliage funiculus: the stalk of the ovary fusiform: spindle-shaped; swollen in the middle and narrowing toward each end gall: an abnormal growth or swelling caused by insects, fungus, etc. gametophytic: haploidal or bearing one half the normal number of chromosomes gamma diversity: a measure of the overall diversity for the different ecosystems within a region gamopetalous: having the petals of the corolla more or less united gamophyllous: composed of coalescent leaves or leaf-like organs geocarpic: a plant whose seeds are produced below the ground geophyte: a plant whose perennial buds are found underground, usually attached to a bulb, corm, tuber, etc. glabrescent: becoming glabrous in age glabrous: smooth, without hairs or protrusions gland: a secreting cell or group of cells on or within a plant structure glaucous: bluish gray or bluish green often as a result of a wax-like bloom globose: spherical or spheroidal gymnosperm: a plant that produces bare (not enclosed) seeds; non-flowering plants gynodioecious: having both bisexual flowers and female flowers, but on separate plants gynoecium: a collective term for all female organs of a flower; the pistil or pistils considered as a group

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gynostegial corona: the collective term for the staminal and interstaminal coronas (in Asclepiadaceae), both of which are associated with the gynostegium gynostegium: the crown of united stamens in the family Asclepidaceae habit: the general growth pattern (form) of a plant; e.g., creeping, trees, shrubs, vines, etc. heliophytic: flourishing in sunlight hemiepiphyte: a plant that spends part but not all of its life as an epiphyte; seeds typically germinate in the canopy and send rooting branches towards the ground, once rooted and securely anchored in the ground the plant is independent of its original host hemitropous: turned half round; half inverted hilum: the scar on the seed coat comprising a corky abscission layer where the seed or ovule was attached to the funiculus holarctic: of or relating to a biogeographic region comprising the Nearctic and Palearctic regions combined, these two regions have been linked intermittently by the Bering land bridge holotype: one pressed herbarium specimen designated by the author as the plant on which the description and name are based hydrophyte: a plant adapted to growing in water, waterlogged soil, or on a substrate that becomes inundated on a regular basis hypanthium: a cup-like base of a flower to which the stamens, sepals, and petals are attached hypocotyls: the main axis of the seed embryo just below the cotyledons and continuing into the developing primary root of the seedling hypogeal: happening, living, or remaining below ground; describing the emergence of cotyledons below the surface of the ground or within the seed coat imbricate: with overlapping edges imparipinnate: having an uneven number of pinnae; lacking a terminal pinna inaperturate: without an aperture; lacking any pores of furrows, typically used in reference to pollen indumentum: a massing of fine hairs, glands, or prickles inflexed: bent inwards inflorescence: a flower cluster, including bracts, on a stalk infrafoliar: on the stem below the leaves infructescence: the grouping or arrangement of fruits integuments: a protective structure that develops from the base of an ovule and encloses it almost entirely except for an opening (the micropyle) at the tip of the nucellus interfoliar: on the stem between the leaves internode: the region of a stem between two nodes where there is no branching of the vascular tissue intramarginal: within and near the margin introrse: turned inward or toward the axis involucral: belonging to an involucre

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involucre: a circle or collection of bracts surrounding a flower cluster or head, or a single flower kerangas: heath forest key: a dry, one-seeded fruit with a wing labellum: a lip; the peculiar upper petal of the Orchidaceae laciniate: slashed; divided into narrow pointed lobes lamina: any broad and flattened region of a plant or alga, which allows for increased photosynthetic surface area laticifer: specialized cell or row of cells containing latex latrorse: directed laterally or sideways leaf: an organ found in most vascular plants consisting of a flat lamina (blade) and a petiole (stalk); many flowering plants also have a pair of small stipules near the base of the petiole leaflet: the individual blades in a compound leaf lectotype: a specimen selected from the original type material to serve as the nomenclatural type when a holotype was not originally designated, or is missing lepidote: beset with small scurfy scales liana: woody, climbing vine; also liane lichenicolous fungi: fungi that grows on fungi lignified: to become hard and woody; often used to describe herbaceous stems liguliform: tongue-shaped or strap-shaped lobate: having numerous lobes lobe: the partially rounded portion of a leaf or other organ; separated from the whole by a deep indentation that does not break the continuity of the structure lobed flower: a tubular or funnel-shaped flower opening into petal-like segments locule: a compartment or cavity of an ovary, anther, or fruit lomatorrhizal: laterally besides each other malpighiaceous: straight and oppressed but attached by the middle megasporangium: the female sporangium containing the megaspores megaspore: the female spore of a heterosporous plant megasporophylls: the leaf bearing the megasporangia mensuration: the measuring of geometric magnitudes, lengths, areas, and volumes mesocarp: the fleshy, middle portion of the wall of a succulent fruit between the skin and the stony layer mesophytic: growing in or adapted to a moderately moist environment meteoranemochory: wind dispersal where diaspore is blown by wind through the air micropyle: a small pore in a seed that allows water absorption; a minute opening in the wall of an ovule through which the pollen tube enters microsporangium: the male sporangium in which microspores are developed microsporophylls: the leaf bearing the microsporangi

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microtherm: a type of plant life that requires an annual mean temperature of 0–14C for normal growth mitriform: shaped like a miter or cap monoaxial: bearing a single axis monocarpellate: formed from one carpel monochasial: with branches on only one side monocotyledon: a large group of flowering plants with a single cotyledon that initially emerges from the seed monoecious: with both male and female reproductive organs on the same plant monophyletic: describing a group of organisms sharing and including a common ancestor, and including all the descendents of that common ancestor; a clade monopodial: describing the main axis of a stem or rhizome that maintains a single direction of growth and giving off lateral branches or stems mycelium: the mass of interwoven filamentous hyphae that forms especially the vegetative portion of the thallus of a fungus mycorrhiza: a symbiotic relationship between certain fungi and the roots of plants, including orchids and many types of tree; mycorrhiza can help the roots to absorb essential nutrients and in some species are essential to the normal growth of the plant myrmecochory: dispersal by ants where diaspore is carried intentionally myrmecophilous: refers to plants which have symbiotic relationships with ants nautohydrochory: water dispersal, where diaspore is carried either floating or submerged in fresh or saltwater currents nectary: a multicellular glandular structure secreting nectar found in flowers and on vegetative parts in some species neotropics: Central and South America, excluding the southern parts of Chile and Argentina, i.e., the New World tropics neotype: a specimen selected to serve as the taxonomic species type when the original type material is missing notorrhizal: above each other; referring to cotyledons nucellus: the central part of the ovule enclosing the female gametophyte obconate: conical with the attachment at the apex of the cone obconic: inversely conical, having the attachment at the apex obnapiform: inverse turnip-shaped oboval: obovate obovate: ovate but attached at the narrow end obturbinate: inverse top-shaped ombrohydrochory: water dispersal where diaspore is propelled by action of rain on plant structure or through wetting by rain or dew opposite: parts of plant arranged in pairs on opposite sides of the stem; used to describe the arrangement of leaves or other parts orthotropic: tending to grow or form along a vertical axis orthotropous: describes an ovule or seed that is erect, with the orifice or micropyle at the apex

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ovate: having an outline in the shape of an egg, with the stalk or point of attachment at the large end ovule: the outgrowth in the ovary that develops into a seed following fertilization of the egg cell within it ovuliferous: bearing ovules paleotropics: Africa, tropical Asia, New Guinea, and many Pacific islands, excluding Australia and New Zealand; i.e., the Old World tropics palmate: divided with radial lobes like the fingers of a hand palminerved: exhibiting palmate venation panicle: a branched flower cluster paniculate: borne in a panicle; resembling a panicle papilla: small, rounded protuberance from any plant surface paraphyletic: describing a group of organisms sharing and including a common ancestor, but not including all the descendents of that common ancestor parasitoids: an insect whose larvae live as parasites that eventually kill their hosts parenchyma: undifferentiated cellular tissue with thin cell walls, which may differentiate to form other types of cell paripinnate: having an even number of pinnae pedicel: the short stalk connecting a flower to the main stem of a plant or branches of its inflorescence pedicelled: borne on a pedicel peduncle: a primary flower stalk, supporting either a solitary flower or cluster of flowers pedunculate: borne upon a peduncle peltate: 1. describing a leaf attached to the petiole from near the center of the lower surface, and not at the margin; 2. shield-shaped pendent: hanging down, overhanging penninerved: having conspicuous lateral veins which are divergent from the midrib and approximately parallel to one another perianth: the sepals and petals of a flower pericarp: the wall of the matured ovary perigynous: adnate to the perianth, and therefore around the ovary and not at its base persistent: remaining attached instead of falling away perulate: describes leaf buds which are covered with scales petiolate: having a petiole petiole: the stalk of a leaf phanerocotylar: exposed cotyledons phanerogam: a term used in early classifications for a plant whose reproductive organs are easily visible as either flowers or cones; phanerogams are currently more commonly referred to as spermatophytes pinna: one of the primary divisions of a pinnate or compoundly pinnate frond or leaf; first order leaflets on a compound leaf

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pinnate: referring to a compound leaf with the leaflets along both sides of the leaf stalk pistil: the central set of organs in a flower; composed of one or more carpels pistillate: possessing pistils and without stamens pistillode: a sterile pistil, often underdeveloped placenta: any part of the interior of the ovary which bears ovules placentation: the arrangement of placentas within an ovary plagiotropic: having the longer axis inclined away from the vertical line planar: two-dimensional planoconvex: flat on one side and convex on the other pleurogram: a U-shaped line on both seed faces resulting from the modification of the face of the ovule during seed development pluriseriate: many-ranked, as applied to leaves arranged in several rows along the stem pneumatophore: a specialized root in certain aquatic plants which performs respiratory functions pollinarium: the male reproductive system of an orchid pollinia: a mass of fused pollen produced by many orchids polygamodioecious: describes a plant group which has bisexual and male flowers on some plants, and bisexual and female flowers on others polymorphic: having more than two distinct morphological variants polyphyletic: a group of organisms not including the most recent common ancestor, often because this common ancestor is not characteristic of the group or because the group originated from more than one evolutionary line polyploidy: with more than two sets of the basic chromosome number poricidal: a type of dehiscence in which the pollen is released through pores at the tip of the anther prismatic: describing the shape of a prism; angular, with flat sides, and of nearly uniform size throughout prophylls: a rudimentary leaf at the base of a leafy shoot protandrous: referring to a flower where the shedding of the pollen occurs before the stigma is receptive protogynous: referring to a flower where the shedding of the pollen occurs after the stigma has ceased to be receptive protogyny: development of female organs before male to avoid self-fertilization pseudostipules: the lower leaflets on pinnate leaf which resemble stipules pseudoverticillate: describing an arrangement where spirally-arranged leaves are congested at the end of each innovation and appear to form a whorl; i.e., pseudowhorled pseudowhorled: describing an arrangement where spirally-arranged leaves are congested at the end of each innovation and appear to form a whorl; i.e., pseudoverticillate pteridophyte: a plant in which the sporophyte generation is the larger phase and in

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which the gametophyte lives an existence independent of its parent sporophyte; the general name for the ferns and related genera ptyxis: describing the way in which a leaf is folded in the bud pubescent: covered with short, soft downy hairs pulvinate: cushion-like pulvinus: a swelling at the base of the stalk of a leaf or leaflet punctate: having translucent spots or depressions punctiform: having the form of a point quincuncial: the arrangement of five objects, with one on each of four corners and one in the center raceme: an elongated cluster of flowers on an unbranched inflorescence racemiform: referring to an inflorescence with the outward appearance of a raceme racemose: resembling a raceme rachis: 1. the main stalk of a flower cluster or the main leafstalk of a compound leaf; 2. the continuation of the stipe through a compound frond in ferns radial symmetry: describing an organism or organ which can be cut into equal halves along many axes passing through the center radicle: the end of a plant embryo which gives rise to the first, embryonic, root, i.e., the caudicle ramiflorous: bearing flowers directly from large branches and leafless twigs, but not on the trunk rays: radial strands of cells in wood and phloem reflexed: abruptly curved backwards; abruptly bent or turned downward reniform: describing an oval with the ends curved around in the same direction like a kidney; kidney-shaped resupinate: turned upside down rostrate: with a beak-like projection rotate: radiating horizontally like the spokes of a wheel ruminate: having a surface which is coarsely wrinkled, appearing as though chewed by a cow and then spit out, but not with the margins appearing gnawed sagittate: shaped like an arrow head with the two basal lobes pointing backwards salverform: having a slender tube which expands abruptly samara: a dry, one-seeded, indehiscent winged fruit sarcotesta: the fleshy outer layer of the seed coat in cycads saxicolous: growing among or upon rocks scalariform: having a ladder-like pattern scale: the outermost structures of a flower; a thin, membrane-like covering of the bud or twig base scandent: climbing sclerophyll: a woody plant with small, leathery, evergreen leaves that retain water and typically found in hot, dry climates sclerophyllous: having leaves stiffened by sclerenchyma sensu lato: in the broad sense (Latin) sensu stricto: in the strict or narrow sense (Latin)

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sepal: one of the separate perianth leaves of the calyx sepaloid: like a sepal septal: of or relating to a septum septate: divided by partitions septicidal: describing a capsule that dehisces through partitions and between cells septum: a partition or wall especially in an ovary serrate: having sharp teeth pointing forward serrulate: finely serrate sessile: stalkless; immobile simple: of one piece; not compound sinutate: wavy-edged sorediate: bearing soredia soredium: a patch of granular bodies on the surface of the thallus of lichens spermatophyte: any plant of the division Spermatophyta, the higher plants that produce seeds, including the gymnosperms and angiosperms spicate: arranged in or resembling a spike spike: a flower cluster in which sessile flowers grow along part of the length of the peduncle spiral: winding in a continuous pattern around an axis; used to describe the arrangement of leaves or other parts sporangium: a tiny sphere in which the spores are produced; often applied to the capsule, but by some authors restricted to the spore sac, or inner sac of the capsule containing the spores sporophytic: diploidal; i.e., bearing the full number of chromosomes stamen: a part of a flower, the tip of which produces pollen (the anther) staminode: a sterile stamen, or any structure without anther corresponding to a stamen stellate: star-shaped stigma: the sticky tip of a pistil stilt-roots: adventitious support roots stipules: paired appendages resembling small leaves found at the base of the leaves of many flowering plants style: the usually slender structure that connects the ovary and the stigma subligneous: describing a plant or plant part that has a texture and appearance of wood but somewhat less robust subopposite: almost opposite but one leaf or leaflet of each pair slightly above the other subtended: to be just below and close to, or enclosed in, its axil succulent: describing a plant having fleshy stems or leaves, often adapted to dry conditions suckers: shoots or small plants arising from the base of a larger plant suffruticesent: very low and woody; diminutively shrubby superior: with the flower parts growing from below the ovary supervolute: having a convolute arrangement in the bud

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supra-axillary: borne above the axil syconium: an urn-shaped receptacle lined with flowers and enclosed at the apex by involucral bracts; a fleshy, multiple fruit having a hollow center, e.g., a fig sympetalous: with petals joined at the margins or at the base, often forming a tube or funnel sympodial: describing the primary axis without a single, persistent growing point that develops from a series of lateral branches which change direction in succession and give it a zigzag form synandrium: with the androecium having the anthers connate syncarpous: with the carpels of the gynoecium united in a compound ovary syndromes: a group of characteristics that consistently occur together synzoochory: animal dispersal where diaspore is carried intentionally taproot: the main root of a plant, having a single, dominant axis and often serving the functions of structural support and food storage teliomorph: the sexual form of a fungus, possessing reproductive structures tepal: refers collectively to the sepals and petals of a flower when they are indistinguishable from each other terete: smooth, cylindrical, and tapering terminal: apical terricolous: living in, on, or near the ground testa: the outer protective seed-coat, commonly hard and brittle tetrads: grains organized in cohering groups of four; a group of four thallus: in cryptogams, a cellular expansion taking the place of stem and foliage and forming the main body fungi and lichens theca: a case, usually referring to the pollen sac in flowering plants or the capsule in bryophytes thrips: very small, thin insects that feed on sap and can cause damage to foliage and flowers and that pollinate some tropical forest plants thyrse: a contracted cylindrical or ovoid and usually compact panicle thyrsoid: resembling a thyrse transverse: across or at right angles to the vertical; cross-wise in position trioecious: having three sorts of flowers on the same or on different plants, some of the flowers being staminate, others pistillate, and others both staminate and pistillate truncate: ending abruptly, as if cut off transversely twiner: a climbing plant with no tendrils or suckers, in which the stem winds around other plants or objects for support ultramafic: ultra basic, containing very low silica content (less than 45%) umbel: a flower cluster in the shape of an umbrella; an inflorescence in which the peduncles or pedicels of a cluster spring from the same point umbellate: in or like an umbel umbelliform: in the shape of an umbel uniseriate: arranged in a single row united: fused together

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

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PS

PAGE 1384

Glossary of Botanical Terms / 1385

urceolate: hollow and cylindrical or ovoid, and contracted at or below the mouth, like an urn valvate: opening by valves, as a capsule; meeting by the edges without overlapping variegated: marked, striped, or blotched with some color in addition to the plant’s general overall color venation: the arrangement and pattern of veins in a leaf vernation: the arrangement of leaves in the bud versatile: describes an anther that is attached near the middle and turning freely on its support verticillate: arranged in a whorl viviparous: 1. producing young plants upon various parts of the parent plant through vegetative reproduction; 2. describing seeds or fruit which sprout before they fall from the parent plant whorl: an arrangement of appendages, such as branches or leaves, such that all are equally spaced around the stem at the same point whorled: not alternate or opposite but placed in a circle around the stem; used to describe the arrangement of leaves or other parts xeromorphic: describing structural features of plants adapted to dry conditions xerophyte: a plant adapted to dry conditions zoochory: the animal dispersal of fruits zygomorphic: bilateral symmetry, usually referring to flowers Sources: Definitions taken from the following sources, which provide more complete lists of botanical terms and additional relevant information. Bailey, J. (ed.). 1999. The Penguin Dictionary of Plant Sciences. 2nd ed. Penguin Books, London. Harris, J.G., and M.W. Harris. 2001. Plant Identification Terminology: An Illustrated Glossary. 2nd ed. Spring Lake Publisher, Spring Lake, New Jersey, U.S. Hickey, M., and C. King. 2000. The Cambridge Illustrated Glossary of Botanical Terms. Cambridge University Press, London. Mabberley, D.J. 1997. The Plant-Book: A Portable Dictionary of the Vascular Plants. 2nd ed. Cambridge University Press, London. van Balgooy, M.M.J. 1998. Malesian Seed Plants. Vol. 2. Rijksherbarium, Leiden, The Netherlands. Additional definitions and information were taken from the following web-based sources: The Garden Web Glossary of Botanical Terms http://glossary.gardenweb.com/glossary/ The Succulent Plant Page http://www.succulent-plant.com/plantglossary.html The UCMP Glossary of Botany http://www.ucmp.berkeley.edu/glossary/gloss8botany.html

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

APP1

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PS

PAGE 1385

Migratory to/from Australia? n n n

yes

resident resident resident resident resident resident resident resident resident

n n n n n n n n n

yes

resident resident resident

n n n

w w w w am

resident resident migrant resident vagrant resident resident resident vagrant resident resident migrant resident

y n y y y possibly possibly n n n y n n

maculosa

aim

resident

n

Aceros

[plicatus] plicatus

im

resident

n

Eurystomus

orientalis

w

resident

y

Family

Genus

Species

Casuariidae

Casuarius Casuarius Casuarius

bennetti casuarius unappendiculatus

Aepypodius Aepypodius Talegalla Talegalla Talegalla Megapodius Megapodius Megapodius Megapodius

arfakianus bruijnii cuvieri fuscirostris jobiensis [freycinet] reinwardt [freycinet] freycineti [freycinet] affinis wallacei

Phasianidae

Coturnix Coturnix Anurophasis

ypsilophora [chinensis]chinensis monorthonyx

ai w

Anatidae

Anseranas Dendrocygna Dendrocygna Dendrocygna Cygnus Tadorna Nettapus Salvadorina Anas Anas Anas Anas Aythya

semipalmata guttata eytoni [bicolor] arcuata atratus radjah pulchellus waigiuensis penelope gibberifrons superciliosa querquedula [nyroca] australis

a w a ai a ai ai

Turnicidae

Turnix

Bucerotidae Coraciidae

Megapodiidae

Endemic to Papua?

resident resident resident

Extralimital

Resident/migrant

Appendix 8.2. Provisional species list of birds occurring in Papua

a

yes yes

ai

i

yes

1386

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

APP2

03-15-07 07:29:47

PS

PAGE 1386

Provisional Species List of Birds Occurring in Papua / 1387 Alcedinidae

Alcedo Alcedo Alcedo Ceyx

[atthis] atthis [azurea] azurea [coerulescens] pusilla lepidus

w a aim w

resident resident resident resident

n n n n

Dacelonidae

Dacelo Dacelo Dacelo Clytoceyx Todiramphus Todiramphus Todiramphus Todiramphus Todiramphus Melidora Syma Syma Tanysiptera Tanysiptera Tanysiptera Tanysiptera Tanysiptera Tanysiptera Tanysiptera

leachii tyro gaudichaud rex nigrocyaneus macleayii [chloris] chloris saurophaga sanctus macrorrhina [torotoro] torotoro [torotoro] megarhyncha hydrocharis [galatea] galatea [galatea] ellioti [galatea] riedelii [galatea] carolinae [nympha] nympha sylvia

a

a

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n y n n y n n n n n n n n n y

Meropidae

Merops Merops

[superciliosus] philippinus ornatus

w a

resident migrant

n y

Cuculidae

Cuculus Cuculus Cacomantis Cacomantis Cacomantis Ramphomantis Chrysococcyx Chrysococcyx Chrysococcyx Chrysococcyx Chrysococcyx Caliechthrus Microdynamis Eudynamis Scythrops

saturatus pallidus variolosus castaneiventris flabelliformis megarhynchus [minutillus] russatus lucidus ruficollis meyeri osculans leucolophus parva [scolopacea] scolopacea novaehollandiae

w a ai a am

migrant vagrant resident resident resident resident resident migrant resident resident vagrant resident resident resident migrant

n y n n y n y y n n y n n y y

Centropus Centropus Centropus Centropus

menbeki [phasianinus] phasianinus [phasianinus] bernsteini [phasianinus] chalybeus

resident resident resident resident

n n n n

Centropodidae

a w i am a

i a

yes yes yes

yes

ai am

a

yes

w ai ai yes

(continued)

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

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03-15-07 07:29:47

PS

PAGE 1387

1388 / appendix 8. 2

Species

Chalcopsitta Chalcopsitta Chalcopsitta Eos Eos Pseudeos Trichoglossus Psitteuteles Lorius Charmosyna Charmosyna Charmosyna Charmosyna Charmosyna Charmosyna Charmosyna Oreopsittacus Neopsittacus Neopsittacus Probosciger Cacatua Cacatua Micropsitta Micropsitta Micropsitta Micropsitta Cyclopsitta Cyclopsitta Psittaculirostris Psittaculirostris Psittaculirostris Psittacella Psittacella Psittacella

[atra] atra [atra] duivenbodei sintillata [histrio] squamata [histrio] cyanogenia fuscata [haematodus] haematodus goldei [lory] lory multistriata wilhelminae rubronotata placentis pulchella josefinae papou arfaki musschenbroekii pullicauda aterrimus [sulphurea] galerita [sanguinea] sanguinea [pusio] keiensis [pusio] geelvinkiana [pusio] pusio bruijnii gulielmitertii diophthalma desmarestii edwardsii salvadorii brehmii picta modesta

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

APP2

03-15-07 07:29:48

yes

i

yes yes

aim

i

a a a yes m a

yes

PS

Migratory to/from Australia?

Genus

Psittacidae

Resident/migrant

Family

Endemic to Papua?

Extralimital

Appendix 8.2. (Continued)

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident migrant resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n n n n n n n possibly n n n n n n n n n n n n

PAGE 1388

Provisional Species List of Birds Occurring in Papua / 1389 Psittacella Geoffroyus Geoffroyus Tanygnathus Eclectus Psittrichas Alisterus Alisterus Aprosmictus Loriculus Apodidae

Collocalia Collocalia

madaraszi geoffroyi simplex megalorynchos roratus fulgidus [amboinensis] amboinensis [amboinensis] chloropterus [erythropterus] erythropterus [stigmatus] aurantiifrons

resident resident resident resident resident resident resident resident resident

n n n n n n n n n

resident

n

resident resident

n n

m

resident resident

n n n n n n

ai i ai

yes

i

yes

a

w

Collocalia Mearnsia Hirundapus Apus

[esculenta] esculenta [spodiopygius] hirundinacea [orientalis] nuditarsus [vanikorensis] vanikorensis papuensis novaeguineae [caudacutus] caudacutus [pacificus] pacificus

w w

resident resident resident migrant

Hemiprocnidae

Hemiprocne

mystacea

im

resident

n

Tytonidae

Tyto

[tenebricosa] multipunctata [novaehollandiae] novaehollandiae alba [capensis] longimembris

a

resident

n

a

resident

n

w w

resident resident

n n

i a ai im

resident resident resident resident

n n n n

Ninox Uroglaux

[magicus] magicus [strenua] rufa connivens [novaeseelandiae] novaeseelandiae theomacha dimorpha

resident resident

n n

Aegotheles Aegotheles Aegotheles Aegotheles Aegotheles Aegotheles

insignis cristatus bennetti wallacii [albertisi] albertisi [albertisi] archboldi

resident resident resident resident resident resident

n n n n n n

Podargus Podargus

ocellatus papuensis

resident resident

n n

Collocalia Collocalia

Tyto Tyto Tyto Strigidae

Aegothelidae

Podargidae

Otus Ninox Ninox Ninox

yes

a

am a

(continued)

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PS

PAGE 1389

1390 / appendix 8. 2

Species

Migratory to/from Australia?

Eurostopodus Eurostopodus Eurostopodus Eurostopodus

argus mystacalis papuensis archboldi

a a

yes

migrant migrant resident resident

n y n n

Caprimulgidae

Caprimulgus Caprimulgus

indicus [macrurus] macrurus

w w

yes

vagrant resident

n n

Columbidae

Columba Macropygia

[leucomela] vitiensis [amboinensis] amboinenesis [ruficeps] nigrirostris reinwardtsi indica stephani albifrons [striata] placida humeralis nicobarica rufigula jobiensis beccarii terrestis nobilis magnificus perlatus ornatus aurantiifrons wallacii [superbus] superbus coronulatus pulchellus regina [rivoli] rivoli [rivoli] solomonensis

im i

resident resident

n n

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident vagrant resident resident

n n n n n n n n n n n n n n n n n n n n n n n n

Macropygia Reinwardtoena Chalocphaps Chalcophaps Henicophaps Geopelia Geopelia Caloenas Gallicolumba Gallicolumba Gallicolumba Trugon Otidiphaps Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus Ptilinopus

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

APP2

03-15-07 07:29:49

Endemic to Papua?

Genus

Eurostopodidae

Extralimital

Family

Resident/migrant

Appendix 8.2. (Continued)

i w im a a w m m

a

i w

ai i

PS

yes

PAGE 1390

Provisional Species List of Birds Occurring in Papua / 1391 Ptilinopus Ptilinopus Ptilinopus Ducula Ducula Ducula Ducula Ducula Ducula Ducula Ducula Ducula Gymnophaps Goura Goura Goura

[viridis] viridis [hyogastra] iozonus naina concinna myristicivora [rufigaster] rufigaster chalconota pinon mullerii zoeae [bicolor] bicolor [bicolor] spilorrhoa albertisii [cristata] cristata [cristata] victoria [cristata] scheepmakeri

i

Otididae

Ardeotis

[kori] australis

a

Gruidae

Grus

rubicunda

a

resident

possibly

Rallidae

Rallina Rallina Rallina Rallina Rallina Gallirallus Gallirallus Lewinia Gymnocrex Amaurornis Porzana Porzana Porzana Eulabeornis Megacrex Porphyrio Gallinula Fulica

rubra [leucospila] leucospila [leucospila] forbesi [leucospila] mayri tricolor [torquatus] torquatus [philippensis] philippensis [pectoralis] pectoralis plumbeiventris [olivaceus] moluccanus [pusilla] pusilla [tabuensis] tabuensis cinerea castaneoventris inepta porphyrio [chloropus] tenebrosa [atra] atra

ai w w ai i a w w w a w ai w

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n y n y

Scolopax Gallinago Gallinago Limosa Limosa Numenius Numenius

saturata hardwickii megala [limosa] limosa lapponica [borealis] minutus phaeopus

i w w w w w w

resident migrant resident migrant migrant migrant migrant

n n n n n n n

Scolopacidae

i i

yes yes

w a i

yes

yes

yes

yes

yes

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n y n n n n

resident

probably not

(continued)

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APP2

03-15-07 07:29:50

PS

PAGE 1391

1392 / appendix 8. 2

Migratory to/from Australia? n n n n n n n n n n n n n n n n n n n n

w

resident

n

a w

resident resident

possibly n

a ai

migrant resident

y y

w w w w w w w

migrant migrant resident migrant migrant migrant migrant

n n n n n n n

Species

Numenius Tringa Tringa Tringa Tringa Tringa Tringa Tringa Tringa Tringa Arenaria Calidris Calidris Calidris Calidris Calidris Calidris Calidris Limicola Phalaropus

madagascariensis totanus stagnatilis nebularia ochropus glareola cinerea hypoleucos [incana] brevipes [incanca] incana interpres tenuirostris canutus alba [ruficollis] ruficollis [minutilla] subminuta acuminata ferruginea falcinellus lobatus

w w w w w w w w w w w w w w w w w w w w

Jacanidae

Irediparra

gallinacea

Burhinidae

Burhinus Burhinus

grallarius [recurvirostris] giganteus

Charadriidae

Haematopus Himantopus

[longirostris] longirostris [himantopus] leucocephalus [dominica] fulva squatarola dubius [alexandrinus] ruficapillus mongolus leschenaultii [asiaticus] veredus

Family

Pluvialis Pluvialis Charadrius Charadrius Charadrius Charadrius Charadrius

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

APP2

03-15-07 07:29:51

PS

Endemic to Papua?

migrant vagrant migrant migrant vagrant migrant migrant migrant migrant migrant migrant migrant migrant migrant migrant vagrant migrant migrant migrant migrant

Genus

Extralimital

Resident/migrant

Appendix 8.2. (Continued)

yes

yes

PAGE 1392

Provisional Species List of Birds Occurring in Papua / 1393 Erythrogonys Vanellus

cinctus miles

a a

migrant resident

y n

Glareolidae

Glareola Stiltia

[pratincola] maldivarum isabella

w a

resident migrant

n y

Laridae

Stercorarius Larus

pomarinus [novaehollandiae] novaehollanidae ridibundus nilotica caspia bengalensis bergii dougallii sumatrana hirundo [albifrons] albifrons anaethetus fuscata hybridus leucopterus stolidus [tenuirostris] minutus alba

w a

migrant migrant

n n

w w w w w w w w w w w w w w w w

vagrant migrant migrant resident resident resident resident resident migrant resident resident migrant migrant resident resident resident

n y y n n n n n n n n y n n n n

haliaetus [madagascariensis] subcristata [longicauda] longicauda alcinus [caeruleus] caeruleus [migrans] migrans sphenurus indus [leucogaster] leucogaster [aeruginosus] spilonotus [aeruginosus] approximans soloensis novaehollandiae fasciatus [rufitorques] melanochlamys [poliocephalus] poliocephalus [cirrocephalus] cirrocephalus meyerianus

w ai

resident resident

n n

w w w a w w w am w yes ai aim

resident resident resident resident resident resident resident resident migrant migrant resident resident resident

n n n n n n n n y n n n n

resident

n

resident

n

resident

n

Larus Sterna Sterna Sterna Sterna Sterna Sterna Sterna Sterna Sterna Sterna Chlidonias Chlidonias Anous Anous Gygis Accipitridae

Pandion Aviceda Henicopernis Macheiramphus Elanus Milvus Haliastur Haliastur Haliaeetus Circus Circus Accipiter Accipiter Accipiter Accipiter Accipiter Accipiter Accipiter

a

(continued)

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

APP2

03-15-07 07:29:51

PS

PAGE 1393

1394 / appendix 8. 2

Migratory to/from Australia? n n n n n y n

a a w a w

resident resident resident migrant resident

n y n y n

[ruficollis] novaehollandiae aim [ruficollis] ruficollis w

resident resident

y n

Phaethon Phaethon

rubricauda lepturus

w w

vagrant vagrant

n n

Sulidae

Sula Sula Sula

dactylatra sula leucogaster

w w w

resident resident migrant

n n n

Anhingidae

Anhinga

[melanogaster] novaehollandiae

w

resident

y

Phalacrocoracidae

Phalacrocorax Phalacrocorax

melanoleucos sulcirostris

aim ai

resident resident

Phalacrocorax

[carbo] carbo

w

vagrant

y y, extent unclear y

Egretta Egretta Egretta Ardea Ardea Ardea

novaehollandiae [garzetta] garzetta sacra pacifica [sumatrana] sumatrana picata

aim w w a w ai

migrant migrant resident vagrant resident resident

y y n y possibly y

Genus

Species

Erythrotriorchis Megatriorchis Butastur Harpyopsis Aquila Aquila Hieraaetus

[radiatus] buergersi doriae [indicus] indicus novaeguineae gurneyi audax morphnoides

Falconidae

Falco Falco Falco Falco Falco

berigora [tinnunculus] cenchroides [subbuteo] severus [subbuteo] longipennis [peregrinus] peregrinus

Podicipedidae

Tachybaptus Tachybaptus

Phaethontidae

Ardeidae

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

APP2

03-15-07 07:29:51

w

PS

Endemic to Papua?

a a

resident resident migrant resident resident resident resident

Family

Extralimital

Resident/migrant

Appendix 8.2. (Continued)

yes

PAGE 1394

Provisional Species List of Birds Occurring in Papua / 1395 Casmerodius Mesophoyx Bubulcus Butorides Nycticorax Zonerodius Ixobrychus Ixobrychus Ixobrychus

albus intermedia ibis [striatus] striatus [nycticorax] caledonicus heliosylus [minutus] sinensis [minutus] novaezelandiae flavicollis

w w w w aim w a w

resident migrant resident resident resident resident migrant migrant resident

y y y n y n n y n

Threskiornithidae

Plegadis Threskiornis Threskiornis Platalea

[falcinellus] falcinellus [aethiopicus] molucca spinicollis [leucorodia] regia

w ai a ai

migrant resident migrant migrant

y y y y

Pelicanidae

Pelecanus

conspicillatus

a

migrant

y

Ciconiidae

Ephippiorhynchu

sasiaticus

w

resident

n

Fregatidae

Fregata Fregata

minor ariel

w w

resident resident

n n

Procellariidae

Pseudobulweria Calonectris Oceanites

[rostrata] becki leucomelas oceanicus

w w w

vagrant vagrant vagrant

n n n

Pittidae

Pitta Pitta

w w

resident resident

n y

Pitta

[sordida] sordida [erythrogaster] erythrogaster [versicolor] versicolor

a

migrant

y

Climacteridae

Cormobates

[leucophaeus] placens

resident

n

Ptilonorhynchidae

Ailuroedus Ailuroedus Archboldia Amblyornis Amblyornis Amblyornis Sericulus Chlamydera Chlamydera

buccoides [crassirostris] melanotis [papuensis] papuensis inornatus macgregoriae flavifrons aureus lauterbachi cerviniventris

resident resident resident resident resident resident resident resident resident

n n n n n n n n n

Maluridae

Clytomyias Sipodotus Malurus Malurus Malurus

insignis wallacii [grayi] grayi alboscapulatus cyanocephalus

resident resident resident resident resident

n n n n n

Meliphagidae

Myzomela Myzomela Myzomela

eques obscura cruentata

resident resident resident

n n n

a yes yes

a

ai

(continued)

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

APP2

03-15-07 07:29:52

PS

PAGE 1395

1396 / appendix 8. 2

nigrita adolphinae [erythrocephala] erythrocephala rosenbergii fulvigula griseigula megarhynchus fallax [argentauris] argentauris [incana] indistincta [squamata] alboauricularis montana mimikae orientalis albonotata aruensis [analoga] analoga gracilis flavirictus subfrenatus obscurus [virescens] versicolor flaviventer polygramma chrysogenys albogularis ixoides cinereus stictocephalus meyeri [citreogularis] brassi [citreogularis] citreogularis

Myzomela Timeliopsis Timeliopsis Melilestes Glychichaera Lichmera Lichmera Lichmera Meliphaga Meliphaga Meliphaga Meliphaga Meliphaga Meliphaga Meliphaga Meliphaga Lichenostomus Lichenostomus Lichenostomus Xanthotis Xanthotis Oreornis Melithreptus Pycnopygius Pycnopygius Pycnopygius Philemon Philemon Philemon

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

APP2

03-15-07 07:29:53

a

a i a

yes

a

a a yes a

yes a

PS

Migratory to/from Australia?

Species

Myzomela Myzomela Myzomela

Resident/migrant

Genus

Endemic to Papua?

Family

Extralimital

Appendix 8.2. (Continued)

resident resident resident

n n n

resident resident resident resident resident resident resident resident

n n n n n n n n

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n n n n n n n

PAGE 1396

Provisional Species List of Birds Occurring in Papua / 1397 Philemon Philemon Ptiloprora Ptiloprora Ptiloprora Ptiloprora Ptiloprora Ptiloprora Melidectes Melidectes Melidectes Melidectes

Pardalotidae

Eopsaltriidae

Melidectes Melidectes Melidectes Melipotes Melipotes Ramsayornis Conopophila Entomyzon

[buceroides] novaeguineae corniculatus plumbea meekiana erythropleura [guisei] mayri [guisei] guisei perstriata fuscus [nouhuysi] nouhuysi ochromelas [leucostephes] leucostephes [leucostephes] belfordi [leucostephes] rufocrissalis torquatus [gymnops] gymnops [gymnops] fumigatus modestus albogularis cyanotis

Crateroscelis Crateroscelis Crateroscelis Sericornis Sericornis Sericornis Sericornis Sericornis Sericornis Sericornis Sericornis Acanthiza Gerygone Gerygone Gerygone Gerygone Gerygone Gerygone Gerygone

murina nigrorufa robusta [magnirostris] beccarii [magnirostris] virgatus [magnirostris] nouhuysi [rufescens] rufescens [rufescens] perspicillatus arfakianus papuensis spilodera murina cinerea chloronotus palpebrosa chrysogaster magnirostris [fusca] ruficollis [fusca] levigaster

Amalocichla Amalocichla Monachella Microeca Microeca

sclateriana incerta muelleriana [flavigaster] flavigaster griseoceps

a a

resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n

resident resident resident resident resident resident resident resident

n n n n n n n n

a

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n n n n n

a a

resident resident resident resident resident

n n n n n

yes

yes yes

yes a a a

a

yes

a a a

(continued)

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PAGE 1397

1398 / appendix 8. 2

Migratory to/from Australia? n n n n n n n n n n n n n n n n

a

resident resident resident

n n n

temminckii

a

resident

n

Pomatostomatiidae Pomatostomus Pomatostomus

isidori temporalis

a

resident resident

n n

Laniidae

Lanius

[cristatus] cristatus

w

yes

vagrant

n

Corvidae

Androphobus Cinclosoma Ptilorrhoa Ptilorrhoa Ptilorrhoa Ifrita Daphoenositta Daphoenositta Rhagologus

viridis ajax leucosticta caerulescens castanonota kowaldi chrysoptera miranda leucostigma

yes

resident resident resident resident resident resident resident resident resident

n n n n n n n n n

Orthonychidae

Genus

Species

Microeca Microeca Eugerygone Petroica Petroica Tregallasia Eopsaltria Poecilodryas Poecilodryas Poecilodryas Poecilodryas Peneothello Peneothello Peneothello Peneothello Heteromyias Pachycephalopsis Pachycephalopsis Drymodes

flavovirescens papuana rubra bivittata archboldi [capito] leucops pulverulenta brachyura [superciliosa] hypoleuca placens albonotata sigillatus cryptoleucus cyanus bimaculatus [albispecularis] albispecularis hattamensis poliosoma superciliaris

Orthonyx

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APP2

yes a a

yes

a

03-15-07 07:29:53

Endemic to Papua?

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

Family

Extralimital

Resident/migrant

Appendix 8.2. (Continued)

PS

PAGE 1398

Provisional Species List of Birds Occurring in Papua / 1399 Pachycare Aleadryas Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Pachycephala Colluricincla Colluricincla Colluricincla Pitohui Pitohui Pitohui Pitohui Pitohui Pitohui Eulacestoma Corvus Corvus Corvus Melampitta Melampitta Loboparadisea Cnemophilus Macgregoria Manucodia Manucodia Manucodia Manucodia Paradigalla Paradigalla Epimachus Epimachus Epimachus Epimachus Lophorina Parotia Parotia Ptiloris

flavogrisea rufinucha [grisola] phaionotus hyperythra meyeri [simplex] griseiceps [caledonica] pectoralis [caledonica] soror [caledonica] lorentzi [caledonica] melanura schlegelii aurea [rufiventris] monacha umbrina megarhyncha harmonica kirhocephalus dichrous incertus cristatus ferrugineus nigrescens nigropectus fuscicapillus tristis orru lugubris gigantea sericea loriae pulchra atra [chalybata] chalybata jobiensis keraudrenii [carunculata] carunculata [carunculata] brevicauda fastuosus meyeri [albertisi] albertisi [albertisi] bruijnii superba sefilata carolae [magnificus] magnificus

i

yes yes

a aim

a

a a

yes ai

a yes

yes a

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n (continued)

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PAGE 1399

1400 / appendix 8. 2

magnificus respublica regius nigra splendidissima alberti melanoleuca [apoda] rubra [apoda] minor [apoda] apoda mentalis [cassicus] cassicus quoyi tibicen [leucorynchus] leucorynchus [leucorynchus] maximus cinereus [blainvillii] blainvillii [blainvillii] montanus [sagittatus] szalayi [sagittatus] sagittatus flavocinctus [caledonica] novaehollandiae caeruleogrisea lineata boyeri [papuensis] papuensis longicauda [tenuirostris] tenuirostris [morio] incerta schisticeps melas

Artamus Artamus Peltops Peltops Oriolus Oriolus Oriolus Coracina Coracina Coracina Coracina Coracina Coracina Coracina Coracina Coracina Coracina

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03-15-07 07:29:55

yes yes

yes

a a a w

ai

a ai a

a a aim

PS

Migratory to/from Australia?

Species

Cicinnurus Cicinnurus Cicinnurus Astrapia Astrapia Pteridophora Seleucidis Paradisaea Paradisaea Paradisaea Cracticus Cracticus Cracticus Gymnorhina Artamus

Resident/migrant

Genus

Endemic to Papua?

Family

Extralimital

Appendix 8.2. (Continued)

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n n n n n n

resident vagrant resident resident resident migrant migrant migrant

n y n n n y n y

resident resident resident resident resident resident resident resident resident

n n n n n y n n n

PAGE 1400

Provisional Species List of Birds Occurring in Papua / 1401 Coracina Campochaera Lalage Lalage Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Rhipidura Chaetorhynchus Dicrurus Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Monarcha Arses

montana sloetii [aurea] atrovirens [aurea] leucomela leucophrys [rufiventris] rufiventris threnothorax maculipectus leucothorax atra [fuliginosa] hyperythra [fuliginosa] albolimbata [fuliginosa] phasiana brachyrhyncha [rufidorsa] rufidorsa [rufifrons] rufifrons papuensis [balicassius] bracteatus axillaris rubiensis cinerascens [melanopsis] frater [melanopsis] melanopsis guttulus [trivirgatus] trivirgatus [manadensis] julianae [manadensis] manadensis [manadensis] brehmii chrysomela [telescophthalmus] telescophthalmus Arses [telescophthalmus] insularis Myiagra atra Myiagra [rubecula] rubecula Myiagra ruficollis Myiagra cyanoleuca Myiagra inquieta Myiagra [alecto] alecto Machaerirhynchus flaviventer Machaerirhynchus nigripectus Grallina cyanoleuca Grallina bruijni Muscicapidae

Zoothera Turdus

[dauma] lunulata poliocephalus

a ai ai

a

aim w

i a a ai yes yes a

yes a ai a a ai a a a w

resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident resident migrant resident migrant resident resident resident resident resident

n n n n n n n n n n n n n n n n n n n n n y y n y n n n n n

resident

n

resident resident resident migrant resident resident resident resident resident resident

n y n y n n n n n n

resident resident

n n (continued)

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PAGE 1401

1402 / appendix 8. 2

Migratory to/from Australia? n n

resident resident resident resident resident resident resident

n n y n n n n

w w ai a

migrant resident migrant vagrant

n n y y

[ juncidis] juncidis exilis

w w

resident resident

n n

Zosterops Zosterops Zosterops Zosterops Zosterops

[chloris] chloris [atrifrons] minor mysorensis fuscicapillus novaeguineae

i

resident resident resident resident resident

n n n n n

Locustella Acrocephalus Acrocephalus Acrocephalus Phylloscopus Megalurus Megalurus Megalurus

[fasciolata] fasciolata [arundinaceus] orientalis [arundinaceus] stentoreus [arundinaceus] australis [trivirgatus] poliocephalus timoriensis albolimbatus gramineus

w w w a m w

migrant vagrant resident migrant resident resident resident resident

n n n n n n n n

Alaudidae

Mirafra

[ javanica] javanica

w

resident

n

Nectariniidae

Dicaeum Dicaeum

[erythrothorax] pectorale [erythrothorax]

resident resident

n n

Species

Muscicapa Saxicola

griseisticta caprata

w w

Aplonis Aplonis Aplonis Aplonis Aplonis Mino Mino

[cantoroides] cantoroides mysolensis [metallica] metallica [metallica] magna mystacea antais dumontii

Hirundinidae

Hirundo Hirundo Hirundo Hirundo

[rustica] rustica [rustica] tahitica nigricans [ariel] ariel

Cisticolidae

Cisticola Cisticola

Zosteropidae

Family

Sturnidae

Sylviidae

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i ai

Endemic to Papua?

migrant resident

Genus

Extralimital

Resident/migrant

Appendix 8.2. (Continued)

yes yes

yes yes

a

PS

yes

PAGE 1402

Provisional Species List of Birds Occurring in Papua / 1403

Nectarinia Nectarinia

geelvinkianum [hirundinaceum] hirundinaceum [sperata] aspasia jugularis

Melanochartidae

Melanocharis Melanocharis Melanocharis Melanocharis Melanocharis Melanocharis Toxorhamphus Toxorhamphus Toxorhamphus Oedistoma

arfakiana nigra longicauda versteri striativentris crassirostris novaeguineae poliopterus iliolophus pygmaeum

resident resident resident resident resident resident resident resident resident resident

n n n n n n n n n n

Paramythiidae

Oreocharis Paramythia

arfaki montium

resident resident

n n

Passeridae

Motacilla Motacilla Anthus

w w a

migrant migrant resident

n n n

Anthus Oreostruthus Neochima Passer

flava [cinerea] cinerea [novaeseelandiae] novaeseelandiae gutturalis fuliginosus phaetona montanus

n n n n

Erythrura Erythrura Lonchura Lonchura Lonchura Lonchura Lonchura Lonchura Lonchura Lonchura Lonchura Lonchura

[tricolor] trichroa papuana [tristissima] tristissima [tristissima] leucosticta grandis [caniceps] vana nevermanni spectabilis castaneothorax stygia teerinki montana

aim

resident resident resident vagrant exotic resident resident resident resident resident resident resident resident resident resident resident resident

Dicaeum

ai

yes

i w

w

yes

yes

a yes

resident

n

resident resident

n n

n n n n n n n n n n n n

Note: Abbreviations in Extralimital column indicate Australia (a), Moluccas/Indonesia (i), Polynesia/Melanesia (m), widespread (w). Source: Taxonomy follows Sibley and Monroe (1990).

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PAGE 1403

Appendix 8.3. The vertebrate fauna of the Fly and Purari Deltas and Bintuni Bay mangroves Fauna

Common name

Scientific name

Sea cow Irrawaddy dolphin Common water rat Lesser forest wallaby Rufescent bandicoot Papuan bandicoot Bush pig Dusky wallaby Sugar glider Spotted cuscus Common striped possum Tree kangaroo Common cuscus Bismarck flying fox Big-eared flying fox

Dugong dugon Orcaella brevirostris Hydromys chrysogaster Dorcopsis hageni Echymipera rufescens Peroryctes papuensis Sus scrofa pauensis Thylogale bruijni Petaurus breviceps Phalanger maculatus Dactylopsila trivirgata Dendrolagus sp. Phalanger orientalis Pteropus neohibernicus Pteropus macrotis

Black duck Australian darter Magpie goose Spotted tree duck Australian pelican Black cormorant Royal spoonbill Glossy ibis Black-necked stork Scheepmaker’s crown pigeon Dwarf cassowary Double wattled cassowary Common wildfowl Black-billed brush turkey Lesser yellow-billed kingfisher Racket-tailed kingfisher Australian paradise kingfisher Sulfur-crested cockatoo King bird of paradise Torres Strait pigeon Dusky lory Rainbow lory Barn owl Papuan hornbill White-throated pigeon

Anas superciliosa Anhinga rufa Anseranas semipalmata Dendrocygna guttata Pelecanus conspicillatus Phalacrocorax sulcirostris Platalea regia Plegadis falcinellus Xenorhynchus asiaticus Goura scheepmakeri Casuarius bennetti Casuarius casuarius Megapodius Freycinet Talegalla fuscirostris Halcyon torotoro Tanysiptera galatea Tanysiptera sylvia Cacatua galerita Cicinnurus regius Ducula spilorrhoa Pseudeos fuscata Trichoglossus haematodus Tyto alba Aceros plicatus Columba vitiensis

Mammals

Birds

1404

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PAGE 1404

Vertebrate Fauna of Fly and Purari Deltas and Bintuni Bay Mangroves / 1405 Purple-tailed imperial pigeon Black-belted fruit pigeon Black-shouldered fruit pigeon Collared fruit pigeon Bare-eyed crow New Guinea eagle Black-capped lory Glossy-mantled manucode Palm cockatoo Pesqueet’s parrot Raggiana bird of paradise Twelve-wire bird of paradise

Ducula rufigaster Ducula zoeae Ducula pinon Ducula mulleri Gymnocorvus tristis Harpyopsis novaguineae Lorius lory Manucodia ater Probosciger aterrimus Psittrichas fulgidus Paradisaea raggiana Seleucides melanoleuca

Javanese file snake Freshwater crocodile Saltwater crocodile Hawksbill turtle Green turtle Pitted-shell turtle Siebenrock’s snake-neck turtle New Guinea short-neck turtle Red short-neck turtle Short-shelled turtle Python Common monitor lizard

Acrochordus javanicus Crocodylus novaeguinea Crocodylus porosus Caretta caretta Chelonia mydas Carettachelys insculpta Cheodina siebenrocki Elseya novaguinea Emydura subglobosa Pelochelys bibroni Python sp. Varanus indicus

River whaler shark Pig-eyed whaler Whaler shark Hammerhead shark Spotted eagle-ray Long-tailed ray Cowtail ray Shovelnose ray Sawshark Indo-pacific Tarpon Long-horned anchovy Black-tipped sardinella Anchovy Freshwater anchovy Long-jawed anchovy Anchovy Indian Anchovy Deep-bodied herring Hamilton’s thryssa

Carcharhinus leucas Carcharhinus gangeticus Carcharhinus glyphis Sphyrna blochii Aetobatus narinari Himantura uar-nak Dasyatis sephen Rhizobatos armatus Pristis microdon Megolops cyprinoids Trissocles setirostris Sardinella melanura Scutengraulis hamiltoni Scutengraulis scratchleyi Stolephorus commersoni Stolephorus bataviensis Stolephorus indicus Kowala macrolepis Thryssa hamiltoni

Reptiles

Fish

(continued)

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PAGE 1405

1406 / a p p e n d i x n o. 8 . 3

Appendix 8.3. (Continued) Fauna

Common name

Scientific name

Blackspot herring Eel-tail catfish Goldspot catfish Broad-nosed catfish Lesser thick-lipped catfish Sharp-nosed catfish Grey catfish Threadfin catfish Ridge-backed catfish Yellow catfish High-nose catfish Long-faced catfish Granular-toothed catfish Long-ridged catfish Beach catfish Flesh-lipped catfish Spoon-snouted catfish Catfish Catfish Catfish Catfish Halfbeak Three-spined frogfish Freshwater moray eel Eel Snake eel Long tom Fly river garfish Short-nosed river garfish River garfish River garfish Deep-bodied flounder Large-toothed flounder Waite’s flounder Three-spot flounder Freshwater sole Sole Patterned tongue sole Tongue sole Tongue sole Spotted tongue sole Short-tailed pipefish Freshwater pipefish Banded freshwater pipefish

Macrura brevis Plotosus canius Hexanematichthys leptaspis Hexanematichthys latirostris Hexanematichthys carinatus Hexanematichthys danielsi Hexanematichthys proximus Hexanematichthys stirlingi Hexanematichthys australis Hexanematichthys sp. Cinetodus froggatti Pseudarius sp. Pseudarius colcloughi Nedystoma sp. Netuma sagoroides Hemipimelodus sp. Dioichthys novaguineae Arius armiger Arius leptaspis Euristhmus lepturus Euristhmus nudiceps Zenarchopterus buffonis Batrachomoeus trispinosus Gymnothorax polyuranodon Anguilla interioris Muraenichthys macropterus Strongylura strongylura Zenarchopterus novaguineae Zenarchopterus brevirostris Zenarchopterus buffonis Zenarchopterus dunckeri Pseudorhombus elevatus Pseudorhombus arsius Arnoglossus waitei Grammatobothus polyphthalmus Aseraggodes klunzingeri Aseraggodes sp. Paraplagusia bilineata Cynoglossus heterolepis Cynoglossus cynoglossus Cynoglossus punticeps Oostethus brachyurus Bombonia djarong Bombonia spicifer

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PAGE 1406

Vertebrate Fauna of Fly and Purari Deltas and Bintuni Bay Mangroves / 1407 Goldie river mullet Skip jack mullet Diamond-scale mullet Large-scaled mullet Brown-banded mullet Tade Grey mullet Grey mullet Mullet Sea mullet Mullet Mullet Blue-spotted mullet Troschel’s mullet Fourfinger threadfin Threadfin Nursery fish Trevally Great trevally Lowly trevally Shadow trevally Trevally Leatherskin Leatherskin Silver tooth ponyfish Black-tipped ponyfish Narrow-banded ponyfish Pug-nosed ponyfish Common ponyfish Ponyfish Pipe-fish Seven-banded cardinal-fish Yellow cardinal-fish Ocellated cardinal-fish Cardinal-fish Spotted flagtail Barramundi Crescent grunter-perch Giant perchlet Glassy perchlet Perchlet Perchlet Perchlet Perchlet Dusky tripletail Four-banded tripletail Papua black bass

Cestraeus goldiei Rhinomugil nasutus Liza vaigiensis Liza oligolepis Liza dussumieri Liza tade Liza subviridis Pseudomugil inconspicuus Mugil cephalus Mugil engeli Valamugil buchanani Valamugil seheli Liza macrolepis Eleutheronema tetradactylum Polydactylus sheridani Kurtus gulliveri Caranx ignobilis Caranx sexfasciatus Caranx ignobilis Carangoides dinema Carangoides sp. Chorinemus lysan Chorinemus sp. Gazza achlamys Leiognathus splendens Leiognathus equulus Secutor ruconis Equula equula Equulites berbis Hippichthys penicillus Apogon septemstriatus Apogon hartzfeldi Apogon carinatus Apogon hyalosoma Kuhlia marginata Lates calcarifer Terapon jarbua Paraambassis gulliveri Priopidichthys (Ambassis) gymnocephalus Ambassis nalua Ambassis macracanthus Ambassis interruptus Ambassis buruensis Lobotes surinamensis Datnioides quadrifasciatus Lutjanus sp. (continued)

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PAGE 1407

1408 / a p p e n d i x n o. 8 . 3

Appendix 8.3. (Continued) Fauna

Common name

Scientific name

Mangrove jack Moses perch Javelin-fish Yellow-finned javelin-fish White-finned javelin-fish Blotched javelin-fish Crescent perch Pikey bream Bony-bream Yellow-tipped threadfin-bream Sunrise goatfish Yellow-striped goatfish Goatfish Lumpy jewfish Green-backed jewfish Silver jewfish Drab jewfish Paper-headed jewfish Northern whiting Beach salmon Archer fish Estuary rock-cod Rock-cod Carpet rock-cod Yellow-lipped rock-cod Rock-cod Spotted sicklefish Spotless butterfly-cod Deep queenfish Spotted Scat Butterfish Spotted butterfish Flap-gilled dragonet Milk-spotted dragonet Dragonet Goby Goby Goby Goby Goby Goby Goby Goby Goby

Lutjanus argentimaculatus Lutjanus johnii Pomadasys hasta Pomadasys kaakan Pomadasys argenteus Pomadasys maculatum Therapon jarbua Acanthopagrus berda Anodontostoma chacunda Nemipterus nemaopus Upeneus sulphureus Upeneus vittatus Upeneus sp. Johnius belengeri Johnius dussumieri Pseudosciaena soldado Pseudosciaena sina Collichthys novaeguineae Sillago sihama Leptobrama mulleri Toxotes chatareus Epinephelus tauvina Epinephelus suillus Epinephelus fuscoguttatus Epinephelus amblycephalus Epinephelus suillus Drepane punctata Pterois russelli Scomberoides tala Scatophagus argus Selenotoca multifasciata Scatophagus argus Eleutherochir opercularis Callionymus sagitta Callionymus macdonaldi Oxyurichthys jaarmani Acentrogobius gracilis Acentrogobius janthinopterus Drombus sp. Mugilogobius tagala Pandaka lidwilli Parioglossus palustris Zappa confluentus Oxyurichthys tentacularis

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PAGE 1408

Vertebrate Fauna of Fly and Purari Deltas and Bintuni Bay Mangroves / 1409 Flat-headed goby Goby Bearded Goby Giant mudskipper Mudskipper Mudskipper Mudskipper Mudskipper Gudgeon Gudgeon Gudgeon Gudgeon Gudgeon Flying-gurnard Gudgeon Spangled gudgeon Green-backed gauvina Black-backed gauvina Loter Loter Purple-spotted gudgeon Bar-tailed flathead Flathead Silver toadfish Toadfish Toadfish Milk-spotted toadfish Reticulated toadfish Indian tripodfish Short-nosed tripodfish Long-nosed tripodfish Threadfin leatherjacket Heart-headed flathead Checkered lizard-fish Greater lizard-fish Lizard-fish Half-smooth golden pufferfish Round-headed pennant-fish Sharp-toothed hammer croaker Spotted croaker Blenny Blenny Java rabbit fish Long-spined porcupine-fish

Glossogobius giurus Glossobius circumspectus Scaetelaos viridis Periophythalmus schlosseri Periophthalmus weberi Periothalmus freycineti Periophthalmus koelreuteri Periophthalmus vulgaris Prionobutis microps Prionbutis koilomatodon Bostrichthys sinensis Butis butis Ophiocera aporos Dactylopena sp. Eutis amboinensis Ophiocara porocephala darwiniense Bunaka gyrinoides Bunaka herwerdeni Oxyeleotris urophthalmus novaeguineae Oxyeleotris lineolatus Mogurnda mogurnda Platycephalus indicus Suggrundus sp. Gastrophysus lunaris Torafuga pleurostictus Takifugu meraukensis Chelonodon patoca Arothron reticularis Triacanthus indicus Tricanthus biaculeatus Trixiphichthys weeri Paramonacanthus filicauda Sorsogona tuberculata Saurida undosquamis Saurida tumbil Saurida gracilis Lagocephalus spadiceus Alectis ciliaris Johnius vogleri Protonibea diacanthus Omobranchus ferox Omobranchus zebra Siganus javus Cyclichthys jaculiferus

Note: Only common species are listed. Sources: White and White (1976); Liem and Haines (1977); Bayley (1980); Pernetta and Burgin (1980); Cragg (1983); Liem (1983); Pernetta (1983); Ecology Team (1984); Erftemeijer (1989); Parenti and Allen (1991); Alongi et al. (1992).

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PAGE 1409

Appendix 8.4. Preliminary checklist of amphibians and reptiles reported from Papua and the Aru Islands Litoria rubella (Gray, 1842) Litoria sanguinolenta (van Kampen, 1909) • Litoria thesaurensis (Peters, 1878) Litoria umarensis Gu¨nther, 2004 • Litoria umbonata Tyler and Davies, 1983 • Litoria vagabunda (Peters and Doria, 1878) Litoria verae Gu¨nther, 2004 • Litoria wapogaensis Richards and Iskandar, 2001 • Litoria wisselensis (Tyler, 1968) • Litoria wollastoni (Boulenger, 1914) Nyctimystes fluviatilis Zweifel, 1958 • Nyctimystes granti (Boulenger, 1914) • Nyctimystes humeralis (Boulenger, 1912) Nyctimystes montanus (Peters and Doria, 1878) • Nyctimystes pulcher (Wandolleck, 1911)

FROGS Family Bufonidae Bufo melanostictus Schneider, 1799 * Family Myobatrachidae Crinia remota (Tyler and Parker, 1974) Lechriodus aganoposis Zweifel, 1972 Lechriodus melanopyga (Doria, 1874) Lechriodus platyceps Parker, 1940 • Limnodynastes convexiusculus (Macleay, 1878) Family Hylidae Litoria amboinensis (Horst, 1883) Litoria angiana (Boulenger, 1915) Litoria arfakiana (Peters and Doria, 1878) Litoria aruensis (Horst, 1883) • Litoria bicolor (Gray, 1842) Litoria brongersmai (Loveridge, 1945) • Litoria caerulea (White, 1790) Litoria chloronota (Boulenger, 1911) • Litoria congenita (Peters and Doria, 1878) Litoria dorsivena (Tyler, 1968) Litoria elkeae Gu¨nther and Richards, 2000 • Litoria eucnemis (Lo¨nnberg, 1900) Litoria genimaculata (Horst, 1883) Litoria graminea (Boulenger, 1905) Litoria infrafrenata (Gu¨nther, 1867) Litoria iris (Tyler, 1962) Litoria longicrus (Boulenger, 1911) Litoria macki Richards, 2001 • Litoria micromembrana (Tyler, 1968) Litoria modica (Tyler, 1968) Litoria multicolor Gu¨nther, 2004 • Litoria mystax (van Kampen, 1906) • Litoria napaea (Tyler, 1968) • Litoria nasuta (Gray, 1841) Litoria nigropunctata (Meyer, 1874) Litoria obtusirostris Meyer, 1874 • Litoria pratti (Boulenger, 1911) • Litoria pygmaea (Meyer, 1874) Litoria quadrilineata Tyler and Parker, 1974 • Litoria rothii (de Vis, 1884)

Family Microhylidae Albericus laurini Gu¨nther, 2000 • Albericus variegatus (van Kampen, 1923) • Asterophrys turpicola (Schlegel, 1837) Austrochaperina basipalmata (van Kampen, 1906) Austrochaperina blumi Zweifel, 2000 Austrochaperina derongo Zweifel, 2000 Austrochaperina kosarek Zweifel, 2000 • Austrochaperina macrorhyncha (van Kampen, 1906) • Callulops eurydactylus Zweifel, 1972 Callulops fuscus Peters, 1867 Callulops robustus (Boulenger, 1898) Choerophryne proboscidea van Kampen, 1914 Cophixalus balbus Gu¨nther, 2003 • Cophixalus biroi (Me´hely¨, 1901) Cophixalus tetzlaffi Gu¨nther, 2003 • Copiula expectata Gu¨nther, 2002 • Copiula major Gu¨nther, 2002 • Copiula obsti Gu¨nther, 2002 •

1410

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PAGE 1410

Preliminary List of Amphibians & Reptiles from Papua and Aru Islands / 1411 Hylophorbus nigrinus Gu¨nther, 2001 • Hylophorbus picoides Gu¨nther, 2001 • Hylophorbus rufescens Macleay, 1898 Hylophorbus sextus Gu¨nther, 2001 • Hylophorbus tetraphonus Gu¨nther, 2001 • Hylophorbus wondiwoi Gu¨nther, 2001 • Liophryne schlaginhaufeni (Wandolleck, 1911) Mantophryne lateralis Boulenger, 1887 Oreophryne albopunctata (van Kampen, 1909) • Oreophryne asplenicola Gu¨nther, 2003 • Oreophryne atrigularis Gu¨nther, Richards and Iskandar, 2001 • Oreophryne biroi (Me´hely¨, 1897) Oreophryne brevicrus Zweifel, 1956 Oreophryne clamata Gu¨nther, 2003 • Oreophryne crucifer (van Kampen, 1913) • Oreophryne flava Parker, 1934 • Oreophryne idenburgensis Zweifel, 1956 • Oreophryne kapisa Gu¨nther, 2003 • Oreophryne minuta Richards and Iskandar, 2000 • Oreophryne pseudasplenicola Gu¨nther, 2003 • Oreophryne sibilans Gu¨nther, 2003 • Oreophryne unicolor Gu¨nther, 2003 • Oreophryne waira Gu¨nther, 2003 • Oreophryne wapoga Gu¨nther, Richards and Iskandar, 2001 • Oxydactyla brevicrus van Kampen, 1913 • Sphenophryne cornuta Peters and Doria, 1878 Xenobatrachus anorbis Blum and Menzies, 1988 Xenobatrachus arfakianus Blum and Menzies, 1988 • Xenobatrachus bidens (van Kampen, 1909) Xenobatrachus giganteus (van Kampen, 1915) • Xenobatrachus macrops (van Kampen, 1909) •

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

Xenobatrachus multisica Blum and Menzies, 1988 • Xenobatrachus obesus Zweifel, 1960 Xenobatrachus ocellatus (van Kampen, 1913) • Xenobatrachus ophiodon Peters and Doria, 1878 • Xenobatrachus rostratus (Me´hely¨, 1898) Xenobatrachus scheepstrai Blum and Menzies, 1988 • Xenobatrachus schiefenhoeveli Blum and Menzies, 1988 • Xenorhina adisca Kraus and Allison, 2003 • Xenorhina bouwensi (de Witte, 1930) • Xenorhina eiponis Blum and Menzies, 1988 • Xenorhina minima (Parker, 1934) • Xenorhina oxycephala (Schlegel, 1858) Xenorhina parkerorum Zweifel, 1972 Xenorhina similis (Zweifel, 1956) Family Ranidae Limnonectes cancrivorus (Gravenhorst, 1829) * Limnonectes grunniens (Daudin, 1802) Platymantis batantae Zweifel, 1969 • Platymantis bimaculata Gu¨nther, 1999 • Platymantis cheesmanae Parker, 1940 • Platymantis cryptotis Gu¨nther, 1999 • Platymantis papuensis Meyer, 1874 Platymantis punctata Peters and Doria, 1878 • Rana arfaki Meyer, 1874 Rana aurata Gu¨nther, 2003 • Rana daemeli Steindachner, 1868 Rana garritor Menzies, 1987 Rana grisea van Kampen, 1913 Rana jimiensis Tyler, 1963 Rana novaeguineae van Kampen, 1909 Rana papua Lesson, 1826 Rana supragrisea Menzies, 1987 Rana volkerjane Gu¨nther, 2003 • TURTLES Family Chelidae Chelodina novaeguineae Boulenger, 1888 Chelodina reimanni Philippen and Grossmann, 1990 •

APP4

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PS

PAGE 1411

1412 / appendix 8 .4 Chelodina rugosa Ogilby, 1890 Elseya branderhorsti (Ouwens, 1914) Elseya novaeguineae (Meyer, 1874) Emydura subglobosa (Krefft, 1876) Family Cheloniidae Caretta caretta (Linnaeus, 1766) Chelonia mydas (Linnaeus, 1766) Eretmochelys imbricata (Linnaeus, 1766) Lepidochelys olivacea (Eschscholtz, 1829) Natator depressus (Garman, 1880) Family Dermochelyidae Dermochelys coriacea Linnaeus, 1766 Family Carettochelidae Carettochelys insculpta Ramsay, 1886 Family Trionychidae Pelochelys bibroni Owen, 1853 Pelochelys signifera Webb, 2002 CROCODILES Family Crocodylidae Crocodylus novaeguineae Schmidt, 1928 Crocodylus porosus Schneider, 1801 LIZARDS Family Agamidae Chlamydosaurus kingii Gray, 1825 Diporiphora australis (Steindachner, 1867) Diporiphora bilineata Gray, 1842 Hydrosaurus amboinensis (Schlosser, 1768) Hypsilurus auritus (Meyer, 1874) Hypsilurus binotatus (Meyer, 1874) • Hypsilurus bruijnii (Peters and Doria, 1878) • Hypsilurus dilophus (Dume´ril and Bibron, 1837) Hypsilurus geelvinkianus (Peters and Doria, 1878) • Hypsilurus modestus (Meyer, 1874) Hypsilurus nigrigularis (Meyer, 1874) • Hypsilurus schultzwestrumi (Urban, 1999) Lophognathus temporalis (Gu¨nther, 1867) Physignathus lesueurii (Gray, 1831)

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

APP4

Family Gekkonidae Cosymbotus platyurus (Schneider, 1792) Cyrtodactylus aaroni Gu¨nther and Ro¨sler, 2002 • Cyrtodactylus irianjayaensis Ro¨ssler, 2001 • Cyrtodactylus marmoratus Gray, 1831 Cyrtodactylus mimikanus (Boulenger, 1914) Cyrtodactylus novaeguineae (Schlegel, 1844) Cyrtodactylus papuensis (Brongersma, 1934) Cyrtodactylus sermowaiensis (de Rooij, 1915) Gehyra dubia (Macleay, 1877) Gehyra interstitialis Oudemans, 1894 Gehyra leopoldi Brongersma, 1930 • Gehyra marginata Boulenger, 1887 Gehyra mutilata (Wiegmann, 1835) Gehyra oceanica (Lesson, 1826) Gehyra papuana Meyer, 1874 Gehyra vorax Girard, 1857 Gekko gecko (Linnaeus, 1758) * Gekko monarchus (Dume´ril and Bibron, 1836) Gekko vittatus (Houttuyn, 1782) Hemidactylus frenatus (Dume´ril and Bibron, 1836) Hemidactylus garnotii (Dume´ril and Bibron, 1836) Hemiphyllodactylus typus Bleeker, 1860 Lepidodactylus lugubris (Dume´ril and Bibron, 1836) Lepidodactylus novaeguineae Brown and Parker, 1977 Nactus ‘‘pelagicus’’ Nactus vankampeni (Brongersma, 1933) Family Pygopodidae Lialis burtonis Gray, 1834 Lialis jicari Boulenger, 1903 Family Dibamidae Dibamus novaeguineae Dume´ril and Bibron, 1839

03-15-07 07:29:59

PS

PAGE 1412

Preliminary List of Amphibians & Reptiles from Papua and Aru Islands / 1413 Family Scincidae Carlia digulensis (Kopstein, 1926) • Carlia fusca (Dume´ril and Bibron, 1839) • Carlia longipes (Macleay, 1877) Carlia pulla (Barbour, 1911) Cryptoblepharus aruensis Mertens, 1928 • Cryptoblepharus litoralis (Mertens, 1958) Cryptoblepharus novaeguineae Mertens, 1928 Cryptoblepharus pallidus Mertens, 1928 Ctenotus robustus Storr, 1970 Ctenotus spaldingi (Macleay, 1877) Emoia aenea Brown and Parker, 1985 Emoia atrocostata (Lesson, 1826) Emoia aurulenta Brown and Parker, 1985 Emoia battersbyi (Proctor, 1923) Emoia baudini (Dume´ril and Bibron, 1839) • Emoia bogerti Brown, 1953 • Emoia brongersmai Brown, 1991 Emoia caeruleocauda (de Vis, 1892) Emoia callisticta (Peters and Doria, 1878) Emoia cyclops Brown, 1991 • Emoia digul Brown, 1991 • Emoia irianensis Brown, 1991 • Emoia jakati (Kopstein, 1926) Emoia jamur Brown, 1991 • Emoia klossi (Boulenger, 1914) Emoia kordoana (Meyer, 1874) Emoia longicauda (Macleay, 1877) Emoia loveridgei Brown, 1953 Emoia maxima Brown, 1953 Emoia obscura (de Jong, 1927) Emoia oribata Brown, 1991 Emoia pallidiceps de Vis, 1890 Emoia paniai Brown, 1991 • Emoia physicina Brown and Parker, 1985 Emoia pseudopallidiceps Brown, 1991 Emoia reimschisseli Tanner, 1950 Emoia sorex (Boettger, 1895) Emoia tropidolepis (Boulenger, 1914) Emoia veracunda Brown, 1953 Eugongylus rufescens (Shaw, 1802) Eugongylus unilineatus (de Rooij, 1915) Glaphyromorphus crassicaudis (Dume´ril and Dume´ril, 1851)

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

Glaphyromorphus nigricaudis (Macleay, 1877) Lamprolepis smaragdina (Lesson, 1826) Lipinia cheesmanae (Parker, 1940) Lipinia longiceps (Boulenger, 1895) Lipinia noctua (Lesson, 1826) Lipinia nototaenia (Boulenger, 1914) • Lipinia occidentalis Gu¨nther, 2000 • Lipinia pulchra (Boulenger, 1903) Lipinia septentrionalis Gu¨nther, 2000 Lipinia venemai Brongersma, 1953 • Lobulia brongersmai (Zweifel, 1972) Lobulia elegans (Boulenger, 1897) Lygisaurus macfarlani (Gu¨nther, 1877) Lygisaurus novaeguineae (Meyer, 1874) Mabuya multifasciata (Kuhl, 1820) Papuascincus morokanus (Parker, 1936) Papuascincus stanleyanus (Boulenger, 1897) Prasinohaema flavipes (Parker, 1936) Prasinohaema parkeri (Smith, 1937) • Prasinohaema prehensicauda (Loveridge, 1945) Prasinohaema semoni (Oudemans, 1894) Prasinohaema virens (Peters, 1881) Sphenomorphus aruensis (Doria, 1874) • Sphenomorphus derooyae (de Jong, 1927) Sphenomorphus fuscolineatus Greer and Shea, 2004 Sphenomorphus jobiensis (Meyer, 1874) Sphenomorphus latifasciatus (Meyer, 1874) Sphenomorphus longicaudatus (de Rooij, 1915) Sphenomorphus maindroni (Sauvage, 1878) • Sphenomorphus meyeri (Doria, 1874) Sphenomorphus mimikanus (Boulenger, 1914) • Sphenomorphus minutus (Meyer, 1874) Sphenomorphus muelleri (Schlegel, 1837) Sphenomorphus nigriventris (de Rooij, 1915) Sphenomorphus oligolepis (Boulenger, 1914) Sphenomorphus pratti (Boulenger, 1903)

APP4

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PS

PAGE 1413

1414 / appendix 8 .4 Sphenomorphus rufus (Boulenger, 1887) • Sphenomorphus schultzei (Vogt, 1911) Sphenomorphus simus (Sauvage, 1879) Sphenomorphus solomonis (Boulenger, 1887) Sphenomorphus undulatus (Peters and Doria, 1878) Sphenomorphus wollastoni (Boulenger, 1914) • Tiliqua gigas (Schneider, 1801) Tribolonotus gracilis de Rooij, 1909 Tribolonotus novaeguineae (Schlegel, 1834)

Family Cylindrophiidae Cylindrophis aruensis Boulenger, 1920 • Family Pythonidae Apodora papuana (Peters and Doria, 1878) Leiopython albertisii (Peters and Doria, 1878) Morelia amethistina (Schneider, 1801) Morelia boeleni (Brongersma, 1953) Morelia spilota (Lace´pe`de, 1804) Morelia viridis (Schlegel, 1872) Family Boidae Candoia aspera (Gu¨nther, 1877) Candoia carinata (Schneider, 1801)

Family Varanidae Varanus beccarii (Doria, 1874) • Varanus boehmei Jacobs, 2003 • Varanus doreanus (Meyer, 1874) Varanus indicus (Daudin, 1802) Varanus jobiensis Ahl, 1932 Varanus kordensis (Meyer, 1874) • Varanus macraei Bo¨hme and Jacobs, 2001 • Varanus panoptes Storr, 1980 Varanus prasinus (Schlegel, 1839) Varanus reisingeri Eidenmu¨ller and Wicker, 2005 • Varanus salvadorii (Peters and Doria, 1878) Varanus similis Mertens, 1958

Family Acrochordidae Acrochordus arafurae McDowell, 1979 Acrochordus granulatus (Schneider, 1799)

SNAKES Family Typhlopidae Ramphotyphlops braminus (Daudin, 1803) Ramphotyphlops erycinus (Werner, 1901) Ramphotyphlops flaviventer (W. Peters, 1865) • Ramphotyphlops multilineatus (Schlegel, 1839) Ramphotyphlops olivaceus (Gray, 1845) Ramphotyphlops polygrammicus (Schlegel, 1839) Ramphotyphlops similis (Brongersma, 1934) • Ramphotyphlops supranasalis (Brongersma, 1934) •

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Typhlops ater Schlegel, 1839 Typhlops diardii Schlegel, 1839

APP4

Family Colubridae Boiga irregularis (Merrem, 1802) Brachyorrhos albus (Linnaeus, 1758) Brachyorrhos jobiensis (Meyer, 1875) • Cantoria annulata de Jong, 1927 Cerberus rynchops (Schneider, 1799) Dendrelaphis calligastra (Gu¨nther, 1867) Dendrelaphis gastrostictus (Boulenger, 1894) Dendrelaphis lorentzi (van Lidth de Jeude, 1911) Dendrelaphis pictus (Gmelin, 1789) Dendrelaphis punctulatus (Gray, 1827) Enhydris polylepis (Fischer, 1886) Fordonia leucobalia (Schlegel, 1837) Heurnia ventromaculata Jong, 1926 • Myron richardsoni Gray, 1849 Stegonotus cucullatus (Dume´ril, Bibron and Dume´ril, 1854) Stegonotus diehli Lindholm, 1905 Stegonotus modestus (Schlegel, 1837) Stegonotus parvus (Meyer, 1875) Tropidonophis doriae (Boulenger, 1897) Tropidonophis elongatus (Jan, 1865) Tropidonophis mairii (Gray, 1841)

03-15-07 07:30:00

PS

PAGE 1414

Preliminary List of Amphibians & Reptiles from Papua and Aru Islands / 1415 Tropidonophis mcdowelli Malnate and Underwood, 1988 Tropidonophis montanus (van Lidth de Jeude, 1911) • Tropidonophis multiscutellatus (Brongersma, 1948) Tropidonophis novaeguineae (van Lidth de Jeude, 1911) Tropidonophis picturatus (Schlegel, 1837) Tropidonophis statisticus Malnate and Underwood, 1988 Tropidonophis truncatus (Peters, 1863) Family Elapidae Subfamily Elapinae Acanthophis laevis Macleay, 1877 Acanthophis rugosus Loveridge, 1948 Aspidomorphus muelleri (Schlegel, 1837) Aspidomorphus schlegelii (Gu¨nther, 1872) Demansia vestigiata (de Vis, 1884) Furina tristis (Gu¨nther, 1858) Micropechis ikaheka (Lesson, 1826) Oxyuranus scutellatus (Peters, 1867) Pseudechis papuanus Peters and Doria, 1878 Pseudechis rossignolii (Hoser, 2000) Pseudonaja textilis (Dume´ril, Bibron and Dume´ril, 1854)

Rhinoplocephalus boschmai (Brongersma and Knaap van Meeuven, 1961) Toxicocalamus grandis (Boulenger, 1914) • Toxicocalamus loriae (Boulenger, 1898) Toxicocalamus preussi (Sternfeld, 1913) Toxicocalamus stanleyanus Boulenger, 1903 Subfamily Laticaudinae Laticauda colubrina (Schneider, 1799) Laticauda laticauda (Linnaeus, 1758) Subfamily Hydrophiinae Acalyptophis peronii (Dume´ril, 1853) Aipysurus eydouxi (Gray, 1849) Aipysurus laevis Lace´pe`de, 1804 Astrotia stokesii (Gray, 1846) Disteira major (Shaw, 1802) Emydocephalus annulatus Krefft, 1869 Enhydrina schistosa (Daudin, 1803) Hydrelaps darwiniensis Boulenger, 1896 Hydrophis atriceps Gu¨nther, 1864 Hydrophis belcheri (Gray, 1849) Hydrophis elegans (Gray, 1842) Hydrophis ornatus (Gray, 1842) Hydrophis pacificus Boulenger, 1896 Lapemis hardwickii Gray, 1834 Parahydrophis mertoni (Roux, 1910) Pelamis platurus Linnaeus, 1766

Note: * indicates introduced species, • indicates endemic species. Checklist is current to September 2005; updates are available at www.bishopmuseum.org/research/pbs/pngherps/.

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APP4

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PS

PAGE 1415

1416

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

APP5

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PS

PAGE 1416

Specific name Charhinus amboinensis Carcharhinus leucas Glyphis sp. C Pristis microdon Himantura chaophraya Scleropages jardinii Megalops cyprinoides Moringua penni Anguilla bicolor Clupeoides papuensis Nematalosa erebi Nematalosa flyensis Nematalos papuensis Thryssa rastrosa Thryssa scratchleyi Arius augustus Arius berneyi Arius danielsi

Family

Carcharhinidae

Pristidae

Dasyatidae

Osteoglossidae

Megalopidae

Moringuidae

Anguillidae

Clupeidae

Engraulidae

Ariidae

Short-barbelled Catfish Berney’s Catfish Daniel’s Catfish

Fly River Thryssa Freshwater Anchovy

Toothed River-herring Freshwater Herring Fly River Herring Strickland River Herring

Indian Short-finned Eel

River Worm-eel

Oxeye Herring or Tarpon

Saratoga

Freshwater Whipray

Largetooth Sawfish

Pigeye Shark Bull Shark New Guinea River Shark

Fly River Fly River Fly River

Fly River Bensbach, Oriomo, Fly rivers

Digul, Fly rivers Bensbach River Fly River Fly River

Daru

Fly River

Digul, Bensbach, Fly, Aramia rivers

Digul, Fly, Bensbach rivers

Fly-Strickland River (Lake Murray)

Middle Fly River

Bensbach River Bensbach River Fly River

Common name Locality Native Species

Appendix 8.5. Fish records in the Trans-Fly

2,13,19 14,19 2,13,19

2,13,19 2,6,7,9,11

2,13,19,32 6,32 2,13,19 2,19

2,32

13,19,32

2,6,7,9,13,19

2,6,11,13,19

4

6,11,13,19

6,11 2,6,11 3

Source (recent period 1975–2004)

Source (early Dutch period 1900–1955)

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APP5

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PS

Spotted Blue-eye Swamp Blue-eye Delicate Blue-eye

Pseudomugil paludicola Pseudomugil tenellus

2,7,9 2,32

2,6,7,9,13,19,32

2,6,7,9,11

8,9,19 2,6,12,19 2,6,13,19,32 13 2,13,32 2,7,9,13,19 2,7,9 2,13,19

Bensbach, Fly, Aramia rivers

Bensbach, Pahoturi, Binaturi, Fly rivers Binaturi, Pahoturi, Morehead rivers Bensbach River

2,6,32

2,32

2,6,7,9,13,19,32

2 2,6,32 2,6,7,9,13,19,32

2,6,7,9,13,19,32

2,6,7,9,11,13,19

Merauke River and Balimo area 2 Fly River 2,7,9,13,19

Digul, Bensbach, Fly, Aramia rivers Merauke, Bensbach, Fly, Aramia rivers Oriomo, Fly rivers Fly River

Fly River Fly, Bensbach rivers Bensbach, Oriomo, Fly rivers Fly River Fly River Fly, Digul rivers Aramia River Fly River

Merauke, Bensbach, Morehead, Pahoturi, Fly rivers Goldie River Rainbowfish Merauke River Macculloch’s Rainbowfish Bensbach River Red-striped Rainbowfish Merauke, Bensbach, Morehead, Fly, Aramia rivers

Threadfin Rainbowfish

Pseudomugilidae Pseudomugil gertrudae

Melanotaenia goldiei Melanotaenia maccullochi Melanotaenia splendida rubrostriata

Melanotaeniidae Iriathernia werneri

Freshwater Longtom

Strongylura kreffti

Obbes’ Tandan Papua Tandan

Porochilus obbesi Plotosus papuensis

Belonidae

Merauke Tandan

Porochilus meraukensis

Zenarchopterus caudovittatus Long-jawed River-garfish Zenarchopterus novaeguineae Fly River-garfish

Narrow-fronted Tandan

Neosilurus ater

Giant Catfish Lesser Salmon Catfish Triangular Shield Catfish Duck-billed Catfish Comb-spined Catfish Froggatt’s Catfish Spoon-snouted Catfish Day’s Catfish

Hemiramphidae

Plotosidae

Arius dioctes Arius graeffei Arius leptaspis Arius spatula Cinetodus carinatus Cinetodus froggatti Doiichthys novaeguineae Nedystoma dayi

(continued)

26,33

25,33

22,33

Fish records in the Trans-Fly / 1417

PAGE 1417

Macleay’s Glassfish Nalua Glassfish Bleeker’s Glassfish Pennyfish Giant Glassfish

Lates calcarifer Ambassis agrammus Ambassis interruptus Ambassis macleayi Ambassis nalua Ambassis urotaenia Denariusa bandata Parambassis gulliveri

Centropomidae

Ambassidae

PS

Amniataba affinis Hephaestus raymondi

Long-spined Glassfish

Ophisternon bengalense Ophisternon gutturale

Synbranchidae

03-15-07 07:29:59

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

Terapontidae

Sailfin Glassfish

Crenimugil heterocheilus Liza alata Liza macrolepis Liza subviridis

Mugilidae

APP5

Tiger Grunter Raymond’s Grunter

Barramundi

One-gilled Eel Swamp Eel

Fringe-lipped Mullet Basket Mullet Large-scaled Mullet Greenback Mullet

Kubuna Hardyhead

Craterocephalus randi

Atherinidae

Bensbach, Morehead, Fly rivers Bensbach, Morehead rivers

Merauke, Bensbach, Fly, Aramia rivers Merauke, Bensbach, Oriomo rivers Bensbach, Oriomo, Fly, Aramia rivers Bensbach River Bensbach River Bensbach, Oriomo, Fly rivers Bensbach, Oriomo, Fly, Aramia rivers

Bensbach, Binaturi, Fly rivers

Bensbach River Bensbach River

Fly River Fly River Bensbach River Bensbach River

Bensbach, Morehead, Fly, Aramia rivers

Common name Locality Native Species

Specific name

Family

Appendix 8.5. (Continued)

2,6,11,32 2,6,32

6,7,9 2,6,32 2,6,7,9,13,19,32 2,6,7,9,11,13,19,32

2,6,7,9,13,19,32

2,6,7,9,32

2,6,7,9,13,19,32

2,6,7,9,11,13,19

6 2,32

2,13,19 16,19 2,6,32 2,6,11,32

2,6,7,9,13,19,32

Source (recent period 1975–2004)

23,33

Source (early Dutch period 1900–1955)

1418 / appendix 8 .5

PAGE 1418

Digul, Bensbach, Morehead, Fly rivers Merauke River Bensbach, Morehead, Fly, Aramia rivers

Barred Gudgeon Crimson-tipped Goby Brown Gudgeon Ebony Gudgeon Empire Gudgeon Trout Gudgeon Snakehead Gudgeon

Acanthopagrus berda Nibea squamosa Toxotes chatareus Toxotes lorentzi Omobranchus punctatus Bostrichthys zonatus Butis butis Eleotris fusca Eleotris melanosoma Hypseleotris compressa Mogurnda mogurnda Ophieleotris aporos

Sparidae

Sciaenidae

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

Toxotidae

Blenniidae

APP5

Eleotridae

Muzzled Blenny

Seven-spot Archerfish Lorentz’s Archerfish

Sharpnose Croaker

Black Bream

New Guinea Tigerfish

Coius campbelli

Datnioididae

Papuan Blackbass

Merauke area Oriomo, Fly rivers Oriomo, Burei, Fly rivers Fly River Bensbach, Binaturi Merauke, Bensbach, Fly, Aramia rivers Binaturi, Fly rivers

Merauke River

Bensbach, Fly, Aramia rivers Merauke, Bensbach, Fly, Aramia rivers

Fly River

Fly River

Bensbach, Oriomo, Fly, Aramia rivers

Fly River

Bensbach, Oriomo, Fly, Aramia rivers Slender Mouth Almighty- 2,6,13,19,32 Bensbach, Fly rivers

Mouth Almighty

Lutjanus goldiei

Glossamia narindica

Glossamia aprion

Crescent Perch Lake Grunter

Terapon jarbua Variichthys lacustris

Lutjanidae

Apogonidae

Lorentz’s Grunter

Pingalla lorentzi

03-15-07 07:30:00

PS

2,32

2 2,7,9,13,19,32 2,7,9,13,19,32 2,32 2,6,32 2,6,7,9,32

2,6,7,9,11,13,19,32 2,6,7,9,32

2,18,19

13,19

2,6,7,9,11,32

2

2,6,7,9,13,19,32

2,6,13,19,32

2,6,7,9,32

(continued)

23,33

21,33

27,33

23,33

Fish records in the Trans-Fly / 1419

PAGE 1419

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APP5

Brachyamblyopus urolepis Gymnoamblyopus novaeguineae Taenioides cirratus Scatophagus argus

Gobioidae

03-15-07 07:30:00

PS

Scatophagidae

Spotted Scat

Bearded Wormgoby

Scaleless Wormgoby Fly River Wormgoby

Weber’s Mudskipper Speckled Goby Spotfin Goby Bearded Goby Barcheek Goby Slender Mudskipper

Munro’s Goby Dwarf Goby Larval Goby Longtail Goby Jaarman’s Goby Freycinet’s Mudskipper New Guinea Mudskipper

Black-banded Gauvina Small-eyed Sleeper

Oxyeleotris selheimi Prionobutis microps Glossogobius sp. 1 Glossogobius sp. 2 Gobiopterus semivestitus Oligolepis acutipennis Oxyurichthys jaarmani Periophthalmodon freycineti Periophthalmus novaeguineaensis Periophthalmus weberi Redigobius bikolanus Redigobius chrysosoma Scartelaos histiophorus Stenogobius psilosinionus Zappa confluentus

Aru Gudgeon Fimbriate Gudgeon Poreless Gudgeon Few-pored Gudgeon

Oxyeleotris aruensis Oxyeleotris fimbriata Oxyeleotris nullipora Oxyeleotris paucipora

Merauke, Fly rivers

Fly, Aramia rivers

Binaturi, Oriomo, Fly rivers Fly River

Digul, Binaturi, Fly rivers Bensbach, Upper Fly rivers Fly River Oriomo, Burei rivers Fly River Bensbach River Merauke, Bensbach, Morehead rivers Oriomo, Fly rivers Bensbach, Oriomo, Fly rivers Oriomo River Merauke River Fly River Fly River

Bensbach, Fly rivers Bensbach River Bensbach, Fly rivers Digul, Bensbach, upper Fly rivers Bensbach, Fly rivers Merauke, Oriomo, Fly rivers

Common name Locality Native Species

Specific name

Gobiidae

Family

Appendix 8.5. (Continued)

13

2,7,9,32

2,13,19,32 2,10,32

2,20,32 13,19

2,13,32 2,6,32 2,32

2,32 2,6,32 15,19 2,32 13,19 6,12 2,6,32

2,6,7,9,13,19,32 2,32

2,6,32 2,6,11,32 2,6,13,19,32 2,6,13,19,32

Source (recent period 1975–2004)

23,33

28,33

23,33

Source (early Dutch period 1900–1955)

1420 / appendix 8 .5

PAGE 1420

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Anabas testudineus Channa striata

Anabantidae

Channidae

Walking Catfish

Striped Snakehead

Climbing Perch

Tilapia

Merauke River, Daru Merauke, Morehead, Oriomo, Aramia rivers

Oriomo, Fly rivers Merauke River

Merauke, Bensbach, Fly rivers Merauke, Fly rivers

Bensbach, Oriomo, Fly, Aramia rivers

Bensbach River

Bensbach River

Bensbach River

Bensbach River

Introduced species

Darwin’s Toadfish Merauke Toadfish

Freshwater Tongue-sole Spotted Tongue-sole

Tailed Sole Velvety Sole

Nurseryfish

2,6,11

2,6,32

2,6,11

2,6,32

5 2,7,9,19,32

2,13,19,32

2,6,13,19,32 2,13,19

2,6,7,9,12,13,19,32

30,33 31,33

29,33

23,33

Sources: From Allen, G.R. (2004), A review of the freshwater fish fauna of the Trans-Fly Ecoregion (unpublished report to WWF South Pacific Programme). Numbers indicate the following sources: 1 Allen (1975); 2 Allen, G.R. (1991), Field Guide to the Freshwater Fishes of New Guinea (Christensen); 3 Compagno, L.J.V., V.H. Niem (1998), Carcharhinidae, in K.E. Carpenter and V.H. Niem, eds., FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western Central Pacific, vol. 2, Cephalopods, Crustaceans, Holothurians and Sharks (Rome: FAO); 4 Compagno and Roberts (1982); 5 Hardy, G.S. (1982), Two new generic names for some Australian pufferfishes (Tetraodontiformes: Tetraodontidae), with species’ redesciptions and osteological comparisons, Aust. Zoologist 21(1): 1–26; 6 Hitchcock, G. (2002), Fish fauna of the Bensbach River, southwest Papua New Guinea, Mems. Queensland Mus. 48(1): 119–122; 7 Kailola, P.J. (1975), A catalogue of the fish reference collection at the Kanudi Fisheries Research Laboratory, Port Moresby, Research Bulletin 16 (Dept. Agriculture, Stock and Fisheries, Port Moresby): 1–275; 8 Kailola, P.J. (2000), Six new species of fork-tailed catfishes (Pisces, Teleostei, Ariidae) from Australia and New Guinea, The Beagle (Rec. Mus. Art Galleries N. Terr.) 16: 127–144; 9 Kanudi Fisheries Research Laboratory, Port Moresby; 10 Murdy, E.O., and C.J. Ferraris Jr. (2003), Gymnoamblyopus novaeguineae, a new genus and species of worm goby from Papua New Guinea (Gobiidae: Amblyopinae), Zootaxa no. 150: 1–6; 11 photo; 12 Queensland Museum, Brisbane, Australia; 13 Roberts, T.R. (1978), An ichthyological survey of the Fly River in Papua New Guinea with descriptions of new species, Smithsonian Contrib. Zool. 281: 1–72; 14 Roberts (1978) as A. cleptolepis; 15 Roberts (1978) as Gobiopterus sp.; 16 Roberts (1978) as Liza diadema; 17 Roberts (1979); 18 Sasaki, K. (1992), Two new species of Nibea (Sciaenidae) from northern Australia and Papua New Guinea, Japan. J. Ichthyol. 39(1): 1–7; 19 United States National Museum of Natural History, Washington, D.C.; 20 Watson, R.E. (1991), A provisional review of the genus Stenogobius with descriptions of a new subgenus and thirteen new species (Pisces: Teleostei: Gobiidae), Rec. West. Aust. Mus. 15(3): 571–654; 21 Weber, M. (1907). Su¨sswasserfische von Neu-Guinea ein Beitrag zur Frage nach dem fru¨heren Zusammenhang von NeuGuinea und Australien, in Nova Guinea. Re´sultats de l’expe´dition scientifique Ne´erlandaise a` la Nouvelle-Guine´e. Su¨sswasserfische Neu-Guinea, vol. 5 (Zool.), pt. 2: 201–267, pls. 11–13.; 22 Weber, M. (1913), Su¨sswasserfische aus Niederla¨ndisch Su¨d- und Nord-Neu-Guinea, in Nova Guinea. Re´sultats de l’expe´dition scientifique Ne´erlandaise a` la NouvelleGuine´e. Zoologie, vol. 9, livr. 4 (Leiden: Zool. Nouvelle-Guine´e), 513–613, pls. 12–14.; 23 Weber (1917); 25 Weber (1917) as Rhobatractus kochi; 26 Weber (1917) as Melanotaenia maculata; 27 Weber (1917) as Petroscirtes kochi; 28 Weber (1917) as Boleophthalmus viridis; 29 Weber (1917) as Symphurus vittatus; 30 Weber (1917) as Tetrodon fasciatus; 31 Weber (1917) as Tetrodon staigeri; 32 Western Australia Museum, Perth; 33 Zoological Museum, University of Amsterdam, The Netherlands.

Oreochromis mossambica

Marilyna darwinii Marilyna meraukensis

Tetraodontidae

Cichlidae

Cynoglossus heterolepis Paraplagusia bilineata

Cynoglossidae

Clarias batrachus

Aseraggodes klunzingeri Synaptura villosa

Soleidae

Clariidae

Kurtus gulliveri

Kurtidae

Fish records in the Trans-Fly / 1421

APP5

03-15-07 07:30:00

PS

PAGE 1421

Appendix 8.6. Mammal records in the Trans-Fly Specific name

Common name

Published habitat description WO SW LF MA SA SF GL HS

RF

Tachyglossus aculeatus Short-beaked Echidna

X

Dasyurus spartacus

Bronze Quoll

Myoictis wallacei

Three-striped Dasyure

P,E

Antechinus melanurus

Black-tailed Antechinus

P,E

Planigale novaeguineae Papuan Planigale

X

X

X

X

X

X

X

X?

Sminthopsis archeri

Chestnut Dunnart

X

Sminthopsis virginiae

Red-cheeked Dunnart

X

X

X

Echymipera echinista

Menzies’ Echymipera

Echymipera kalubu oriomo

Common Echymipera

P,E

X

X

E

Echymipera rufescens

Long-nosed Echymipera

P,E

E

E

X

Isoodon macrourus

Northern Brown Bandicoot

X

X

X

X

Lagorchestes conspicillatus

Spectacled Hare-wallaby

X

X

Dorcopsis luctuosa phyllis

Grey Dorcopsis

Macropus agilis

Agile Wallaby

X

X

Thylogale brunii

Dusky Pademelon

P,E

Thylogale stigmatica oriomo

Red-legged Pademelon

X

Phalanger mimicus

Southern Common Cuscus

X

X

Spilocuscus maculatus

Common Spotted Cuscus

X

X

X

Distoechurus pennatus

Feather-tail Possum

P,E

Dactylopsila trivirgata kataui

Striped ossum

X

X

X

Petaurus breviceps

Sugar Glider

X

X

P

Hydromys chrysogaster Common Water-rat

X

X

E X X X

X

Any permanent water, either marine or fresh

Xeromys myoides

False Water-rat

X

Leptomys signatus

Fly River Leptomys

Conilurus penicillatus

Brush-tailed Rabbit-rat

X

Pseudomys delicatulus

Delicate Mouse

X

Chirumys vates

Lesser Tree-mouse

P,E

Lorentzimys nouhuysi

Long-footed Tree-mouse

P,E

E

X

X

1422

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

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PAGE 1422

Mammal Records in the Trans-Fly / 1423 Paramelomys leucogaster

White-bellied Melomys

P,E

P,E

Melomys levipes

Long-nosed Melomys

P,E

Melomys platyops

Lowland Melomys

P,E

Melomys burtoni

Grassland Melomys

X

Melomys rufescens niviventer

Black-tailed Melomys

X

Paramelomys lorentzi

Lorentz’s Melomys

X

X

Paramelomys moncktoni Monckton’s Melomys

P

Pogonomelomys brassi

E

Chestnut Tree-mouse

Pogonomys mollipilosus Large Pogono-melomys

E

Uromys caudimaculatus White-tailed Giant Rat

P,E

X

Rattus leucopus ringens Cape York Rat

X

X

Rattus sordidus aramia Canefield Rat

X

X

X

X

Dobsonia minor

Lesser Bare-backed Fruit Bat E

Dobsonia moluccensis

Great Bare-backed Fruit Bat

Macroglossus minimus

Northern Blossom Bat

X

X

X

Nyctimene albiventer

Common Tube-nosed Bat

X

X

X

Nyctimene aello

Greater Tube-nosed Bat

E

Nytimene draconilla

Lesser Tube-nosed Bat

Nytimene robinsoni

Palla’s Tube-nosed Bat

X

Paranyctimene raptor

Green Tube-nosed Bat

E

Pteropus alecto

Black Flying Fox

Pteropus macrotis

Big-eared Flying Fox

Pteropus neohibernicus Greater Flying Fox

X

X

X X X

X X

E

X

X

X

E

X

X

X

Pteropus scapulatus

Little Red Flying Fox

X

Rousettus amplexicaudatus

Common Rousette Bat

E

P

Syconycteris australis

Common Blossom Bat

X

X

Mosia nigrescens

Lesser Sheath-tailed Bat

E

Saccolaimus mixtus

Papuan Sheath-tailed Bat

Saccolaimus saccolaimus

Naked-rumped Sheathtailed Bat

E

E

E

Taphozous australis

Southern Sheath-tailed Bat E

E

E

E

X X

X

(continued)

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

APP6

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PS

PAGE 1423

1424 / appendix 8 . 6

Appendix 8.6. (Continued) Specific name

Common name

RF

Published habitat description WO SW LF MA SA SF GL HS

Hipposideros ater

Dusky Leaf-nosed Bat

E

E

Hipposideros maggietaylorae

Maggie Taylor’s Leaf-nosed E Bat

E

Hipposideros muscinus

Fly River Leaf-nosed Bat

E

Hipposideros cervinus

Fawn Leaf-nosed Bat

E

E

Hipposideros diadema

Diadem Leaf-nosed Bat

P,E

E

Chalinolobus nigrogriseus

Pied Wattled Bat

E

Myotis moluccarum

Arafura Large-footed Bat

Most habitat types so long as they are proximate to water

Pipistrellus angulatus

New Guinea Pipistrelle

Pipistrellus papuanus

Papuan Pipistrelle

X

Scotorepens sanborni

Sanborn’s Broad-nosed Bat

X

E

Nyctophilus microtis

Papuan Big-eared Bat

P,E

Nyctophilus timoriensis Greater Big-eared Bat

P,E

P,E

X?

Kerivoula muscina

Fly River Woolly Bat

P,E

Minopteris australis

DOT in Bonaccorso map, but P,E no matching specimen

P,E

P,E

Chaerephon jobiensis

Northern Mastiff-bat

X

X

X

Mormopterus beccarii

Beccari’s Mastiff-bat

X

P,E

E

E

E X E

X

P,E

Note: Published habitat description abbreviations: RF, rainforest; WO, woodland (includes monsoon forest); SW, swamp woodland with Melaleuca; LF, littoral forest; MA, mangroves; SA, savanna; SF, savanna with gallery forest; GL, grassland; HS, herbaceous swamp. Cell entries: X, written data on habitat for collected specimens; P, prediction based on collection locality for specimen; E, prediction based on extralimital ecological data. Source: From Helgen, K.M., and P.M. Oliver (2004), A review of the mammal fauna of the Trans-Fly Ecoregion (unpublished report to WWF South Pacific Programme).

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

APP6

03-15-07 07:30:04

PS

PAGE 1424

Appendix 8.7. IUCN Protected Area Management categories i u cn h a s de f i n ed a series of six protected area management categories, based on primary management objectives. These are summarized below. Category Ia: Strict Nature Reserve: protected area managed mainly for science Area of land and/or sea possessing some outstanding or representative ecosystems, geological or physiological features and/or species, available primarily for scientific research and/or environmental monitoring. Category Ib: Wilderness Area: protected area managed mainly for wilderness protection Large area of unmodified or slightly modified land, and/or sea, retaining its natural character and influence, without permanent or significant habitation, which is protected and managed so as to preserve its natural condition. Category II: National Park: protected area managed mainly for ecosystem protection and recreation Natural area of land and/or sea, designated to (a) protect the ecological integrity of one or more ecosystems for present and future generations, (b) exclude exploitation or occupation inimical to the purposes of designation of the area and (c) provide a foundation for spiritual, scientific, educational, recreational and visitor opportunities, all of which must be environmentally and culturally compatible. Category III: Natural Monument: protected area managed mainly for conservation of specific natural features Area containing one, or more, specific natural or natural/cultural feature which is of outstanding or unique value because of its inherent rarity, representative or aesthetic qualities or cultural significance. Category IV: Habitat/Species Management Area: protected area managed mainly for conservation through management intervention Area of land and/or sea subject to active intervention for management purposes so as to ensure the maintenance of habitats and/or to meet the requirements of specific species. Category V: Protected Landscape/Seascape: protected area managed mainly for landscape/seascape conservation and recreation Area of land, with coast and sea as appropriate, where the interaction of people and nature over time has produced an area of distinct character with significant aesthetic, ecological and/or cultural value, and often with high biological diversity. Safeguarding the integrity of this traditional interaction is vital to the protection, maintenance and evolution of such an area. Category VI: Managed Resource Protected Area: protected area managed mainly for the sustainable use of natural ecosystems Area containing predominantly unmodified natural systems, managed to ensure long term protection and maintenance of biological diversity, while providing at the same time a sustainable flow of natural products and services to meet community needs. 1425

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

APP7

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PAGE 1425

1426

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

APP8

03-15-07 07:30:08

PS

PAGE 1426

TN TNL CA CA CA CA

Wasur

Teluk Cenderawasih

Rawa Biru

P. Batanta Barat

P. Biak Utara

Peg. Cyclops/Dafonsoro

CA CA SM SM SM SM SM

P. Waigeo Barat

Peg. Weyland

Enarotali

P. Sabuda dan P. Tataruga

P. Anggrameos

S. Rouffaer

Pulau Dolok (P. Kimaam)

Peg. Jayawijaya TW

CA

P. Misool Selatan

Sorong

CA CA

P. Yapen Tengah

CA

TN

Lorentz

P. Salawati Utara

Reserve Type

Protected Area

Ib

Ib

Ib

Ib

Ib

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

II

II

II

IUCN Category

Appendix 8.8. Protected areas in Papua

G

G

G

G

G

G

G

G

G

G

G

G

G

G

G

G

G

G

G

Status

945

800,000

664,627

310,000

2,500

5,000

300,000

300,000

153,000

84,000

59,000

57,000

22,500

6,138

116,749

4,000

1,453,500

413,810

2,450,000

Area (ha)

180–4,646

150

200

0–205

sea level

1,750–4,000

900–3,892

0–777

0–565

500–1,496

0–93

90–2,160

0–695

0–1,183

0–90

sea level

0–90

0–5,031

Elevation (m)

397/Kpts/Um/I/1981 (7 Mei)

914/Kpts/Um/10/1981(30 Okt.)

37/Kpts/Um/6/1978 (13 Juli)

820/Kpts/Um/1/1982 (24 Feb.)

547/Kpts/Um/6/1981(30 Juni)

82/Kpts-II/1993 (16 Februari)

84/Kpts/Um/2/1980 (11 Feb.)

84/Kpts/Um/2/1980 (11Feb.)

395/Kpts/Um 1.5/1981 (7 Mei)

716/Kpts/Um/1982 (12 Okt.)

755/Kpts/Um/1982 (12 Okt.)

114/Kpts/Um/1982 (14 Jan.)

56/Kpts/Um/1/1978 (26 Jan.)

212/Kpts/Um/1982 (8 Apr.)

912/Kpts/Um/1981(30 Okt.)

252/Kpts/Um/1/1978 (2 Mei)

472/Kpts-II 1993 (2 September)

448/MenHut-IV/1990 (6 Maret)

154/Kpts-II/1997 (19 Maret)

Decree

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

APP8

03-15-07 07:30:08

CA CA

Pantai Mubrani-Kaironi

Pantai Wewe-Koor

SM

Inggresau CA

CA

Teluk Bintuni Mangrove

CA

CA

Peg.Tamrau Utara

Pantai Sidei-Wibain

CA

Peg. Tamrau Selatan

Pantai Sausapor

CA CA

Kumbe-Merauke

CA

Peg. Kumawa

S. Kais

CA

CA

Gunung (Pantai) Wagura-Kote

Peg.Wandamen/Wondiwoi

CA

P. Numfoor

CA

CA

P. Pombo

CA

TWA

Teluk Yotefa

Peg. Fakfak

TWA

Gunung Meja

P. Supiori

TWA

Nabire

PS

Ia

Ia

Ia

Ia

Ib

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

Ia

P

P

P

P

P

P

P

P

P

P

P

G

P

G

P

P

P

G

G

G

TGHK 820/Kpts/Um/11/1982

100 (20km  50m) 100 (28km  50km)

820/Kpts/Um/11/1982

Rek.Bupati No.072/492/SET (4 Des’97)

TGHK

TGHK

TGHK

TGH

82/Kpts-II/1993 (16 Februari)

TGHK

525/Kpts/Um/7/1982 (17 Juli)

TGHK

?

TGHK

372/Kpts/Um/1/1978 (12 Agt.)

19/Kpts/Um/1980 (19 Jan.)

TGHK

0–3,825

0–3,825

0–150

0–1,442

0–2,222

275–1,620

0–1,034

100

sea level

sea level

50–250

21/Kpts/Um/1/1980 (11 Jan.)

90 (18 km  50 m)

(14km  50m)

(12km  50m)

300,000

368,365

247,875

126,810

122,000

188,150

73,022

191,000

42,000

15,000

1,500

100

1,675

460

100

(continued)

Protected Areas in Papua / 1427

PAGE 1427

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

SM SM SM

P. Mapia

Kep. Asia dan Ajoe

Kep. Asia

APP8

03-15-07 07:30:08

TW TW TW TW

Sausiram

Parieri (Biak)

Beriat

Klamono

Danau Bian SM

SM SM

Kep. Raja Ampat

Teluk Lelintah

TW

P. Sayang

SM

SM

Mamberamo/Peg. Foja

Kep. Ajoe

CA

SM

Pantai Jamursba-Mendi

Peg. Arfak

Reserve Type

Protected Area

Appendix 8.8. (Continued)

Ib

Ib

Ib

Ib

Ib

Ib

Ib

Ib

Ia

Ib

IUCN Category

PS

G

G

P

P

P

P

P

P

P

P

P

P

G

G

P

Status

1,909

9,193

2,000

1,000

50,000

69,390

96,000

168,630

7,000

1,531

4,015

1,018,000

68,325

10,000

Area (ha)

50

50

sea level

sea level

0–2,103

1,500–2,941

Elevation (m)

SK Menhut No.219/Kpts-II/1992 (27 Mart’93)

SK Menhut No. 850/Kpts-II/1992 (31 Agust’92)

TGHK

TGHK

SK Menhut No. 81/Kpts-II/1993

Srt.No.1208/PPA.030 VII/SBKSDA IRJA I/1992 (20 Juli’ 92)

Rek. Bupati No.050/5/11 (9Jan’94)

Srt.No.151/PPA.030/IV/SBKSDA IRJA I/92 (16/4/97)

914/Kpts/Um/10/1981(30 Okt.)

783/Kpts-II/92 (11 Agustus 1992)

Rek.Bupati No.522.5/1010 (5 Nov’94)

Decree

1428 / appendix 8. 8

PAGE 1428

SML SML SML SML SM TWL TWL

Pulau Numfoor/Pulau Manem

Pulau Ayawi

Pulau Venu

Mangima

Pulau Misool

Pulau Kofiau

TB

Pulau Rumberpon SML

TWA

Moraid

Pulau Kofiau

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

Misool Selatan

CA

P. Waigeo Timur

Ib

Ib

Ib

Ib

Ib

Ib

Ia

P

P

P

P

P

P

P

P

P

P

G

7,747

12,549

3,800

16,320

1,547

2,924

7,197

4,319

4,205

6,300

119,500

APP8

03-15-07 07:30:08

PS

(continued)

Srt.No.1208/PPA.030 VII/SBKSDA IRJA I/1992 (20 Juli’ 92)

Rek.Bupati No.552.5/447 (25 Mei 1992)

Rek.Bupati No.503/1204 (22 Agust. 1995)

Rek.Bupati No. 660.1/1489 (9 Sept. 1993)

Rek.Bupati No.660.1/1826 (29 Nov. 1993)

Srt.No.3075/PPA.030/XII/SBKSDA IRJA I/91 (21 Des.)

Rek.Bupati No.552.5/447 (25 Mei 1992)

TGHK

TGHK

SK Menhut No.251/Kpts-III/1996

Protected Areas in Papua / 1429

PAGE 1429

TWL TWL TWL TWL CAL

Kep. Dua

Teluk Broe/P. Numfoor

Kep. Padaidori

Pulau Insobabi

Pulau Misool Selatan

APP8

03-15-07 07:30:09

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

Ia

IUCN Category

P

P

P

P

P

Status

2,756

24,600

183,000

2,572

2,084

Area (ha)

Elevation (m)

Rek.Bupati No. 552.5/477 (25 Mei 1992)

Rek.Bupati No.660.1/1489 (9 Sept. 1993)

Rek. Bupati No.522.5/1462 (21 Agust. 1992)

Rek.Bupati No.660.1/1826 (29 Nov. 1993)

Srt.No.2469/II-SBKSDA IRJA I/1992 (21 Okt.)

Decree

Note: General abbreviations: P, Pulau (Island/s); S, Sungai (River); Peg, Pegunungan (Mountains); TGHK, Tata Guna Hutan Kesepakatan (Land Use Agreement). Category abbreviations: CA, Cagar Alam (strict nature reserve); CAL, Cagar Alam Laut (marine strict nature reserve); TW, Taman Wisata (recreational park); TWA, Taman Wisata Alam (recreational nature park); TWL, Taman Wisata Laut (recreational marine park); TNL, Taman Nasional Laut (marine national park); TN, Taman Nasional (national park); TB, Taman Buru (hunting park); SM, Suaka Margasatwa (wildlife sanctuary); SML, Suaka Margasatwa Laut (marine wildlife sanctuary). Status abbreviations: P, proposed; G, Gazetted. Sources: 1. Laporan tahunan Sub BKSDA IrJa I Sorong tahun 1995/1996 (Lamp.1&2); 2. Laporan tahunan Sub BKSDA IrJa II tahun 1996/1997 (Lamp.2); 3. National Conservation Plan for Indonesia, Maluku and Irian Jaya; Field Report of UNDP/FAO (hal 4–23); 4. Data Kawasan Konservasi di Kabupaten Sorong s/d tahun 1997; 5. Konservasi Alam dan Pembangunan di Irian Jaya (Ronald G. Petocz), hal 72 & Lamp. 4; 6. IUCN (199), Guidelines for Protected Areas Management Categories (IUCN, UK and Gland).

Reserve Type

Protected Area

Appendix 8.8. (Continued)

1430 / appendix 8. 8

PS

PAGE 1430

1431

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

APP9

03-15-07 07:30:10

PS

PAGE 1431

507 79,613

CA

Batanta Barat

Misool Selatan CA

3,050

CA CA

Sausafor

CA

Peg. Kumawa

Peg. Tamrau Utara

CA

Peg. Fakfak

CA

CA

Peg. Tamrau Selatan

CA

CA

Waigeo Timur

Peg. Cycloop

CA

Waigeo Barat

Peg. Arfak

CA

Supiori

15,162

308

2,587

8,440

0

0

0

0

0

0

0

0

7,686

208,208 0

60,699 0

73,024

89,929

325

0

862

29,460

0

0

0

0

0

0

0

0

Lower Upper Submontane montane alpine rainforest rainforest forest

250,244 121,999

94,493

35,780

17,413

32,349

92,952

81,793

63,443 37,433

Salawati Utara CA

69,428

CA CA

Bupul

Area Name

Biak Utara

Lowland Reserve evergreen Type rainforest

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0 0

0

671

0

0

0

0

0

0

215

1,600

325

515

2,812

47

Mangrove forest

111

0

0

0

0

0

0

926

4,416

0

899

237

0

2,719

0

Swamp forest

0

0

121

0

0

0

0

0

8

71

0

0

0

8,654

0

Swamp brush

Habitat Area (Ha)

Appendix 8.9. Habitat coverage within protected areas in Papua

93

618

3,558

47

0

937

1,850

468

375

193

649

529

0

5,166

0

Brush

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

47

164

5,802

89

0

625

0

4,552

191

0

0

558

Bare Savanna land

419

1,218

603

0

0

426

270

0

1,030

232

293

0

87

314

243

Other uses

0

7,040

14,880

0

1,099

9,131

5,770

342

7,603

2,070

0

20,475

0

27,948

124

Cloud/ no data

(continued)

9,026

350,971

487,135

110,115

37,188

31,981

78,139

99,455

97,016

40,324

65,799

104,224

641

114,230

3,417

Total

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

APP9

SM SM SM SM SM

Jamursba Medi

MamberamoFoja

P. Dolok

P. Komolom

Yapen Tengah

Danau Bian

CA

Wondiboy

SML

CA

Peg. Wayland

Kep. Raja Ampat

CA

Wagura Kote

SML

CA

Teluk Bintuni

Kep. Panjang

CA

Area Name

03-15-07 07:30:11

PS

837

31,642

735,257

433

55,239

984

3,808

72,601

25,360

44,393

16,534

4,968

Lowland Reserve evergreen Type rainforest

Appendix 8.9. (Continued)

0

0

28,131

0

0

326

0

797

171

11,201

0

0

0

0

26,747

0

0

0

0

0

547

37,347

0

0 0

0

0

0

643

0

0

0

185

0

1,018

71

Swamp forest

29,466

61,677

0

32,917

0

0

0

181

0

0

15

Swamp brush

19,041

2,521

66,481 165,302

11,407 268,713

0

0

0

294

0

41

0

687

64,820

Mangrove forest

0 109,273

0

0

0

0

0

0

0

4,670

Lower Upper Submontane montane alpine rainforest rainforest forest

Habitat Area (Ha)

0

0

0

0

0

0

0

0

0

0

0

9,840

0

0

1,110

15,371

418

0

0

422

65

0

53

3,161

Bare Savanna land

1,432 113,769

897

0

3,830

301

256

1,025

3,087

2,170

13

308

Brush

0

0

0

33,842

0

9,268

524

0

36,952

44,401

37,807

232

3,403

Cloud/ no data

2,806

1,087

420 176,731

169

0

467

25

75

1,100

972

Other uses

66,707

680,422

1,167,257

433

102,365

2,583

4,498

112,475

74,997

140,750

18,484

73,586

Total

1432 / appendix 8 .9

PAGE 1432

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

APP9

03-15-07 07:30:11

PS

0

0

0

0

0

0

0

77,613

0

3,076,242 758,435

234

5,082

1,849

2,908

8,821

10,651

23,016

640,146

14,950

41,745 361,797

0

69,203

852,580

0

0

0

0

0

0

0

404,030

0

0

0

190,310

1,352

0

0

0

188

0

74

0

0

0

0

99

0

0

84

0

0

0

99

0

0

0

0

37

0

0

63,360 104,858 0

638

2,363

508

0

0

0

2

0

0

135

0

0

24,310

0

0

0

0

0

33

0

0

0

159

0

164

3,253

0 326,252

0

0

0

0

104 134,551

7,827 38,053

40

41,415

0

3,691 18,018

3,037

21,164

0

74,738

661

1,013

0

1,158

211

75

2,562

450

0

0

0

0

0

90,814

24,503 103,769

86

529

209

68,767

198,918 434,308 605,197 429,334 87,489 258,192 386,900 110,945 734,701

0

0

0

0

0

0

0

158,775 210,599 176,013

0

0

0

35,473

7,933,240

1,345

6,096

1,851

4,460

9,032

10,923

422,603

2,167,581

20,426

469,012

1,842

743,850

Note: Areas calculated for 39 protected areas, including three marine protected areas. For clarity of presentation, some forest and habitat classifications have been grouped (e.g., mining, plantation, and resettlement combined to Other uses; Alpine forest was added to Subalpine forest; Body of water was added to Swamp forest). For general and category abbreviations, see Appendix 8.8. Sources: Map from Dirjen PHKA (2003a). Forest classifications follow Whitmore (1990) and habitat types follow FWI et al. (2001).

Total

TWA

TWA

Kep. Padaido

Sorong

TWA

Gunung Meja

TWA

TWA

Berjat

TWA

TN

Wasur

Nabire

TN

Lorentz

Klamono

TNL

Teluk Cenderawasih

SM

1,051

Sidaey Wibain SM

Sungai Rouffaer

259,340

SM

Peg. Jayawijaya

Habitat Coverage within Protected Areas in Papua / 1433

PAGE 1433

Appendix 8.10. Nongovernmental institutions in Papua that are working on environmental and conservation issues District

Name and address

Focus

Merauke

Yayasan Alam Lestari Masyarakat Maju dan Sejahtera (ALMAMATER) Jl. Brawijaya No.13 Merauke Telephone: (0971) 32133 / Fax: (0971) 325853

Agriculture Health Environment

Merauke

Yayasan Wasur Lestari (YWL) Jl. Ahmad Yani, Jalur Drainese P.O. Box 220 Telephone : (0971) 325408 / Fax: (0971) 32504 E-mail: [email protected]

Environment Capacity building Economic

Merauke

Yayasan Pengembangan Sosial Ekonomi & Lingkungan (YAPSEL) Jl. Missi Merauke Telephone: (0971) 321489 / Fax: (0971) 323204 E-mail: yps [email protected]

Social and economic Environment Advocation Education

Manokwari

Yayasan Pemberdayaan Masyarakat Pribuni (YAPMI) Telephone / Fax: (0986) 214979 E-mail: [email protected]

Capacity Building Environment Economic

Manokwari

Lembaga Bantuan Pertanian Papua (LBP) Jl. Trikora Gg. Melati No. 10 Wosi Manokwari Telephone: (0986) 21528 / Fax: (0986) 211330 E-mail: [email protected]

Agriculture Environment Education

Manokwari

Yayasan Paradisea Jl. Ciliwung Sanggeng P.O. Box 242 Manokwari Telephone: (0986) 211068 / Fax: (0986) 211068

Environment Economic

Manokwari

Yayasan Lingkungan Hidup Humeibou (YALHIMO) Jl. Ciliwung No. 01 Sanggeng P.O. Box 105 Manokwari Telephone: (0968) 212014 E-mail: [email protected] septer [email protected]

Environment Capacity building

Sorong

Yayasan Sosial Agustinus Jl. Jend. A. Yani No. 83 P.O. Box 183 Sorong

Environment Advocation

Sorong

Yayasan Sosial Faumair Lestari (YESFEL) Jl. Kartini No. 2 Kampung Baru Sorong

Social and economic

Sorong

Komisi Irian Keuskupan Sorong Jl. RA. Kartini No. 3 Pastoran Kampung Baru Sorong

Environment Advocation

Sorong

Yayasan SVEHO Jl. Armanolo—Komplex Satin—Teminabuan—Sorong Selatan Telephone: (0952) 31227

Health Education Environment

1434

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

AP10

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PS

PAGE 1434

Non-governmental Institutions Working on Environmental Issues / 1435 Sorong

Yayasan Konpers Jl. D.I. Panjaitan No. 09, RT.IV, RW.VIII Kel. Rufei—Sorong Telephone / Fax: (0951) 324715 E-mail: [email protected]

Environment Capacity building Economic Advocation

Biak

Yayasan Rumsram Jl. Bosnik Raya STAB Biak Telephone: (0981) 23269 / Fax: (0981) 23269 E-mail: [email protected]

Environment Economic

Fakfak

Yayasan Agribisnis dan Lingkungan Hidup (YALHI) Jl. Imam Bonjol Wagom P.O. Box 176 Fakfak Telephone: (0956) 23310 / Fax: (0956) 22573

Environment

Fakfak

Yayasan Pendidikan Kasih Karunia Indonesia (YASPENKARI) Jl. M. Tata Wagom Telephone: (0956) 24301

Education Environment Social and economic

Fakfak

Yayasan Studi Konsultasi dan Bantuan Hukum (YSKBH) Jl. Cenderawasih Fakfak Telephone: (0956) 22882

Education Environment

Fakfak

Yayasan Sosial Pengembangan Kawasan Timur (YASOBAT) Jl. Yos Sudarso, Wagom Fakfak Telephone: (0956) 23934 / Fax: (0956) 22425

Capacity building Economic Environment

Fakfak

Yayasan Kasih Mulia (YKM) Jl. Imam Bonjol P.O. Box 158 Wagom Fakfak Telephone: (0956) 23908 / Fax: (0956) 22452 (Kandatel Fakfak)

Economic Environment Health Advocation

Nabire

Yayasan Sosial Bina Mandiri UTAMA (YABIMU) Jl. Pipit Kali Harapan Tromol Pos 27 Nabire Telephone: (0984) 23881 / Fax: (0984) 23458

Community development Human rights Environment

Timika

Lembaga Musyawarah Adat Amungme (LEMASA) Jl. P. Magal No. 13 Timika Telephone: (0901) 322383 / Fax: (0901) 322472

Economic Advocation Environment

Jayapura

Yayasan Agrosilvopastoral (ASP) Perumahan Grand Permai No. 14 D Kotaraja Telephone / Fax: (0967) 582997 E-mail: [email protected]

Agriculture Environment

(continued)

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

AP10

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PS

PAGE 1435

1436 / appendix 8. 1 0

Appendix 8.10. (Continued) District

Name and address

Focus

Jayapura

Yayasan Lingkungan Hidup Irian Jaya (YALI) Jl. Ifar No. 62 Abepura Telephone / fax: (0967) 584671 E-mail: [email protected]

Environment

Jayapura

Natural Resources Management Program/ Environmental Policy and Institutional Strengthening Indefinite Quantity Contract (NRMP/EPIQ) Perumahan Grand Permai BTN Blok A. No. 8 Kotaraja Telephone / Fax: (0967) 584670 E-mail: nrm [email protected]

Environment Capacity building Information

Jayapura

Aliansi Mahasiswa Peduli Lingkungan Hidup (AMPPeLH) Jl. Raya Sentani No. M 38 Padang Bulan Telephone / Fax: (0967) 583381

Environment

Jayapura

Conservation International Irian Jaya (CI—Irja) Jl. Bhayangkara I No. 33 Jayapura Telephone: (0967) 523423 E-mail: [email protected]

Environment

Jayapura

Yayasan Pendidikan Lingkungan Hidup Cycloops (YPLHC) Jl. SPG. Teruna Bhakti No. 3 Waena Jayapura Telephone / fax : (0967) 572507 E-mail : [email protected]

Education Environment

Jayapura

Wahana Lingkungan Hidup Papua (WALHI Papua) Jl. Raya Sentani No. M 38 Belakang Kantor Lurah Hedam, Padang Bulan Telephone / Fax: (0967) 583381

Environment

Jayapura

Sistem Hutan Kerakyatan (SHK) Jl. Kalibobo No. M 38 Padang Bulan Telephone / Fax: (0967) 583381 E-mail: [email protected]

Environment

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

AP10

03-15-07 07:30:13

PS

PAGE 1436

Appendix 8.11. Government institutions that are working on environmental and conservation issues in Papua District

Name and address

Focus

Jayapura

Dinas Kehutanan Proinisi Papua Jl. Tanjung Ria Base G Jayapura Telephone: 62-967-541522

Forestry issues

Jayapura

Badan Pengendalian Dampak Lingkungan Pro. Papua Kantor Dinas Otonom Provinsi Papua— Gedung B Lantai I Jl. Raya Abepura-Kotaraja Telephone / Fax: 62-967-587694

Environment issues

Jayapura

Balai Pemantapan Kawasan Hutan Wilayah X Papua (Forestry Mapping Branch) Jl. Raya Abepura—Kotaraja Telephone: 62-967-582529 / Fax: 62-967-582527

Forest mapping

Jayapura

BKSDA Papua I (Nature Conservation Bureau) Jl. Raya Abepura—Kotaraja Telephone: 62-967-581596 / Fax: 62-967-585529

Conservation issues

Sorong

BKSDA Papua II Jl. Jend Sudirman No. 40.-Kotak Pos 1053 Sorong Telephone: 62-951-321986 or 951-323097 Fax: 62-951-321986

Conservation issues

Manokwari

Balai Taman Nasional Teluk Cenderawasih Jl. Trikora Wosi-Rendani Manokwari Kotak Pos 229 Manokwari 98312 Telephone / Fax: 62-986-212303 or 986-214719

Conservation issues

Nabire

Kantor Taman Nasional Teluk Cenderawasih Jl. Semarang No. 112 Nabire Telephone: 62-984-421362

Conservation issues

Merauke

Balai Taman Nasional Wasur Jl. Raya Mandal 69 Spaden No. 2 Merauke 99611

Conservation issues

1437

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

AP11

03-15-07 07:30:15

PS

PAGE 1437

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

AP11

03-15-07 07:30:15

PS

PAGE 1438

Index Abepura, 74, 81, 84, 85, 100, 101, 1286, 1288 Adelbert-Finisterre Terrane, 143, 148, 155, 215, 221 Adelbert Mts, 94, 154, 209, 210, 215, 221, 234, 660, 663, 729 Admiralty Islands, 16, 21, 24, 32, 40, 43, 70, 71, 479, 547, 564, 565, 573, 579, 582, 596, 1200 Aua, 16, 40 Manus, 16, 18, 32, 71, 78, 97, 122, 547, 548 Wuvulu, 16, 40 Africa, 27, 41, 95, 143, 307, 312, 313, 335, 346, 349, 371, 375, 379, 399, 400, 401, 421, 429, 436, 440, 442, 462, 473, 495, 510, 522, 548, 570, 571, 580, 584, 638, 642, 646, 1044, 1240, 1380. See also Madagascar Agathis, 9, 65, 80, 248, 345, 946, 949, 958, 962, 966, 978, 980, 990, 994, 1008. See also Copal Aghawaghon River, 1211 Aglaia, 275, 913, 922, 923, 929, 930, 933, 935, 936, 938, 940, 941, 954, 955, 1356 Agricultural and forestry pests, 55, 67, 97, 517, 521, 522, 670, 677, 829, 1127, 1143, 1156, 1203, 1278, 1332–1333, 1365. See also Diseases, nonhuman; Herbicides; Pesticides Agriculture, commercial, 1103, 1140–1143, 1156–1157, 1165, 1171 employment in, 1111, 1151 See also Agricultural and forestry pests; Agriculture, effects on ecosystems; Cash crops; Pesticides; Plantations Agriculture, effects on ecosystems, 8, 329, 406, 990, 991, 1003 See also Land conversion; Pesticides; Pollution Agriculture, traditional, 114, 1125–1140, 1144–1145, 1332 and climate, 182, 188, 189, 193, 1087–1088 early/origins, 8, 126–127, 130, 1087, 1091– 1094, 1126 and rainforest occupation, 126–128, 1096 and soils, 130, 760, 946, 995, 1126, 1129, 1135 swidden, 8, 1129, 1137, 1360

and work ethic, 118 See also Archeological sites, and agriculture; Food crops; Hunting, early human; Land tenure system; Nontimber forest products; Shifting cultivation Agroforestry. See Plantations Aiduna Formation, 246 Air transport, 45, 56, 57, 58, 59, 60, 67, 73, 75, 77, 645, 817, 1101, 1142, 1286, 1151, 1332 Ajkwa River, 217, 264, 831, 847, 849, 859, 884, 956, 958, 1211, 1212 Bridge, 951, 954, 955 Algae, 36, 303, 314, 509, 590, 591, 649, 814, 884, 902, 953, 1005, 1006 coral reefs, 771, 771, 773, 776, 779, 780, 787, 791 cryovegetation, 157, 184, 1049, 1051–1052 freshwater, 859, 869, 881, 940 mangroves, 833, 839, 842–844 Alien species. See Introduced species Alpha diversity, 404, 654, 1371 Alpine and subalpine ecosystems/vegetation, 28, 82, 83, 89, 92, 93, 167, 168, 175, 184, 186, 193, 249, 258, 260, 264, 282, 303, 304, 307, 308, 315–316, 330, 337–339, 340, 345, 371, 384, 385, 389, 391, 429, 432, 440, 441, 753, 754, 757, 762, 763, 863, 870, 871, 979–980, 1013, 1017, 1018, 1021, 1025–1052, 1089 animals, 587, 589, 593, 660, 664, 712, 713, 714, 715, 716, 718, 736, 1090, 1091, 1092 conservation/ecoregion, 1211, 1246, 1268, 1269, 1314, 1431–1433 Alstonia, 9, 355, 467, 468, 558, 911, 912, 916, 917, 918, 920, 922, 923, 924–927, 929, 930, 933, 935, 936, 941, 948, 949, 950, 951, 953, 954, 955, 963, 964, 968, 974, 975, 988, 998, 1056, 1058 Amamapare, 810 Amazon/Amazonia, 589, 645, 1200 Ambon. See Maluku American Samoa, 777, 784, 789, 790. See also Samoa Amungme, 1105, 1115, 1131, 1136, 1140, 1141, 1143

1439

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

INDX

03-19-07 07:21:57

PS

PAGE 1439

1440 / index Anacardium. See Cashew nuts Andaman Islands, 546, 547, 790 Andes (South America), 307, 435, 447 1044 Angaur. See Palau Anggi Lakes (Vogelkop), 6, 50, 53, 54, 58, 61, 65, 73, 79, 80, 84, 470, 875, 993, 997, 1091, 1129 Anisoptera (dragonflies). See Odonata Anisoptera (tree), 290, 468, 949, 952, 983, 984, 985, 987, 988, 990 Anopheles. See Mosquitoes Antarctica, 19, 73, 246, 248, 255, 657, 774, 1271 Antarctic Beech. See Nothofagus Ants, 97, 167, 519, 1312 Fire (Wasmannia), 678 introduced, 678, 1218 and plants, 87, 335, 367, 377, 385, 396, 397, 401, 412, 413, 414, 415, 461, 588, 841, 949, 963, 967, 991, 998, 1008 (see also Rubiaceae; Symbiosis) as prey, 589, 594, 710 April River (PNG), 285 April Ultramafics, 153 Arafura coast/foreland, 209, 210, 217, 218, 238–239, 241, 249, 488, 489 Arafura Sea, 6, 137, 177, 181, 552, 646, 647, 811, 813, 847, 864, 879, 880, 950, 1164 Arafura Shelf, 181, 255, 260, 264 Aramia River (Trans-Fly), 217, 218, 1416– 1421 Araucaria, 9, 34, 39, 68, 83, 261, 345, 347, 947, 959, 978, 980, 981, 983, 992, 993– 994, 1009 Archbold, Richard/Archbold Expeditions, 5, 12,15, 52, 56, 58, 59, 60, 61, 63, 76–77, 88, 106, 215, 272, 337, 488, 515, 534, 569, 656, 942, 945, 979, 980, 1026 Archeological evidence charcoal, 121, 123, 260, 1087, 1089, 1090, 1091, 1092, 1094 fire, 1087, 1089, 1091, 1126 fire-cracked rocks, 121 human burial, 125 pollen record, 123, 1090, 1093 pottery, 125, 1094 shell midden, 125, 814 stone tools, 121, 1087 waisted blade, 121, 1088 See also Human habitation Archeological sites, and agriculture, 121, 167, 255, 261, 1087, 1089, 1091, 1093, 1095, 1126, 1129, 1130, 1133, 1136, 1137, 1139, 1142, 1352

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

INDX

See also Agriculture, traditional; Human habitation; Peat Archeological sites, specific, 123–130, 249, 250, 251, 264, 1091 Golo Cave, 123 Ijomba Bog, 123, 258, 1094 Kelangur, 249, 251 Kelela Swamp, 261, 1093 Kosipe, 121, 1088, 1089 Kria Cave, 123, 125, 128, 130 Kuk Swamp, 121, 1091, 1126 Kwiyawagi, 172, 249, 250, 251, 736, 1090, 1092 Lachitu Rockshelter, 121, 1092 Lemdubu Cave, 123 Mapala Rockshelter, 123, 251, 1092 Nombe Rockshelter, 121, 249, 1088, 1091 Pamwak Rockshelter, 122 Sirunki, 260, 261 Supulah Hill, 174, 260, 1089, 1090, 1094 Tari Basin, 249, 261, 1090 Toe´ Cave, 123, 125, 126, 128 See also Human habitation Arcs Banda Arc, 142, 144, 153, 158, 197, 200 Bismarck Arc, 142, 152, 155, 158, 198 Mariana Islands/Arc, 141, 149, 313, 580, 596 New Britain Arc, 144, 158 Solomons Arc/Plate/Trench, 137, 142–147, 154–156 Sunda Arc, 138, 153, 158 Tonga Arc, 138, 141 Trobriand Arc, 142, 143, 145 See also Geological formations Arfak Mts, 5, 11, 12, 22, 23, 24, 46, 47, 53, 58, 64, 80, 84, 154, 155, 157, 168, 187, 198, 213, 257, 336, 340, 386, 387, 487, 520, 568, 599, 601, 602, 656, 659, 711, 712, 713, 714, 715, 716, 717, 728, 732, 737, 744, 746, 747, 978, 979, 997, 1140, 1244, 1246, 1247, 1262, 1268, 1270. See also Anggi Lakes Arfak Plain, 468, 950 Arguni Bay (Bird’s Neck), 6, 187, 200, 488, 649, 962 Armed Forces, 1100, 1103, 1188, 1277, 1288, 1290, 1292, 1300, 1307. See also Military Arnhemland (Australia), 590 Arowana. See Bony tongues Arso (people), 112, 114 Arso (Tami River basin), 50, 1286, 1352, 1354 Arthropods, 70, 96, 99, 100, 165, 104, 204, 517, 780, 781

03-19-07 07:21:57

PS

PAGE 1440

index / 1441 in caves, 1067, 1072, 1075 as prey, 589, 592, 593, 620, 623, 624 soil arthropods, 99, 101, 165, 516, 517 See also specific taxa Aru Islands, 16, 22, 23, 47, 49, 54, 62, 123, 197, 202, 209, 210, 211, 218, 242, 250, 252, 255, 263, 382, 465, 479, 487, 488, 489, 537, 548, 564, 579, 580, 581, 586, 594, 599, 600, 601, 602, 603, 625, 628, 648, 656, 723, 772, 792, 809, 811, 813, 829, 902, 903, 904, 1200, 1246, 1410– 1415. See also Maluku Asmat (people), 11, 108, 109, 110, 111, 113, 114, 116, 117, 118, 119 Asmat (Casuarina) Coast, 6, 51 Asmat region, 3, 264, 810, 917, 1244 Astrolabe Bay (PNG), 19, 22, 32, 34, 35, 36, 37, 38, 39, 40, 41 Astrolabe Range, 20, 27, 30, 94 Attorney General, 1288, 1289, 1291, 1292 Aure trough, 143, 662, 663 Australian craton, 138, 143, 144, 149, 150– 152, 153, 156, 158, 160, 197, 198, 199, 202, 204, 213, 214, 221, 286, 483 Australian Plate, 4, 143, 196, 201, 481, 532, 640, 646, 658, 662, 825 Austronesians, 108, 722, 1087, 1093–1094, 1128. See also Lapita Autonomy, 1188 General, 1220 Regional, 1219, 1258, 1281, 1282, 1316 Special, 1099, 1116, 1117–1120, 1121, 1169, 1172, 1189, 1219, 1220, 1258, 1282, 1304, 1308, 1315, 1316 See also Legislation, Laws (UU) Avicennia, 9, 507, 549, 550, 827, 828, 830, 831, 832, 833, 880 Ayamaru (people), 111, 118, 125, 127, 887, 1120 Ayamaru Lakes (Vogelkop), 63, 80, 213, 264, 375, 645, 730, 887 Ayamaru Plateau, 123, 124, 126, 128, 130, 250 Ayawasi (Vogelkop), 80, 387, 442 Bacan Island. See Maluku Bailey, Robert, 127 Bali, 640, 760, 814, 1112, 1239, 1318, 1322, 1349, 1350 Baliem River, 5, 249, 250, 258, 881, 890, 1217 swallet/cave, 881, 1066, 1070 Baliem Valley, 8, 11, 12, 56, 73, 74, 81, 82, 84, 123, 168, 171, 172, 186, 187, 189, 201, 222, 257, 260, 261, 521, 647, 717, 728, 735, 736, 869, 962, 990, 1038, 1087–

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

1093, 1126–1132, 1135, 1139, 1141, 1142, 1144, 1244 Bananas, 91, 588, 624, 956, 1091, 1095, 1126, 1130, 1135, 1136, 1137, 1140, 1360 Banda Arc, 142, 144, 153, 158, 197, 200 Banda Sea, 255, 411, 552 Bandicoots (Echymipera, Isoodon, Microperoryctes, Peroryctes), 200, 250, 406, 693, 694, 708, 713–714, 726, 729, 730, 731, 732, 734, 735, 736, 737, 744, 1404, 1422 Banggai/Togian/Sula Islands (Sulawesi), 78, 157, 247, 496, 497, 503, 639, 793, 794, 1312 BAPPEDA. See Development planning BAPPENAS. See Development planning Barari (Arguni Bay), 187 Barringtonia, 9, 270, 273, 468, 833, 879, 903, 904, 906, 907, 911, 919, 930, 933, 935, 942, 1055, 1057 Basilaki Island, 209, 219, 244 Batanta Island. See Raja Ampat Islands Bats (Chiroptera), 10, 352, 381, 391, 402, 406, 432, 460, 466, 483, 484, 687, 689, 691, 704, 710, 719–721, 724, 725, 726, 731, 734, 748–749, 841, 1066, 1072, 1077, 1078, 1079–1080, 1081 Beehler, Bruce, 96, 105, 198, 670 Bees, 352, 367, 401, 402, 425, 451, 459, 465, 475, 519 Beetles. See Coleoptera Bensbach River, 887, 889, 1328, 1332, 1340, 1416–1421 Berau Bay, 112, 169, 468, 810, 917, 920, 1212, 1213, 1214. See also Bintuni Bay Berau Peninsula, 63, 199 Berrypeckers (Melanocharitidae, Paramythiidae), 655, 658, 673, 1403 Beta diversity, 8, 293, 566, 589, 1235, 1243, 1372 Bewani Mts, 209, 210, 214, 215, 233, 489, 660, 664, 709, 717, 729, 730 Biak (people), 108, 109, 110, 111, 112, 113, 114, 117, 118 Biak/Supiori Island (Cenderawasih Bay), 7, 154, 157, 168, 174, 197, 200–201, 204, 207, 209, 210, 211, 214, 220, 231, 257, 366, 386, 468, 487, 489, 499, 505, 537, 569, 599, 601, 618, 619, 620, 621, 623, 629, 630, 637, 645, 678, 692–702, 704, 705, 712, 715, 716, 720, 726, 730, 733, 745, 746, 748, 749, 792, 808, 809, 813, 816, 891, 994, 1071, 1073, 1075, 1076, 1077, 1079, 1080, 1102, 1104, 1111, 1113, 1120, 1142, 1151, 1161, 1200,

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1442 / index Biak/Supiori Island (Cenderawasih Bay) (cont.) 1220, 1221, 1243, 1246, 1269, 1270, 1271, 1362, 1426, 1428, 1431 Bian River (Trans-Fly), 1054, 1055, 1056, 1327, 1328, 1428, 1432 Bintuni Bay, 6, 110, 112, 169, 255, 273, 275, 468, 488, 568, 645, 648, 660, 661, 662, 663, 810, 827, 831, 833, 838, 839, 849, 850, 880, 890, 917, 920, 962, 1117, 1119, 1151, 1154, 1155, 1206, 1207, 1212, 1213, 1214, 1217, 1270, 1404–1409, 1427, 1432. See also Berau Bay Biodiversity, 1201, 1230 alpha diversity, 404, 654, 1371 beta diversity, 8, 293, 566, 589, 1235, 1243, 1372 and ecosystem function, 761–762 keystone resources/species, 668, 677, 842, 1202 measuring, 1202, 1233–1234 See also Biodiversity, threats to Biodiversity hotspots, 274, 764, 1200, 1205, 1231, 1237 animals, 504, 534, 537, 599, 644, 645, 651, 652 marine, 511, 1237 plants, 274, 317, 359, 435 See also Conservation designation, Hotspots Biodiversity, threats to. See Dams; Endangered and threatened species; Habitat degradation; Habitat loss; Introduced species; Logging, ecological aspects; Mining; Pollution; Wildlife trade Biogeographical history Cathaysian biota, 246 Gondwanan biota, 88, 221, 246, 252, 286, 320, 322, 381, 404, 570, 571, 574, 689, 980 Gondwanan land forms, 246, 287, 288, 659 Laurasian biota, 286, 320, 404, 689 Sahul, 121, 122, 391, 813, 1096 See also Biogeography; Tectonic history, and endemism Biogeography, 28–29, 196–205, 207–223, 228–245, 285–290, 339, 340, 391, 517, 519–521, 552–553, 655, 657–664, 675, 730, 791, 1000, 1218, 1349 theory of island, 785, 1264 See also Speciation; Species radiations; Vicariance BirdLife International/EBA and IBA, 489, 490, 675, 1202, 1240–1241, 1246–1247, 1315, 1327

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Bird’s Head. See Vogelkop Bird’s Head Alliance, 1190, 1192 Bird’s Neck, 5, 6, 48, 49, 69, 147, 150, 156, 159, 197, 200, 202, 203, 213, 488, 568, 651, 652, 690, 692–702, 711, 715, 730, 732, 737, 738, 745, 872, 1246 Birds of Paradise (Paradisaeidae), 7, 10, 14, 16, 19, 199, 201, 202, 204, 396, 406, 413, 655, 658, 660, 661, 670–671, 1359, 1400, 1404, 1405 trade, 11, 23, 61, 658, 675, 1161, 1219, 1285, 1286, 1288, 1313 Bishop Museum Field Station. See Wau Ecology Institute Bismarck Arc, 142, 152, 155, 158, 198 Bismarck Archipelago, 18, 27, 32–33, 37, 158, 208, 221, 269, 292, 382, 386, 387, 404, 416, 419, 420, 421, 424, 442, 467, 479, 541–544, 552, 564, 565, 573, 579, 580, 582, 584, 585, 628, 710, 720, 1200, 1314 BKSDA. See Natural Resources Conservation Bureau (BKSDA) Black walnut. See Dracontomelon Blennies (Blenniidae), 638, 642, 1419 Blue-eyes (Pseudomugilidae), 230, 239, 243, 646, 866, 868, 875, 878, 879, 880, 1407, 1417 Bogs, 338–339, 874, 876, 980, 1039, 1042– 1045. See also Swamps Bokondini, 84, 736 Bomberai Peninsula, 5, 6, 8, 9, 46, 142, 187, 197, 209, 213–214, 229–231, 270, 276, 279, 291, 569, 603, 645, 648, 649, 659, 661, 861, 917, 920, 1244, 1246. See also Kumawa Mts; Onin Peninsula Bony tongues (Scleropages: Arowana, Saratoga), 646, 650, 886, 887, 1164, 1333, 1416 Borneo, 277, 280, 324, 325, 326, 335, 352, 379, 382, 383, 386, 394, 396, 400, 424, 437, 439, 443, 462, 475, 481, 483, 484, 545, 566, 567, 575, 584, 640, 658, 659, 689, 766, 794, 800, 947, 965, 966, 968, 983, 995, 1104, 1239, 1350, 1357 Borobudur Market, 1286, 1287 Bougainville Island. See Solomon Islands Bougainville Mts (near Oinake´), 215, 885, 910 Bowerbirds (Ptilonorhynchidae), 3, 201, 203, 406, 655, 658, 660, 671 Bowutu Mts, 217, 958 Bowutu Terrane, 153 Brazil, 73, 307, 313, 410, 545, 1366 Breadfruit, 406, 951, 1360 British Petroleum, 1155, 1191, 1192, 1212, 1213

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index / 1443 Brugiera, 9, 264 Brush Turkey. See Megapodiidae Bryophytes, 320–332 biology/ecology, 320, 326–329, 954, 963, 968, 981, 982, 1004, 1006, 1028, 1030, 1037, 1039, 1042, 1045, 1046, 1048 collections/surveys, 35, 42, 69, 73, 85, 86, 91, 106, 320–321, 324, 330 conservation, 329–331 endemism, 322, 324, 326, 350 taxa/distribution, 321–326 See also Hepaticae; Mosses Bucerotidae. See Hornbills Bufo (introduced), 586, 893, 1365, 1366, 1367, 1410 Buginese, 1115 Buka. See Solomon Islands Bulbophyllum. See Orchids Bulolo (PNG), 270, 305, 325, 347, 885, 980, 985, 991, 992 Bulolo River, 215, 216 Bunaken Marine Reserve (Sulawesi), 644 Bupul Nature Reserve, 1330, 1431 Buru. See Maluku Butonese, 1115 Butterflies. See Lepidoptera Butterflyfishes (Chaetodontidae), 638, 639, 642, 781 Cacao. See Cocoa Campnosperma, 9, 879, 910, 911, 913, 917, 922, 923, 928–930, 933, 935, 936, 940, 941, 949, 951, 954, 955 Cane Toad. See Bufo Cape Nelson Peninsula, 209, 210, 219, 244 Cape York, 59, 221, 580, 811 Carallia, 9 Carcharinus. See Sharks, freshwater Cardinalfishes (Apogonidae), 239, 241, 638, 642, 644, 650, 882, 1419 Caribbean, 160, 346, 500, 548, 552, 781, 787, 788 Caroline Islands, 34, 547, 552, 580 Caroline Plate, 137, 145–148, 154, 159, 774 Carp (Cyprinus), 651, 871, 887, 889, 890, 1217 Carstensz Glacier, 74, 123, 183, 190–193, 1049, 1050, 1223. See also Glaciers Carstensz Meadow, 1027, 1032, 1033, 1036, 1038–1041, 1043, 1045 Cash crops, 521, 1125, 1129, 1139, 1140, 1141, 1142, 1143, 1156, 1170, 1331, 1331. See also Cashew nuts; Cloves; Cocoa; Coconuts; Coffee; Food crops;

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

Fruits; Nutmeg; Rubber; Tobacco; Vegetables Cash economy/income, 812, 893, 1111, 1144, 1331. See also Cash crops Cashew nuts (Anacardium occidentale), 1139, 1142, 1156, 1157 Cassava (Manihot esculentus), 1128, 1129, 1130, 1137, 1139, 1144, 1156, 1354 Cassowaries, 10, 199, 249, 271, 283, 382, 383, 384, 385, 432, 466, 658, 665, 667, 674, 676, 678, 841, 1219, 1237, 1386, 1404 trade, 1285, 1286, 1287 Castanopsis, 9, 261, 314, 470, 946, 959, 978, 979, 980, 981, 982, 984, 985, 988, 990– 993, 996, 998, 1000, 1090 Casuarina, 6, 248, 903 904–905, 906, 907, 943, 950, 951, 952, 958, 959, 965, 968, 969, 971, 984, 985, 989, 990, 995, 997, 1025, 1092, 1093 Casuarina Coast. See Asmat (Casuarina) Coast Casuarius. See Cassowaries Catfishes (ariid, plotosid), 234, 239, 240, 241, 645–651, 863, 864, 887, 889, 1164, 1406, 1416, 1417. See also Walking catfish (Clarias) Cathaysian biota, 246 Catholics. See Christianity; Christian missionaries Cattle, 119, 778, 886, 1330 Caves, 84, 222, 249, 663–664, 720, 757, 776– 777, 881, 1064–1081. See also Archeological sites, specific Cenderawasih Bay, 3, 156, 157, 179, 181, 186, 187, 197, 200, 261, 325, 479, 489, 505, 534, 537, 568, 574, 591, 637, 659, 660, 719, 724–730, 772, 773, 775, 792, 802, 809, 810, 813, 1077, 1106, 1107, 1110, 1120, 1121, 1206, 1271. See also Biak/ Supiori; Numfoor; Yapen Cenderawasih Bay (people of/culture), 112, 116, 118, 119 Cenderawasih Bay National Park, 809, 814, 816, 1206, 1262, 1271, 1317, 1426, 1433 Cenderawasih University (UNCEN), 74, 81, 84, 85, 1108, 1245, 1312, 1315 Cenozoic, 248, 480, 481, 566. See also Tertiary; Quaternary Central Cordillera (Merauke Range), 4, 5, 6, 7, 9, 12, 47, 49, 177, 178, 183, 186, 187, 197, 200, 201, 202–204, 257, 296, 337, 431, 656, 660, 661, 665, 692–702, 706– 708, 711, 712, 713–718, 719, 720, 721, 723, 724, 728, 729, 732, 733, 735–738,

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1444 / index Central Cordillera (Merauke Range) (cont.) 744, 745, 746, 747, 979, 1095, 1131, 1151, 1199, 1246, 1260, 1268, 1311. See also Jayawijaya Range; Sudirman Range Cervus timoriensis. See Rusa Deer Charles Louis Mts (Central Cordillera), 47, 487, 489, 599, 601, 711, 738 Chimbu, 1128 China, 339, 344, 346, 386, 419, 473, 573, 667, 677, 886, 1115, 1160, 1277, 1285 Chiroptera. See Bats Christensen Research Institute (Madang), 79, 96, 504 Christianity, 78, 112, 113, 1099, 1116, 1117 Christian missionaries, 11, 21, 22, 23, 24, 26, 30, 31, 33, 35, 36, 39, 42, 43, 44, 54, 56, 65, 66, 90, 113, 249, 567, 886, 1103, 1106, 1107, 1117 Cicadas, 160, 202, 518, 520, 532–537 CITES/Appendixes, 629, 632, 633, 1277, 1291, 1299, 1350 Climate, 7, 92, 165, 172, 177–194, 264, 1242, 1327 and agriculture/human habitation, 182, 187, 188, 189, 193, 1087–1088 and biodiversity, 483, 566, 587, 640, 658, 983, 996, 1003, 1048, 1054, 1094 history, 126, 248, 255–260, 263–265, 1000, 1003, 1089, 1091, 1094 regulation as ecosystem service, 760, 761, 1201, 1224 See also Climate change; Cyclones; Drought; Monsoon; Rainfall; Solar radiation; Water balance; Winds Climate change, 190–193, 255, 511, 761, 764, 792, 802, 814, 817, 1202, 1205, 1206, 1216, 1221–1223 glacial changes/retreat, 190–193, 257, 1223 See also Climate, history; Climate, regulation; El Nin˜o events; Glaciers; Sea level, changes in Climax species/communities, 309, 397, 521, 903, 935, 940, 948, 952, 953, 956, 957, 958, 995, 1013, 1046. See also Seral vegetation Climbing Gourami/Perch (Anabas), 651, 889, 892, 1218, 1421 Cloud forest, 404, 416, 980, 996, 1004. See also Mosses/moss forest Cloudy Mountains, 26, 209, 210, 219, 244 Cloves (Eugenia aromatica), 1139, 1141, 1142, 1156, 1161 Cocoa (Theobroma cacao), 670, 1103, 1137, 1141, 1142, 1156, 1157

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Coconut (Cocos nucifera), 29, 37, 1129, 1137, 1139, 1360 as cash crop, 1141, 1142, 1156 copra, 905, 1103, 1142 plantations, 599, 1103, 1156–1157 wild, 359, 903, 905–907, 946, 1137 Coconut beetle, 55, 97 Cocos. See Coconut Code of Criminal Litigation (KUHAP), 1284, 1288, 1293, 1298 Coevolution, examples of, 368–369. See also Pollination; Seed dispersal; Symbiosis Coffee, 589, 670, 1103, 1129, 1141, 1142, 1156, 1157 Coleoptera, 10, 207, 211, 230, 231, 233, 234, 236, 237, 238, 240, 352, 375, 412, 413, 519, 520, 539, 549, 620, 859, 863, 865, 868, 869, 871, 1006, 1076, 1077, 1286, 1287 collections, 30, 40, 46, 47, 62, 70, 83, 97, 98, 99, 101 Colocasia. See Taro Colombia, 142, 435, 547 Columbids (pigeons, doves), 10, 202, 203, 352, 382, 413, 432, 460, 466, 655, 661, 668, 676, 1219, 1268, 1286, 1288, 1328, 1359, 1390, 1391, 1404–1405 Community cooperatives (kopermas), 1160, 1161, 1176, 1209, 1280, 1281, 1282, 1285 Congo Forest, 1200, 1240 Conifers. See Agathis; Araucaria Conservation designations Ecoregions (WWF), 1240, 1245–1246 Endemic Bird Areas (BirdLife), 1240–1241, 1246–1247 Hotspots (CI), 764, 1200,1239–1240 Important Bird Areas (BirdLife), 1240– 1241 Wilderness Areas (CI), 330, 756, 1200, 1201, 1239–1240, 1272, 1314, 1365 Conservation education, 331, 522, 651, 815, 816, 892, 1106–1109, 1218, 1245, 1254, 1307–1308, 1319, 1349, 1434–1436 Conservation International (CI), 75, 1149, 1184, 1241–1245, 1247 biodiversity surveys, 207, 504, 891, 1219, 1286 on enforcement, 1286–1287, 1298–1299, 1301, 1303 FKPTP, 1173 See also Conservation designations; Conservation Priority Setting (CPS); Rapid Assessment Programs (RAP) Conservation priority setting (CPS), 487, 599, 1232–1239, 1241–1246, 1266, 1267,

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index / 1445 1313–1314. See also Biodiversity; Ecosystem services; Keystone resources/species Convention on International Trade in Endangered Species of Wild Flora and Fauna. See CITES Coordinating Ministry of Police and Security, 1288, 1289, 1292 Copal, 65, 80, 962, 994, 1103, 1374 Copper, 559, 849, 959, 1105, 1149, 1150, 1152, 1153, 1154, 1211. See also Freeport McMoRan Copra. See Coconut Coral Sea, 145, 151, 152, 504, 541–544, 552, 783, 848 Coral Triangle, 495–498, 499, 501, 502, 503, 508, 637, 639, 782–785, 790, 1200, 1216 Corals/coral reefs, 10, 62, 174, 247, 263, 308, 467, 495–505, 509–511, 591, 637, 639– 641, 644, 755, 759–760, 771–795, 800, 801, 803, 804, 807, 810, 811, 813, 814, 816, 881, 882, 902, 903, 904, 1200, 1208, 1216, 1223, 1237, 1239, 1312 Crabs, 211, 216, 499, 507, 508, 511, 557, 589, 591, 597, 781, 812, 838, 839, 841, 842, 843, 844, 845, 902, 919, 1074, 1354. See also Crustaceans; Hermit crab Cretaceous, 138, 142, 145, 149–152, 160, 306, 335, 401, 473, 800 Crocodiles, 507, 591–592, 840, 841 farming, 1324, 1333 fossil, 249 taxa/distribution, 480, 482, 565, 567, 574– 575, 1405 trade/threats, 603, 604, 1059, 1161, 1219, 1313, 1333 Crustaceans, 367, 497, 507, 508, 511, 523, 597, 619, 649, 781, 791, 815, 839, 846, 859, 863, 881, 885, 1074, 1078, 1354 stomatopods, 499–502 wood-boring, 539, 546–549, 551–557, 559 See also Crabs; Fisheries; Lobster; Shrimp Cryovegetation, 157, 184, 1049, 1051–1052. See also Algae; Alpine and subalpine ecosystems; Glaciers Cryptocarya, 79, 913, 920, 935, 936, 938, 941, 953, 957, 973, 981, 982, 984, 985, 987, 988, 991, 998 Cryptogams, 47, 63, 89, 90, 92, 290, 309. See also Ferns; Fungi; Mosses CSIRO, 98, 102, 1241 expeditions and surveys, 76, 77–78, 87, 88, 95, 1054, 1327 Cultivation. See Agriculture, traditional; Shifting cultivation

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

Cupressaceae, 345–346, 947, 980, 996, 998, 1029, 1037, 1042. See also Papuacedrus Currents, water/ocean, 177–182, 539, 552, 640, 641, 642, 773, 776, 779, 785, 788, 803, 808, 860, 882. See also Winds Cuscuses (Phalanger, Spilocuscus), 199, 200, 202, 252, 665, 690, 695, 696, 710, 712, 714–715, 726, 728, 730, 731, 733, 734, 735, 745, 1078, 1286, 1287, 1359, 1404, 1422 Customs, 1288, 1292 Cycadaceae/Cycas, 344–345, 903, 983 Cyclones (typhoons), 181, 190, 496, 785, 786, 806, 813, 946, 949, 955, 956, 1221, 1351 Cyclops Mts, 5, 6, 49, 153, 154, 155, 169, 173, 186, 198, 201, 209, 210, 215, 221, 233, 261, 269, 385, 387, 487, 489, 521, 589, 599, 600, 601, 603, 604, 645, 659, 660, 711, 712, 717, 724, 728, 729, 730, 873, 945, 949, 959, 960, 1090, 1094, 1138, 1200, 1209, 1233, 1246, 1268, 1270, 1426 Cyperus, 902, 942, 950, 967, 1057 Cyrtandra, 293, 296, 998, 1008 Dacrycarpus, 9, 248, 252, 258, 346, 978, 997, 998, 1007, 1012, 1018–1021, 1027–1036 Dacrydium, 9, 252, 346, 959, 967–971, 973, 998, 1003, 1009, 1012, 1036 Dactylopsila. See Possums and gliders Dams, 872, 886, 1211 as biodiversity threat, 651, 666, 815, 849, 1211 Mamberamo dam project, 1170, 1182, 1184, 1185, 1186, 1215 Damselfishes (Pomacentridae), 638, 639, 640, 642, 643, 644, 882 Damselflies. See Odonata Dani, 11, 108, 109, 110, 111, 114, 116, 117, 118, 119, 1127, 1131, 1132, 1133, 1134, 1136, 1141, 1142, 1143, 1144 Darai Plateau, 150, 218 Daru (Trans-Fly), 182, 541, 847, 1330, 1333, 1365, 1416, 1421 Darwin, Charles, 19, 656 theory of reef evolution, 771–772 Dasyurids (dunnarts, dasyures, quolls), 202, 248, 483, 692, 708, 710, 711–713, 723, 726, 731, 734, 744, 1422 Deer. See Rusa Deer Deforestation. See Habitat loss, deforestation Dendrobium. See Orchids Dendrolagus. See Kangaroos and wallabies D’Entrecasteaux Islands, 76, 159, 207, 209, 210, 211, 219, 244, 386, 585, 966

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1446 / index Development. See Human/economic development Development planning, 1171, 1172, 1231, 1233, 1243, 1324, 1338, 1342 Biodiversity Action Plan (1993), 285 development plans (REPELITA), 1104, 1107, 1112 National Development Planning Agency (BAPPENAS); 1098, 1104, 1113, 1118, 1313 Provincial Development Planning Bureau (BAPPEDA), 1118, 1243, 1313 Provincial Environmental Impact Management Agency (BAPPEDALDA), 1206, 1362 Diamond, Jared, 3, 11, 84, 95, 670, 745 Digul River, 5, 9, 150, 217, 218, 487, 488, 569, 599, 601, 603, 645, 648, 733, 778, 827, 858, 864, 878, 888, 917, 1247, 1327, 1417, 1419, 1420 Dioscorea. See Yams Diospyros, 9, 350, 826, 833, 879, 904, 905, 906, 911, 913, 922, 928, 929, 933, 935, 936, 941, 948, 953, 954, 956, 958, 963, 968, 969, 971, 972, 973, 982, 984, 985, 987, 989 Diptera (flies), 38, 99, 101, 104, 352, 367, 375, 425, 451, 480, 482, 516, 519, 520, 522, 859, 863, 869, 871, 873, 876, 880, 1077 Diseases, human, 112, 128, 189, 522, 533, 838, 886, 1109 BSE/nvCJD/Kuru, 89, 99 HIV/AIDS, 1098, 1109, 1319 from macaques, 1360–1361 malaria, 40, 96, 522, 645, 667, 838, 892, 1109, 1110, 1357 vectors, 521, 522–523, 886 See also Medical entomology; Mosquitoes Diseases, nonhuman, 678, 1222, 1237, 1263, 1278, 1357, 1360, 1366, 1367 coral/marine, 781, 787, 793 plant, 264, 760, 829, 1004, 1127, 1278 See also Agricultural and forestry pests Dispersal, by wind pollination, 405, 459, 465, 1004, 1006 seeds, 357, 377, 392, 402, 432, 460 See also Currents Disturbed habitat, 130, 345, 350, 353, 363, 367, 381, 385, 408, 468, 522, 557, 599, 677, 711, 720, 803, 829, 892, 923, 951, 953, 956, 958, 965, 980, 985, 990, 991, 995, 1004, 1014, 1030, 1204, 1217, 1351. See also Fire; Pioneer species; Roads; Secondary vegetation; Succession; Undisturbed habitat

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Dobo. See Aru Islands Dogs, 632, 722, 1095, 1330 hunting, 722, 1356 ‘‘singing,’’ 722 Dolok (Kimaam/Kolepom/Yos Sudarso) Island, 6, 7, 49, 62, 114, 521, 810, 1055, 1138, 1139, 1271, 1426, 1432 Dorcopsis. See Kangaroos and wallabies Dracontomelon (inc. black walnut), 10, 949, 953, 957 Dragonflies. See Odonata Drought, 8, 170, 189, 287, 328, 589, 649, 760, 829, 946, 964, 965, 1004, 1025, 1091, 1093, 1127, 1143, 1203, 1221, 1237. See also El Nin˜o events; Water balance Drymys, 968, 970–972, 997, 998, 1000, 1001, 1008, 1018, 1028, 1029, 1033, 1036, 1037, 1038 Dugong, 30, 801, 802, 803, 804, 805, 807, 809, 813, 814, 815, 901, 1404 Dysoxylum, 930, 952, 984 Early humans. See Human habitation; archeological entries East Papua Composite Terrane, 144, 218, 487 Echidnas (Tachyglossidae, Zaglossus), 46, 200, 201, 692, 710–711, 712, 730, 731, 732, 734, 737, 738, 743, 1422 Echymipera. See Bandicoots Economic development. See Human/economic development Ecoregions. See Worldwide Fund for Nature (WWF) Ecosystems. See Alpine and subalpine ecosystems; Bogs; Caves; Cloud forest; Corals/ coral reefs; Cryovegetation; Glaciers; Gondwanan biota; Grasslands; Heath; Mangroves; Monsoon forest; Moss/moss forest; Peat, swamp; Savanna; Seagrass; Swamps/swamp forest Ecosystem services, 320, 327, 329, 515, 670, 753, 760–762, 765, 801, 833–839, 1201, 1202, 1204, 1206, 1208, 1234, 1245, 1276 climate regulation, 760, 761, 1201, 1224 See also Pollination; Seed dispersal Ecuador, 142, 435, 589, 982, 1049 Education, 74, 79, 81, 86, 98, 1098, 1100, 1105, 1106–1109, 1111, 1112, 1113, 1114, 1120, 1165, 1169, 1171, 1172, 1190, 1192, 1317, 1319, 1331. See also Conservation education Eipo, 74 Elaeocarpus, 274, 292, 381, 382–386, 387, 930, 947, 952, 952, 963, 967, 980, 981, 982,

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index / 1447 985, 986–988, 991, 992, 996, 998, 999, 1001, 1008, 1009, 1018, 1021 Elapidae. See Sea snakes; Snakes Eleocharis, 942, 1056, 1057, 1061, 1334 Eleotridae. See Gudgeons El Nin˜o events, 8, 170, 177, 181–182, 188– 189, 194, 497, 946, 953, 1004, 1030, 1091, 1093, 1143, 1221, 1222. See also Climate change; Drought Emoia, 486, 564, 578, 579, 593, 1413 Employment/labor, 1109, 1110–1112, 1113, 1114, 1120, 1151, 1152, 1154, 1157, 1158, 1167, 1184, 1186 of Papuans/migrants, 1098, 1103, 1104, 1110, 1111, 1115, 1152 Enarotali, 1130, 1151, 1178, 1215 Enaratoli Strict Nature Reserve, 1268, 1270, 1426 Endangered and threatened species, 330, 665, 672, 676, 678, 711, 757, 760, 787, 801, 805, 887, 1202, 1233, 1234, 1235, 1240, 1243, 1266, 1267, 1277, 1285, 1286, 1291, 1300, 1304, 1313, 1314, 1324, 1329, 1350 Endemism, 207, 208, 644 animals, 160, 201, 207, 208–220, 228–245, 479, 487–490, 511, 532–537, 599–603, 628–631, 644–645, 648, 649, 651–652, 691, 710, 723, 728, 730, 1064 and biogeography, 264, 280, 281, 290, 307, 483, 520–521, 755 and conservation, 788–792, 1217, 1230, 1232, 1234, 1235, 1237–1238, 1239, 1241, 1242, 1243, 1314, 1365 plants, 274–280, 292, 316, 322, 330, 339, 341, 351, 382, 404, 442–443, 462, 1311 and tectonic history/geology, 158, 160, 196, 201, 202, 220–223, 288–289, 320, 566 See also Biodiversity hotspots; Speciation Enewetak Atoll (Marshall Islands), 771, 772 Enforcement, 643, 644, 678, 679, 815, 887, 1186, 1189, 1214, 1245, 1263, 1277, 1278, 1279, 1280, 1284, 1286–1309, 1333, 1335, 1340 Eocene. See Tertiary, Eocene Erosion, 8, 165, 166, 168, 169, 198, 255, 263, 308, 329, 566, 644, 759, 814, 817, 847, 849, 865, 933, 958, 960, 982, 1013, 1087, 1089, 1206, 1216 protection against, 760, 761, 825, 1254 Ertsberg mine. See Freeport McMoRan, Ertsberg mine Etna Bay, 6, 200, 213, 216, 366, 505, 637, 645, 648, 649, 663, 868, 890, 962, 1247

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

Eucalyptus, 25, 33, 263, 264, 429, 431, 522, 626, 902, 911, 949, 951, 959, 980, 1058 Eugenia. See Cloves Evolution. See Speciation; Species radiations; Vicariance Exotic species. See Introduced species Extractive industries. See Fisheries; Logging; Mining; Oil and gas; Plantations Fakfak, 3, 5, 7, 113, 158, 209, 210, 213, 214, 231, 325, 487, 505, 520, 542, 543, 569, 599, 601, 637, 659, 690, 711, 728, 732, 809, 1066, 1068, 1071, 1073, 1074, 1075, 1076, 1078, 1102, 1106, 1109, 1110, 1142, 1151, 1156, 1157, 1158, 1160, 1213, 1220, 1244, 1246, 1261, 1268, 1270, 1427, 1431 Fam Islands, 792, 808 Fergusson Island. See D’Entrecasteaux Islands Ferns (pteridophytes), 42, 71, 89, 246, 258, 261, 291, 315, 331, 335–341, 442, 824, 869, 903, 911, 934, 935, 940, 942, 943, 948–954, 956, 963, 977, 980–985, 990, 991, 992, 995, 997, 998, 1008, 1012, 1018, 1021, 1028, 1029, 1030, 1036, 1037–1042, 1046, 1091, 1093 Fertilizer, 171, 172, 174, 884, 1208, 1216 in traditional agriculture, 1133, 1134, 1136 Ficus, 292, 296, 396, 404, 405, 406, 407, 468, 558, 668, 677, 904, 914, 918, 923, 925, 928, 930, 938, 943, 947, 948, 949, 951– 958, 963, 984, 985, 989, 990, 995, 1014, 1015, 1356 and fig wasps, 404, 405–406 Fiji, 142, 145, 146, 149, 313, 346, 382, 411, 424, 473, 546, 574, 639, 788, 790, 1107 Finisterre Mts, 154, 209, 210, 215, 234, 468, 660 Fire as ecological disturbance, 8, 193, 260, 432, 919, 946, 947, 953, 1004, 1005, 1018, 1025, 1030, 1056, 1058, 1060, 1061, 1089, 1093, 1094, 1143, 1203, 1206, 1221, 1222, 1237, 1334 human-caused, 8, 760, 794, 814, 1004, 1005, 1037, 1055, 1061, 1126, 1143, 1203, 1208, 1221, 1329, 1334 management, 1061, 1208, 1334, 1335, 1337, 1341, 1345 succession/survivors/resistance, 288, 335, 386, 956, 957, 958, 990, 995, 1013, 1037, 1039, 1040, 1056, 1057, 1330 See also Archeological evidence, fire; Disturbed habitat; Regeneration; Secondary vegetation; Seral vegetation; Succession

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1448 / index Firewood, 920, 995, 1259, 1333 Fish. See Fisheries; Fishing; Freshwater fish; Introduced species, taxa, fish; Reef fish; Sharks Fisheries, 801, 805, 814, 815, 833, 842, 845– 847, 849, 1150–1161, 1163–1164, 1165 lobster, 793, 846, 1164 shrimp (prawn), 813, 846, 850, 1163, 1164 Fishing, 643–644, 787–788, 794, 845, 849, 888, 902, 1216 blast/poison/destructive, 427, 643, 792– 793, 794, 1216 commercial, 12, 802, 814, 1111, 1114 (see also Fisheries) illegal/overfishing, 643–644, 651, 788, 792, 794, 814, 846, 1214 subsistence/traditional, 114, 119, 643, 788, 813, 814, 845, 888, 1129–1130, 1137, 1138, 1340 (see also Hunting, subsistence) Fish poison, 427 Flannery, Tim, 3, 95, 100, 484, 724, 732 Flies. See Diptera Fly River, 150, 216, 217, 218, 382, 483, 489, 532, 648, 656, 661, 714, 746, 824, 827, 829, 830, 831, 832, 838, 843, 845, 848, 1247, 1327, 1404–1409, 1416–1421, 1422, 1424 Foja Mts, 3, 5, 154, 155, 159, 198, 201, 209, 210, 214, 216, 232–233, 261, 487, 489, 599, 601, 603, 659, 660, 690, 711, 718, 728, 729, 730, 732, 745, 873, 1087, 1200, 1242, 1244, 1246, 1268, 1269, 1271, 1428, 1432 Fold Belt, 150, 152–154, 197, 198, 213, 871 Food chains/webs, 632, 649, 666, 779, 780, 833, 842–845, 847, 885, 901, 1072–1073, 1078, 1202 Food crops. See Bananas; Breadfruit; Cash crops; Cassava; Coconut; Fruits; Manioc; Peanuts; Potato; Rice; Sugar cane; Sweet potato; Taro; Vegetables; Yam Food security, 1143–1144 Foreign investment, 1150–1152, 1161 Forest loss. See Habitat loss, forest loss Forestry Bureau, 1254 Forestry Office of Papua Province, 1177, 1255, 1256, 1287, 1292, 1298, 1299, 1301, 1302, 1303, 1316 Forestry Service (Boswezen), 73, 79, 80 Forests, protected Limited Production Forest (Hutan Produksi Terbatas), 1158, 1258 Permanent Production Forest (Hutan Produksi Tetap), 1158, 1258

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Production Forest (Hutan Produksi), 1158, 1160, 1173–1176, 1178, 1180, 1206, 1208, 1209, 1279–1283, 1292, 1302, 1315 Protected Forests (Hutan Lindung), 1174, 1206, 1251, 1254–1255, 1258, 1315 Fossils, 1087 animal, 30, 99, 100, 125, 248–251, 574, 575, 584, 646, 711, 717, 722, 723, 730, 731, 736, 1074, 1090 (see also Archeological sites) marine, 246–247, 774, 775, 881 plant, 100, 149, 246, 247–248, 335, 404, 473, 800, 1089 Fragmentation. See Habitat loss, fragmentation Frederik-Hendrik Island. See Dolok Freeport McMoRan, 384, 810, 814, 849, 1105, 1141, 1150, 1152–1154, 1191–1192, 1210–1211 and fieldwork, 75, 81, 82, 153, 977 Ertsberg mine, 81, 1030, 1152, 1211 Grasberg mine, 849, 869, 884, 885, 889, 1037, 1046, 1081, 1111, 1115, 1117, 1152, 1154, 1167, 1168, 1211 Freshwater fishes blue-eyes (Pseudomugilidae), 230, 239, 243, 646, 866, 868, 875, 878, 879, 880, 1407, 1417 bony tongues (Scleropages: Arowana, Saratoga), 646, 650, 886, 887, 1164, 1333, 1416 catfishes (ariid, plotosid), 234, 239, 240, 241, 645–651, 863, 864, 887, 889, 1164, 1406, 1416, 1417 (see also Walking catfish (Clarias)) gudgeons (Eleotridae), 231, 232, 234, 237– 241, 243, 244, 646–650, 863, 865, 871, 877, 878, 879, 880, 882, 887, 888, 1077, 1409, 1419, 1420 (see also Gobies) lungfish (Neoceratodus), 646 rainbowfish (Melanotaeniidae), 210, 228– 236, 239–242, 646, 647, 648, 649, 650, 651, 863, 868, 871, 872, 873, 875, 887, 891, 892, 1417 See also Fishing Freshwater sharks (Carcharinus), 6, 888, 1405, 1416 Freycinetia, 293, 296, 991, 998, 1008, 1012 Frogs biology/ecology, 203, 587–589 endemism, 200, 201, 479, 564, 566, 600, 601, 602, 603

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PAGE 1448

index / 1449 introduced, 586, 893, 1365, 1366, 1367, 1410, 1411 as prey, 590, 592, 595, 596, 597, 598, 600, 621, 627, 1354, 1366, 1367 surveys/collections, 70, 94, 95, 484, 485, 605, 1311, 1312, 1366 taxa/distribution, 10, 204, 480, 482, 483, 489, 565, 567, 569–574, 1410–1411 conservation/trade, 604, 605, 1368 Fruits, as human food, 1003, 1130, 1137, 1139, 1144, 1352, 1354, 1359, 1360 as cash crop, 1139, 1142, 1156 Fungi, 165, 539, 678, 843, 844, 1005, 1006, 1072, 1143 collections, 42, 58, 78, 85, 86, 87, 89, 91, 100 See also Lichens (lichenized fungi) Gag Island, 18, 46, 705, 706, 719, 733, 749, 793, 1152 Gahavisuka Provincial Park, 90, 314, 315, 316, 325 Galley Reach, 92, 547, 548, 830, 835, 837 Gam Island, 22, 210, 228, 878 Gap analysis, 1236, 1266, 1269 Gauttier Mts. See Foja Mts Gebe Island, 18, 22, 46, 719, 733 Gecko, 489, 575, 577, 586, 592–593, 600, 1353, 1367–1368, 1412 Geelvink Bay. See Cenderawasih Bay Geological eras, 139. See also Cenozoic; Fossils; Mesozoic; Paleozoic; Quaternary; Tertiary Geological formations Arafura Shelf, 181, 255, 260, 264 Fold Belt, 150, 152–154, 197, 198, 213, 871 Mobile Belt, 150, 153, 154–158, 197, 200 Ophiolite Belt, 142, 153, 159, 222 Sorong Fault, 148, 157, 159, 198, 199, 200, 213 Sunda Shelf, 359, 566, 585, 689 See also Arcs; Australian craton; Island arcs; Tectonic plates; Terranes Geology. See Geological formations; Igneous rock; Ophiolites; Sedimentary rock; Tectonic history, and endemism; Ultramafics; Volcanics Gigi Lake. See Anggi Lakes Gingers (Zingiberaceae), 379, 473–475, 934, 940, 948, 952, 956, 998, 1008, 1012 Giragandak Cave, 1071, 1073, 1074, 1075, 1076 Gita Lake. See Anggi Lakes

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Glacial periods, 3, 5, 8, 53, 74, 125, 137, 155, 169, 190–193, 249, 251, 255–256, 257– 260, 261, 263, 264, 282, 388, 443, 519, 566, 568, 640, 647, 765, 825, 870, 962, 1000, 1025, 1038, 1041, 1045–1052, 1090, 1091, 1094, 1223 Glaciers, 3, 5, 8, 53, 74, 123, 169, 183, 190– 193, 257, 260, 282, 388, 568, 765, 870, 1041, 1045, 1048, 1049–1052, 1091, 1223 Carstensz Glacier, 74, 123, 183, 190–193, 1049, 1050, 1223 Meren Glacier/Valley, 190–192, 1033, 1036, 1044–1051 Northwall Firn (glacier), 190–193, 1048 Glassfishes (Ambassis), 649, 868, 878, 882, 1164, 1407, 1418 Gliders (Petauridae). See Possums and gliders Global warming. See Climate change Glomera. See Orchids Gnetum, 344, 347, 914, 936, 938, 949, 951, 953, 957, 984, 993 Gobies (Gobiidae), 229–242, 638, 642, 644, 646, 647, 648, 650, 781, 860, 862, 863, 864, 865, 875, 882, 1164, 1408–1409, 1419, 1420. See also Gudgeons Gold, 26, 38, 849, 884–885, 889, 1081, 1103, 1105, 1115, 1117, 1151–1154, 1210, 1211. See also Freeport McMoRan Golo Cave, 123 Gondwanan biota, 88, 221, 246, 252, 286, 320, 322, 381, 404, 570, 571, 574, 689, 980 Gondwanan land forms, 246, 287, 288, 659 Gondwanan plant taxa. See Dacrycarpus; Dacrydium; Nothofagus Goura. See Columbids Grasberg mine. See Freeport McMoRan, Grasberg mine Grasslands, 9, 167, 173, 193, 249, 250, 264, 309, 315, 335, 338, 345, 371, 389, 440, 441, 442, 587, 593, 660, 664, 710, 711, 713, 714, 715, 716, 717, 722, 735, 754, 758, 942, 958, 959–960, 979, 980, 997, 1003, 1004, 1013, 1014, 1018, 1021, 1025, 1029, 1033, 1036, 1037–1040, 1042, 1044, 1045, 1046, 1047, 1054– 1063, 1087, 1089–1094, 1246, 1251, 1314, 1327, 1329, 1330, 1333, 1334, 1344, 1352. See also Alpine and subalpine ecosystems; Savanna Great Barrier Reef (Australia), 496, 510, 771, 773, 774, 778, 782, 783, 794 Green Sea Turtle, 574, 590, 591, 801, 803, 805, 811, 813, 815, 1286, 1405

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1450 / index Ground nuts. See Peanuts Groupers (Serranidae), 638, 643, 793, 794, 1216 Guam, 546, 596, 678, 783 Gudgeons (Eleotridae), 231, 232, 234, 237– 241, 243, 244, 646–650, 863, 865, 871, 877, 878, 879, 880, 882, 887, 888, 1077, 1409, 1419, 1420. See also Gobies Gulf of Papua, 541, 544–548, 553, 555, 558, 831, 835, 837, 838, 845–848 Gunung Mulu National Park (Borneo), 965 Habitat degradation. See Disturbed habitat; Erosion; Habitat loss; Pollution; Sedimentation; Siltation Habitat loss, 764, 1202, 1203, 1210, 1214, 1222, 1239 deforestation, 329, 523, 794, 814, 1078, 1081, 1159, 1170, 1214, 1276, 1311 forest loss, 765, 1170, 1177, 1205, 1206, 1207, 1215, 1223 fragmentation, 466, 1206, 1210, 1214, 1215, 1233, 1236–1237, 1348, 1349 See also Habitat degradation; Land conversion Halmahera Eddy (current), 180, 181, 808 Halmahera Island. See Maluku Harpy Eagle. See New Guinea Harpy Eagle Hawai’i, 140, 382, 495, 511, 522, 547, 639, 642, 678, 772, 774, 777, 778, 782, 783, 788, 789, 790, 881, 886, 892 Hawksbill Turtle, 574, 591, 793, 813, 1405 Headland, Thomas, 127, 128 Health/health services, 40, 116, 760, 1098, 1103, 1106, 1109–1110, 1111, 1112, 1113, 1114, 1165, 1169, 1171, 1182, 1190, 1192, 1214, 1319, 1331, 1332, 1360, 1434–1435. See also Diseases, human; Traditional medicine Heath (keranga), 338, 350, 408, 948, 962–975, 981, 985, 995, 1021, 1046–1047, 1048, 1049 Hepaticae (liverworts), 320–322, 325, 326, 328, 997, 1014, 1043 Herbarium Bogoriense, 801 Herbicides, 1208, 1216 Hermit crab, 497 Heteroptera, 207, 211, 213–220, 222, 228– 245, 480, 482, 518, 859, 863, 865, 868, 869, 871, 873, 877, 879, 880, 881, 1312 Hinduism, 113 HIV/AIDS, 1098, 1109, 1319 Holocene. See Quaternary, Holocene Hollandia. See Jayapura

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Honeyeaters (Meliphagidae), 201, 375, 447, 655, 658–659, 660, 668–669, 673, 1396 Hopea, 290, 539, 558, 911, 912, 914, 916, 917, 918, 920, 921, 923, 924, 925, 927, 930, 931, 935, 950, 954, 955, 959, 985, 986– 988, 990 Hornbills (Bucerotidae), 352, 382, 406, 413, 432, 659, 667–668, 671, 676, 1404 Horticulture, 1126. See also Agriculture Human/economic development, 282, 1098– 1100, 1103, 1105, 1106–1117, 1170, 1171, 1207, 1231, 1232, 1245, 1311, 1324 cash economy/income, 812, 893, 1111, 1144, 1331 (see also Cash crops) community development, and culture, 111–119, 1116 cultural characteristics conducive/detrimental to, 11, 116, 117, 118 food security, 1143–1144 foreign investment, 1150–1152, 1161 urbanization, 1111, 1113, 1170 See also Education, Employment/labor, Health; Human/economic development indices; Transmigration; specific sectors Human/economic development indices, 1098, 1106, 1113–1115 GDP, 1114, 1149, 1151, 1154, 1155, 1156, 1158, 1163, 1164, 1167, 1168, 1181, 1184 GNrP, 1168, 1184, 1185 Human Development Index (HDI), 1113– 1114 life expectancy, 1110 mortality/morbidity, 760, 1110, 1114 Human groups Amungme, 1105, 1115, 1131, 1136, 1140, 1141, 1143 Arso, 112, 114 Asmat, 11, 108, 109, 110, 111, 113, 114, 116, 117, 118, 119 Ayamaru, 111, 118, 125, 127, 887, 1120 Biak, 108, 109, 110, 111, 112, 113, 114, 117, 118 Buginese, 1115 Butonese, 1115 Cenderawasih Bay, 112, 116, 118, 119 Chimbu, 1128 Dani, 11, 108, 109, 110, 111, 114, 116, 117, 118, 119, 1127, 1131, 1132, 1133, 1134, 1136, 1141, 1142, 1143, 1144 Eipo, 74 Hupla, 1143 Imyan, 1116

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index / 1451 Javanese, 1115, 1120, 1140 Kamoro, 118, 1105, 1115, 1140 Kanum, 1331, 1332, 1339 Kapauku, 1131, 1134, 1136 Kimam, 109, 1138, 1139, 1140 Makassarese, 1115, 1286 Maluku (Moluccans), 11, 1110, 1112, 1120 Mappi, 110 Marind, 1131, 1332, 1339 Marind-Anim, 48, 109, 111, 113 Marori-Mengey, 1331, 1332, 1339 Maya, 108, 112, 114 Me, 109, 110, 111, 113, 114, 116, 117, 118, 119 Meybrat, 108, 109, 110, 111, 113, 116, 118 Mimika, 109, 110, 114 Miyanmin, 93 Moi, 113, 114, 1142 Muyu, 108, 109, 110, 111, 114, 116, 117, 118, 1106 Nduga, 1143 Nimboran, 110, 112, 117, 1106 Ormu, 1138, 1139, 1142 Sentani, 108, 110, 111, 112, 117, 1142 Waropen, 108, 109, 110, 111, 112, 113, 114, 118, 119 Yali, 82, 962, 1128, 1131, 1132, 1134, 1143, 1144 Yei, 1331, 1332, 1339 Yos Sudarso Bay, 109, 112, 117 Human habitation Austronesians, 108, 722, 1087, 1093–1094, 1128 and climate, 182, 187, 189 early/origins, 121–130, 1087, 1088–1094 Lapita, 16, 467, 722 and soils, 130, 760, 946, 995, 1126, 1129, 1135 See also Agriculture, traditional; Archeological evidence; Archeological sites Humboldt Bay, 337, 724, 959 Hunting, subsistence, 114, 127, 1003, 1037, 1057, 1111, 1113, 1116, 1129–1130, 1137, 1138, 1300, 1332, 1334 as biodiversity threat, 1203, 1210, 1214, 1243, 1283, 1286–1287, 1300, 1304, 1333, 1337, 1344 of birds/eggs, 671– 672, 674, 676 early human, 127–130, 251, 264, 1089, 1090, 1091, 1092, 1095 legislation, 1277, 1278, 1279, 1280, 1283, 1300 of macaques, 1357, 1360 of mammals, 10, 127–130, 711, 715, 729, 731, 1333

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in protected areas, 1254, 1259, 1265, 1337, 1338, 1334 and roads, 1177 using dogs, 722 See also Fishing, subsistence Hunting, recreational/tourism, 1254, 1259, 1340 Hunting, for trade/illegal/endangered and protected species, 1283, 1285–1287, 1300, 1304 Huon Peninsula, 121, 148, 215, 220, 221, 305, 322, 324, 326, 327, 328, 331, 662, 663, 729, 737, 774, 775 Hupla, 1143 Huxley, T. H., 21 Hydnophytum. See Rubiaceae Hymenoptera, 70, 406, 519, 620. See also Ants; Bees; Wasps Idenburg River. See Taritatu River Idenburg Valley, 984, 990, 991, 994 Igneous rock, 138, 153, 159, 168, 266, 903, 984, 1005 Ijomba Bog, 123, 258, 1094 Impatiens, 990 Imyan, 1116 India, 145, 149, 312, 339, 346, 352, 382, 386, 400, 411, 419, 436, 462, 541, 545, 547, 548, 575, 582, 584, 1160, 1170, 1251, 1285 Indian Ocean, 177, 181, 429, 495, 496, 497, 501, 502, 504, 511, 519, 552, 582, 640, 667, 784, 789, 800 Indo-Australian Plate, 137, 774, 1151 Indochina, 339, 352, 400 Indonesian Institute of Sciences (LIPI), 1141, 1164, 1245, 1277, 1312, 1313 Infrastructure, 72, 77, 270, 657, 817, 1100, 1103, 1104, 1109, 1111, 1112, 1114, 1116, 1150, 1151, 1156, 1170, 1171, 1184, 1190, 1203, 1213, 1214–1215, 1220, 1242. See also Air transport; Dams; Roads Introduced species, 678–679, 762, 861, 892, 893, 1202, 1203, 1216–1219, 1222, 1243, 1254, 1259 accidental, 849, 889, 1216, 1217, 1219 for food/commercial purposes, 871, 887, 892, 1216 for mosquito control, 889, 891, 892 prehistoric, 722 (see also Dogs; Pigs) as threat to biodiversity, 893, 1216–1219, 1364–1368

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1452 / index Introduced species, taxa birds, 654, 666, 678–679 fish, 646, 649, 650, 651, 869, 871, 874, 878, 887, 888–893, 1217–1218, 1330 (see also Carp; Climbing Gourami; Snakehead; Tilapia; Walking Catfish) herpetofauna, 586–587, 580, 678, 1365– 1368 (see also Bufo) insects, 522, 678, 1218 invertebrates, 869, 1218 mammals, 632, 678, 679, 722–723, 778, 886, 1218 (see also Cattle; Dogs; Macaques; Murids, rats; Pigs; Rusa Deer) plants, 22, 351, 405, 467, 940, 958, 1094, 1218, 1313, 1330, 1345 (see also Food crops) Intsia (inc. merbau), 10, 289, 467, 833, 879, 904, 905, 906, 911, 914, 922, 923, 928, 929, 930, 931, 933, 935, 937, 939, 941, 947, 949, 950, 953, 954, 955, 959, 983, 985, 987, 990, 1158, 1207, 1356 Invasive species. See Introduced species Ipomoea, 940. See also Sweet potatoes Islam, 11, 112, 113, 1112, 1117 Island arcs, 138, 140, 142–148, 150–154, 156, 158, 160, 197, 198, 201, 214, 215, 220– 221, 222, 481, 519, 532, 534, 566, 657, 658, 662, 1199. See also Arcs; Ophiolites IUCN, 675, 1203, 1253–1254, 1260, 1324, 1329, 1333, 1425 Red List, 678, 1202, 1350 See also Endangered and threatened species Jakarta, 678, 887, 1110, 1113, 1115, 1116, 1120, 1121, 1150, 1155, 1318, 1322 Jamaica, 546, 777, 787 Japan, 141, 160, 247, 346, 382, 435, 473, 510, 546–548, 573, 667, 778, 782, 783, 886, 1150, 1160 Japen Island. See Yapen Island Java, 138, 152, 277, 280, 289, 312, 324, 325, 384, 394, 400, 443, 544, 573, 640, 689, 778, 947, 979, 1153, 1161, 1239, 1251, 1286, 1318, 1322, 1350 Javanese people, 1115, 1120, 1140 Jayapura, 153, 214, 215, 263, 366, 459, 488, 505, 521, 537, 542, 545, 587, 604, 628, 630, 637, 651, 679, 722, 723, 729, 809, 811, 874, 887, 889, 958, 1059, 1101, 1102, 1109, 1110, 1111, 1114, 1117, 1119, 1142, 1151, 1156, 1157, 1158, 1161, 1179, 1209, 1210, 1219, 1286, 1288, 1312, 1319, 1320, 1348, 1349,

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

INDX

1352, 1354, 1357, 1360, 1361, 1365, 1366, 1367 Jayawijaya Range, 4, 201–202, 487, 488, 599, 600, 601, 602, 603, 630, 660, 1132. See also Central Cordillera (Merauke Range); Sudirman Range Jayawijaya region/Regency, 325, 339, 488, 1109, 1113, 1114, 1130, 1142, 1150, 1168 Jayawijaya Wildlife Sanctuary, 1242, 1268, 1269, 1271, 1426, 1433 Jurassic, 220, 246, 247 Kai Islands, 537, 546, 547, 579, 580, 592, 628, 811 Kaimana, 112, 181, 213, 880, 1068, 1071, 1078, 1151 Kaimana Bay, 505, 637 Kakadu National Park, 1335, 1336, 1339, 1341 Kalimantan, 277, 289, 645, 760, 794, 844, 933, 934, 946, 947, 948, 962, 967, 977, 1150, 1200, 1205, 1208, 1209, 1276, 1285, 1318, 1322 Kamoro, 118, 1105, 1115, 1140 Kangaroos and wallabies (Macropodidae) fossil, 123, 125, 126, 128, 130, 248–249, 250, 251, 252, 263, 1090, 1092 kangaroos (Dendrolagus), 3, 10, 200, 201, 387, 671, 694, 695, 712, 717–718, 726, 728, 730, 731, 732, 735, 736, 738, 744, 745, 1285, 1286, 1404 wallabies (Dorcopsis, Dorcopsulus, Macropus, Thylogales), 665, 695, 708, 717– 718, 726, 729, 731, 734, 736, 841, 1059, 1060, 1092, 1161, 1333, 1404, 1422 Kania, 429, 431, 959, 1008 Kanum, 1331, 1332, 1339 Kapauku, 1131, 1134, 1136 Kapok, 957, 1103 Kapuas River, 483, 645 Kawe´ Island, 75, 257, 288, 808 Kelangur, 249, 251 Kelela Swamp, 261, 1093 Kemabu Plateau, 260, 1017, 1029, 1030, 1038, 1039, 1043 Keranga. See Heath Keystone resources/species, 668, 677, 842, 1202 Kikori River (PNG), 216, 217, 218, 489, 827, 846, 862, 877, 1213 Kimaam Island. See Dolok Island Kimam (people), 109, 1138, 1139, 1140 Klamono, 1154, 1212, 1428, 1433

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index / 1453 Kofiau Island. See Raja Ampat Islands Kokas (Bomberai Peninsula), 270, 1068, 1074, 1076, 1077 Kolepom Island. See Dolok Island Kopermas. See Community cooperatives (kopermas) Kosipe, 121, 1088, 1089 Kotaraja, 1349, 1351, 1352, 1355, 1357, 1360 Kria Cave, 123, 125, 128, 130 Kuk Swamp, 121, 1091, 1126 Kumawa Mts (Bomberai Peninsula), 5, 209, 210, 213, 214, 231, 487, 599, 601, 603, 659, 690, 732, 1246, 1268, 1270, 1427, 1431 Kumbe River (Trans-Fly), 1054, 1055, 1270, 1327, 1427 Kuru/BSE/nvCJD, 89, 99 Kwiyawagi, 172, 249, 250, 251, 736, 1090, 1092 Labor. See Employment/labor Lachitu Rockshelter, 121, 1092 Lae, 43, 71, 79, 143, 468, 544, 545, 555, 980 University of Technology (UniTech), 87 See also Morobe Lae Herbarium, 85, 86, 89, 93, 94, 98, 269, 270 Lake Habbema, 52, 258, 269, 387, 534, 719, 736, 1021, 1027, 1036, 1091 Lake Hogayaku, 258, 259 Lake Hordorli, 261, 262, 264, 1090, 1092, 1094 Lakekamu River, 217, 589, 890 Lake Kutubu, 62, 211, 217, 1246 Lake Larson, 1040, 1043, 1044, 1045 Lake Sentani, 6, 186, 214, 261, 263, 645, 651, 873, 875, 888, 892, 960, 1089, 1092, 1246, 1314. See also Sentani Lake Wanagon, 1211 Lake Yamur (Bird’s Neck), 6, 645, 858, 875, 888 Lamington (cake), 29 Land conversion, 490, 521, 522, 676, 884, 885, 1003, 1144, 1156, 1158, 1160, 1170, 1177, 1178, 1200, 1202, 1207–1209, 1215, 1236, 1243, 1251, 1312, 1352. See also Habitat loss; Plantations, land conversion to Land tenure system, 110–111, 425, 1134, 1137, 1140, 1144, 1234, 1277, 1344 Languages, 11, 108–109, 1094, 1149, 1205, 1331, 1332, 1336, 1345 Lapita, 16, 467, 722. See also Austronesians Laurasian biota, 286, 320, 404, 689

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

Law enforcement. See Enforcement Leatherback Turtle, 574, 591 Legislation, Government Regulations (PP) 28/1985 (forestry), 1279 13/1994 (Wildlife Hunting), 1279 68/1998 (nature conservation areas), 1279 6/1999 (Production Forest utilization), 1279, 1281 7/1999 (Plant and Animal Preservation), 1280, 1300 8/1999 (Wildlife Utilization), 1280, 1300 25/2000 (Conservation Forests), 1282 34/2002 (Forest Management), 1280, 1282 45/2004 (Forest Protection), 1279 PERPU (illegal logging), 1292 Legislation, Laws (UU) 5/1967 (Basic Forestry), 1279 6/1967 (Basic Forestry), 1283 11/1967 (mining regulation), 1152 8/1981 (criminal litigation), 1284 4/1982 (environmental management), 1278 5/1990 (Conservation Act), 1253, 1258, 1278, 1279, 1280, 1283, 1284, 1300, 1302 5/1991 (protected species), 1283 5/1992 (protected species), 1283 16/1992 (quarantine), 1278 5/1993 (protected species), 1283 23/1997 (environmental management), 1278 20/1999 (BKSDA), 1255, 1256, 1258, 1316 21/1999, 1316 22/1999 (Regional Autonomy), 1168, 1219, 1281, 1282 25/1999 (decentralization), 1168 41/1999 (Forestry Act), 1253, 1258, 1279, 1282, 1283, 1284, 1302 45/1999 (division of Papua), 1119, 1219, 1220 20/2000 (decentralization), 1315 21/2000 (conservation), 1258, 1314 21/2001 (Special Autonomy), 1169, 1219, 1258, 1282, 1316, 1320 26/2002, 2002 Legislation, Ministry of Forestry Decrees (SK), 1281, 1282 Lemdubu Cave, 123 Lengguru Fold Belt. See Fold Belt Lepidoptera, 375, 377, 519, 592, 620, 863 butterflies, 10, 201, 377, 391, 453, 480, 485, 486, 516, 519, 520, 522, 523, 1230, 1241, 1286, 1287, 1313 butterfly farming, 522

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1454 / index Lepidoptera (cont.) collections, 30, 31, 35, 38, 40, 43, 46, 47, 63, 68, 70, 71, 83, 94, 96, 97, 98, 99, 100–101 moths, 10, 391, 396, 519, 520, 521, 522, 1365 Lesser Sunda Islands. See Nusa Tenggara Libocedrus, 9, 1027 Lichens (lichenized fungi), 86, 100, 276, 291, 292, 303–317, 953, 997, 1012, 1014, 1018, 1041, 1042, 1043, 1045, 1046 collections, 38, 39, 40, 58, 86, 91, 100 See also Fungi LIPI. See Indonesian Institute of Sciences (LIPI) Lipinia, 489, 577, 578, 600, 602, 1413 Liquid natural gas (LNG). See Oil and gas extraction; Tangguh LNG project Lithocarpus, 9, 36, 470, 953, 978–988, 990– 993, 996, 998, 1003 Litsea, 911, 922, 929, 933, 937, 941, 951, 954, 955, 967, 981, 982, 992, 1021 Liverworts. See Hepaticae Lizards, 569, 1366 biology/ecology, 592–594 endemism, 489, 600, 601, 602 introduced, 586, 1367–1368 as prey, 589, 596, 597, 598, 599, 1356, 1359 taxa/distribution, 480, 482, 483, 485, 486, 564–567, 575–580, 1412–1414 trade, 604 See also Emoia; Gecko; Skink; Varanids LMAs. See Traditional community associations (LMAs) Lobster, 508, 511, 591, 793, 812, 831, 839, 845, 846, 847, 1164, 1216, 1237. See also Crustaceans Logging clearcutting, 317, 644, 650, 759, 884, 956, 1202, 1209 illegal, 629, 765, 794, 883, 1159–1160, 1207, 1220, 1276, 1282–1285, 1309 large-scale/industrial, 677–678, 883, 1158– 1159, 1160 in protected areas, 644, 674, 1206, 1259, 1283 selective, 314, 317, 397, 677, 850, 883, 885, 954, 1192, 1351, 1352 small-scale, 677, 883, 1160–1161, 1207, 1280, 1281, 1282 See also Community cooperatives (kopermas); Forests, protected; Habitat loss; Non-timber forest products; other logging entries

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

INDX

Logging, ecological aspects, 677–678, 764 as biodiversity threat, 317, 397, 629, 634, 650, 657, 671, 677–678, 764, 1003, 1202, 1205–1207, 1214, 1220, 1236– 1237 and climate change, 764, 1206 and environmental degradation, 510, 644, 764, 793, 806, 810, 814, 850, 872, 878, 883, 885, 965, 1202, 1206, 1207, 1216, 1276, 1352 minimizing impact of, 677–678 succession/regeneration, 397, 668, 677– 678, 1003, 1351 See also Habitat loss Logging, quantitative measures (area, monetary value), 1115, 1158–1160, 1161, 1205–1207, 1276 Logging, societal aspects benefits research, 645 clearing for plantations, 884, 1208–1209 concessions/licenses, 850, 1158, 1175– 1177, 1178, 1208–1209, 1234, 1278, 1280, 1281, 1316 economic aspects, 651, 794, 850, 1115, 1158, 1167, 1170, 1179–1180, 1183, 1191, 1206, 1276 planning/conservation, 1232, 1234, 1242, 1243 policy/legal aspects, 1115, 1117, 1220, 1277, 1280–1285, 1288–1302, 1303, 1304, 1307, 1316 profit from, 765, 883, 1115, 1158, 1285, 1309, 1311 See also Community cooperatives (kopermas); Enforcement; Legislation, Government Regulations (PP); Legislation, Ministry of Forestry Decrees (SK) Long Island, eruption of, 466, 946 Lorentz National Park, 257, 487, 488, 521, 599, 600, 601, 602, 603, 993, 1199, 1221, 1262, 1268, 1269, 1271, 1433 Lorentz River, 52, 217, 733, 734, 736 Lories, 661, 668, 669, 678, 1219, 1285, 1286, 1287–1288, 1388, 1404, 1405 Louisiade Islands (Misima, Rossel, Tagula), 207, 208, 209, 211, 219, 245, 386 Lungfish (Neoceratodus), 646 Lycophytes, 335–341 Macaques, 679, 723, 1218–1219, 1348–1361, 1365 Madagascar, 335, 346, 371, 382, 386, 399, 401, 473, 582, 691, 891, 1077

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index / 1455 Madang (PNG), 303, 305, 314, 315, 317, 325, 373, 417, 468, 504, 543–547, 833 Makassar (Sulawesi), 800, 821, 1287 Makassarese, 1115, 1286 Malaria, 40, 84, 96, 522, 645, 667, 838, 892, 1109, 1110, 1357 avian, 678 Malay Archipelago, 160, 396, 473, 655, 658, 689, 723 Malaysia, 127, 277, 280, 312, 316, 324, 325, 335, 383, 394, 464, 466, 495, 542, 546, 548, 773, 777, 789, 965, 980, 983, 1104, 1160, 1239, 1277, 1285, 1350, 1354, 1360 Maluku, 11, 17, 18, 20, 46, 95–96, 113, 114, 123, 157, 167, 187, 213, 248, 336, 339, 356, 381, 382, 385, 386, 411, 420, 423, 457, 479, 529, 534, 564, 571, 573, 579, 580, 582, 584, 585, 599, 619, 628, 629, 640, 656, 659, 689, 690, 720, 723, 733, 748, 808, 811, 813, 948, 977, 1077, 1101, 1208, 1239, 1246, 1265, 1285, 1318, 1322 Ambon, 14, 17, 24, 213, 571, 582, 1101 Bacan, 157, 533 Buru, 213, 381 (see also Taman Buru) Halmahera, 114, 146, 147, 148, 157, 158, 159, 167, 181, 383, 496, 582, 640, 773, 808 Obi, 157, 496 Seram, 198, 213, 336, 339, 385, 521, 552, 571, 582, 586 Ternate, 23, 46, 167 See also Aru Islands; Gebe Maluku (people of/Moluccans), 11, 1110, 1112, 1120 Mamberamo dam project. See Dams, Mamberamo dam project Mamberamo-Foja Wildlife Sanctuary, 1242, 1244, 1268, 1269, 1271, 1428, 1432 Mamberamo River/Basin, 3, 5, 6, 8, 150, 155, 157, 169, 173, 180, 198, 209, 210, 214, 215, 220, 222, 234, 235, 261, 339, 487, 488, 489, 515, 569, 584, 589, 599, 600, 603, 645, 648, 651, 660, 661, 724, 726, 729, 778, 827, 844, 848, 861, 864, 874, 878, 886, 888, 889, 904, 910, 911, 920, 949, 950, 1136, 1137, 1180, 1182, 1199, 1217, 1246, 1311, 1312 Mangroves, 6, 9, 87, 166, 264, 506–508, 664, 756, 757, 763, 773, 801, 802, 806, 808, 810, 814, 816, 824–850, 879–881, 882, 905, 910, 933, 946, 1174, 1175, 1207, 1208, 1213, 1216, 1223, 1244, 1246, 1270 animals in, 495, 505–508, 539, 541, 542, 544, 546, 548, 549, 552, 555–557, 596,

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

597, 619, 631, 638, 837–847, 1404– 1409 ecosystem services/production, 760–761, 833–837, 847–848, 1204 human uses/impact, 849–850 plants in, 337, 355, 363, 367, 371, 385, 394, 416, 417, 440, 457, 468 Manihot. See Cassava Manioc, 1135, 1137 Manokwari, 220, 260, 325, 337, 366, 467, 520, 541, 586, 630, 645, 730, 809, 259, 1074, 1102, 1107, 1108, 1110, 1111, 1112, 1117, 1142, 1151, 1156, 1157, 1158, 1161, 1209, 1213, 1219, 1261, 1286, 1287, 1319, 1320, 1352, 1365, 1366, 1367. See also State University of Papua (UNIPA) Manokwari Herbarium, 270, 276 Mansuar. See Raja Ampat Islands Manus Island. See Admiralty Islands Mapala Rockshelter, 123, 251, 1092 Mappi, 110 Mariana Islands/Arc, 141, 149, 313, 580, 596 Marind, 1131, 1332, 1339 Marind-Anim, 48, 109, 111, 113 Marine National Park (TNL: Taman Nasional Laut), 1262, 1271, 1430 Marine protected areas, 501, 816, 1269, 1272 Marine Wildlife Sanctuary (Suaka Margasatwa Laut), 1271, 1429–1430 Markham River/Basin, 209, 210, 215, 234– 235, 270, 383, 489, 555, 648, 827, 844 Marori-Mengey, 1331, 1332, 1339 Marshall Islands, 580, 642, 783 Marsupials, 29, 96, 248, 249, 252, 402, 466, 480, 483, 689, 691, 692–698, 711–718, 724, 726, 729, 734, 743–746, 1219 fossil, 123, 125, 126, 128, 130, 248–249, 250, 251, 252, 263, 1090, 1092 See also Bandicoots; Cuscuses; Dasyurids; Kangaroos and wallabies; Possums and gliders Matoa, 9, 458, 1158. See also Pometia Mauritius, 313, 382, 723, 1348–1351, 1353, 1356 Maya, 108, 112, 114 Mayalibit Bay (Waigeo Island), 638, 773, 808, 958, 959 Mayr, Ernst, 37, 61, 62, 64, 66, 200, 515, 656 McCluer Gulf. See Berau Bay Me, 109, 110, 111, 113, 114, 116, 117, 118, 119 Medical entomology, 70, 83, 84, 106, 516 Medical services. See Health/health services

INDX

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1456 / index Medicinal plants. See Traditional medicine Medicine. See Health/health services; Traditional medicine Meervlakte, 210, 234, 663, 874, 878. See also Mamberamo Megapodiidae, 7, 199, 655, 661, 672, 674, 675, 676, 841, 1386, 1404 Melaleuca, 276, 429, 431, 432, 470, 917, 919, 1055–1058, 1060–1061, 1327, 1329, 1334 Melampitta, 663, 664, 674, 757, 1399 Melanotaeniidae. See Rainbowfish Meliphagidae. See Honeyeaters Mentawai Islands, 138, 760 Merauke Range. See Central Cordillera (Merauke Range) Merauke region/Regency, 167, 169, 170, 171, 175, 182, 183, 184, 185, 186, 188, 189, 193, 541–543, 555, 604, 628, 630, 733, 734, 809, 810, 874, 887, 889, 892, 902, 903, 906, 942, 1059, 1109, 1110, 1111, 1112, 1114, 1136, 1138, 1142, 1151, 1156, 1157, 1158, 1160, 1161, 1163, 1178, 1209, 1220, 1262, 1270, 1320, 1330, 1333, 1339, 1341, 1345, 1352, 1417–1421, 1427 Merbau, 9, 1115, 1158, 1207. See also Intsia Meren Glacier/Valley, 190–192, 1033, 1036, 1044–1051. See also Glaciers Mesozoic, 149, 150, 151, 153, 220, 221, 246, 247, 344, 481, 566 Metroxylon sagu. See Sago Meybrat, 108, 109, 110, 111, 113, 116, 118 Microperoryctes. See Bandicoots Military, 883, 885, 1101, 1102, 1105, 1115, 1118, 1119, 1121, 1189, 1286, 1287, 1290, 1307. See also Armed Forces; Navy Milne Bay region/Province (PNG), 218, 485, 503, 546–548, 582, 644, 660, 663, 711, 735, 790, 793, 794, 891, 992 Mimika (people), 109, 110, 114 Mimika district, 630, 809, 810, 1119, 1150, 1158, 1168 Mimika River, 489, 568, 733, 734 Minajerwi River, 811, 861, 865, 905, 906, 911, 935, 940, 942, 957, 995, 1211 Minerals. See Mining Mining, 30, 158, 645, 869 as biodiversity threat, 329, 650–651, 889, 1078, 1081, 1209–1212, 1221, 1255 economic value/policies, 651, 1105, 1114, 1115, 1149–1154, 1167–1170, 1172,

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

INDX

1177, 1180, 1189, 1192, 1215, 1245, 1255 land conversion, 490, 657, 764, 814, 1037, 1046, 1141, 1167 tailings/pollution/sedimentation from, 264, 329, 523, 666, 806, 810, 811, 814, 817, 849, 884–885, 1037, 1141, 1209– 1212, 1221 See also Copper; Freeport McMoran; Gold; Nickel; Sand mining Ministry for the Environment, 1311 Ministry of Forestry, 1158, 1176, 1257, 1277, 1279, 1280, 1281, 1282, 1289, 1291, 1292, 1298, 1302, 1313, 1316, 1330, 1333 Ministry of Industry and Trade, 1282, 1288, 1290, 1292 Ministry of Interior, 1254 Ministry of Justice, 1288, 1290, 1291 Ministry of Mining and Energy, 1151 Ministry of Transportation, 1288, 1290, 1292 Miocene. See Tertiary, Miocene Misima Island. See Louisiade Islands Misool Island. See Raja Ampat Islands Misool Terrane, 149, 151, 288 Missionaries. See Christian missionaries Mites, 70, 71, 83, 414, 461, 497, 517, 522 Miyanmin, 93 Mobile Belt, 150, 153, 154–158, 197, 200 Moi, 113, 114, 1142 Mollusks, 495–511, 641, 781, 782, 783, 812, 838, 839, 840 early collectors, 21, 38, 99 fossils/middens, 123, 246, 247, 814 introduced, 893, 1218 as prey, 620, 845, 846 terrestrial/freshwater, 517, 519, 523, 620, 859, 869, 872 Moluccan ironwood. See Intsia Moluccan Islands. See Maluku Monitor lizards. See Varanids Monkeys. See Macaques Monotremes, 96, 480, 483, 689, 691, 692, 710, 724, 726, 730, 734, 743–744. See also Echidnas Monsoon, 7, 177, 178, 181, 188, 189, 263, 457, 496, 506, 802, 808, 996, 1054, 1091, 1327 Monsoon forest, 9, 350, 384, 617, 624, 661, 710, 711, 713, 717, 733, 754, 917, 946, 979, 1054, 1055, 1327, 1329, 1330, 1344 Moray eels (Muraenidae), 639, 1406 Morehead River (Trans-Fly), 892, 1054, 1327, 1328, 1331, 1417–1421

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index / 1457 Morobe, 209, 210, 216–217, 238, 325, 373, 417, 421. See also Lae Mosquitoes, 506, 520, 522, 838 collections, 62, 64, 70, 71, 84, 96, 99 control, 96, 873, 889, 891, 892 See also Malaria Mosses/moss forest, 9, 28, 308, 320–331, 441, 470, 534, 589, 720, 859, 860, 869, 978, 979, 980, 981, 982, 990, 991, 992, 996, 997, 998, 1004, 1006, 1007, 1008, 1012, 1014, 1021, 1029, 1030, 1039, 1041, 1043, 1045, 1046, 1047, 1048, 1049. See also Bryophytes; Cloud forest Moss mouse, 689, 732, 735 Moths. See Lepidoptera Motupore Island, 87, 540, 546, 835, 837 Mt Arfak Strict Nature Reserve, 1262. See also Arfak Mts Mt Carstensz. See Mt Jaya Mt Gahavisuka. See Gahavisuka Provincial Park Mt Giluwe (PNG), 216, 260, 1003, 1006, 1013, 1039, 1044 Mt Idenburg, 5, 1007, 1021 Mt Jaya, 5, 12, 17, 51, 53, 123, 169, 177, 183, 184, 185, 186, 187, 188, 190, 191, 222, 246, 250, 251, 258, 260, 269, 336, 337, 568, 589, 736, 962, 979, 1004, 1005, 1025, 1026, 1028, 1030, 1033, 1037, 1040, 1043, 1044, 1045, 1046, 1047, 1049, 1051, 1052, 1092, 1094, 1223. See also Carstensz Glacier; Central Cordillera (Merauke Range); Sudirman Range Mt Kinabalu, 307, 312, 350 Mt Lorentz, 339, 1311 Mt Mandala, 5, 15, 186, 190, 257 Mt Meja, 959, 1271, 1427, 1433 Mt Sumuri (Weyland Range), 714, 728, 737, 738 Mt Trikora, 5, 49, 51, 168, 190, 257, 258, 269, 713, 719, 736, 881, 1025, 1026, 1027, 1029, 1036, 1038, 1066, 1091. See also Central Cordillera (Merauke Range); Sudirman Range Mt Wilhelm (PNG), 88, 216, 304, 305, 308, 311, 314, 315, 325, 330, 980, 1025, 1026, 1040, 1043, 1045, 1046, 1047, 1048 Mt Wilhelmina. See Mt Trikora Mt Wondiwoi, 221, 603, 690, 728, 745, 1268, 1270, 1427. See also Wandammen Mts Murids, 200, 201, 202, 689, 699–703, 708, 718–719, 726, 729–732, 734–737, 746– 748, 1404, 1422 moss mouse, 689, 732, 735

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

rats, 10, 201, 406, 671, 672, 676, 678, 689, 702, 703, 718–719, 722, 723, 726, 730, 731, 732, 734, 736, 747, 748, 841, 1218, 1423 Museum Zoologense Bogoriense, 486, 569, 648, 735, 738 Mutualism. See Symbiosis Muyu, 108, 109, 110, 111, 114, 116, 117, 118, 1106 Myristica, 290, 292, 296, 408–413, 415, 668, 833, 879, 904, 911, 912, 914, 916, 923, 924–926, 928, 929, 931, 933, 935, 936, 937, 939, 940, 947, 948, 949, 951, 953, 954, 955, 973, 981, 982, 984, 988, 996, 1142. See also Ants, and plants; Nutmeg Myrmecodia. See Rubiaceae Myrmecophily. See Ants, and plants Nabire, 156, 257, 261, 366, 521, 569, 630, 809, 1151, 1152, 1209, 1210, 1220, 1221, 1246, 1427, 1433 National Intelligence Board (BIN), 1119 National Parks (Taman Nasional), 1179, 1253, 1254, 1255, 1257, 1259, 1260, 1262, 1271, 1272, 1273, 1283, 1317, 1318, 1338, 1340 legislation, 1261 outside of region, 1251, 1312 size, 763, 1256, 1257, 1265, 1318, 1433 See also Protected Areas National Parks management, 92, 1318, 1340 National Park Bureau (BTN), 1255, 1256, 1261, 1262, 1317, 1337, 1339, 1340 See also Natural Resources Conservation Bureau (BKSDA) National Police, 1288, 1289, 1291, 1292. See also Police Natural Resources Conservation Bureau (BKSDA), 1255–1257, 1261–1262, 1287, 1288, 1291, 1302, 1303, 1317, 1318, 1319, 1320, 1322, 1324, 1437 Nature Conservation Area (Kawasan Konservasi Alam). See Protected Areas Nauclea, 264, 911, 920, 929, 931, 933, 935, 941, 943, 952, 955, 963, 964, 995, 1054, 1055, 1056, 1057, 1058, 1089 Navy, 1101, 1290, 1293, 1295, 1298. See also Military Nduga, 1143 New Britain, 127, 145, 147, 148, 167, 198, 221, 305, 346, 384, 385, 386, 399, 404, 417, 421, 543, 662, 1076, 1365 New Britain Arc, 144, 158

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1458 / index New Caledonia, 145, 149, 151, 152, 248, 306, 381, 382, 384, 386, 387, 424, 498 New Guinea Harpy Eagle, 632, 665, 675, 1237, 1264, 1328, 1394, 1405 New Ireland, 122, 385, 399, 548, 573, 662, 1365 Nickel, 142, 168, 793, 958, 959, 1152 Nimboran, 110, 112, 117, 1106 Nombe Rockshelter, 121, 249, 1088, 1091 Non-timber forest products (NTFP), 110, 1125, 1132, 1136, 1161, 1214, 1259, 1332, 1333, 1337 license to collect (IHHBK), 1161 See also Coconut, copra; Copal; Firewood; Kapok; Pandanus; Rattan; Resins; Rubber; Sago; Traditional medicine; Ylang-ylang Northwall Firn (glacier), 189, 190–193, 1048. See also Glaciers Nothofagus, 9, 60, 66, 248, 252, 260, 261, 264, 306, 327, 346, 441, 445, 470, 946, 947, 959, 979, 980, 981, 982, 990, 992, 996, 998, 1000, 1003–1009, 1010, 1011, 1012, 1017, 1090 Numfoor Island (Cenderawasih Bay), 7, 114, 168, 197, 200–201, 204, 212, 487, 489, 537, 599, 601, 630, 692–696, 699–702, 704–705, 720, 730, 733, 749, 809, 1077, 1120, 1246, 1269 Nurseryfish (Kurtus), 650, 1407, 1421 Nusa Tenggara (Lesser Sunda Islands), 138, 386, 468, 571, 579, 580, 640, 720, 723, 873, 948, 977, 1110, 1239, 1246, 1265, 1318, 1322, 1350 Nutmeg (Myristica fragrans), 411, 413, 1103, 1142, 1156, 1157. See also Myristica Nypa, 9, 359, 363, 366–367, 597, 827, 828, 831–833, 836, 845, 880, 1351, 1424 Oaks. See Castanopsis; Lithocarpus Obi Island. See Maluku Ocean currents. See Currents Octomeles, 9, 923, 929, 933, 935, 939, 940, 949, 951, 952, 953, 954 Odonata, 211, 215, 219, 220, 228–245, 480, 518, 859, 863, 865, 868, 869, 877, 878, 879, 884 damselflies (Zygoptera), 207, 211, 213, 214, 215, 216, 218, 219, 480, 518, 863, 868, 871, 892, 1312 dragonflies (Anisoptera), 211, 480, 518, 863, 873, 949, 952 Oil and gas extraction, 850, 1103, 1154–1155, 1172, 1212–1214, 1234, 1242, 1243, 1245

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

INDX

economics, 1103, 1105, 1111, 1114, 1118, 1149, 1150, 1154–1155, 1169, 1170 environmental effects, 813, 816, 849, 1212– 1214 See also British Petroleum; Pertamina; Tangguh LNG project Oil palm as biodiversity threat, 676, 884, 1208 and fertilizers/pesticides, 667, 884, 1208 land clearing/conversion, 676, 884, 1145, 1177, 1178, 1202, 1207, 1213 plantations, 814, 1142, 1156–1157, 1178, 1209 Oligocene. See Tertiary, Oligocene Olive Ridley Turtle, 574, 591 Onin Peninsula, 5, 47, 64, 112, 117, 197, 792 Ontong Java Plateau, 143, 145 Ophiolite Belt, 142, 153, 159, 222 Ophiolites, 141, 142, 144, 145, 149, 152, 153, 154, 156, 158, 159, 212, 215, 218, 220, 222, 223, 566. See also Geology; Island arcs Orchids, 46, 66, 91, 291, 296, 435–455, 967, 981, 1012 Bulbophyllum, 296, 436, 437, 443, 444, 445, 446, 447, 448, 451, 452, 453 Dendrobium, 29, 64, 296, 436, 437, 441, 443, 444, 446, 447, 449, 450, 451, 452, 453, 455, 919, 949, 967, 1028, 1029, 1030, 1036, 1038, 1286 Glomera, 296, 436, 439, 441, 442, 443, 447, 1029, 1038 Phreatia, 296, 436, 437, 1028, 1036 trade, 453–455, 1219 Oriomo Plateau, 470, 1054, 1055, 1327 Oriomo River, 1329, 1416–1421 Ormu, 1138, 1139, 1142 Orthoptera (grasshoppers, katydids, crickets), 518, 624, 1072, 1074, 1076, 1356 Otomona River, 264, 952, 954, 955, 956, 958, 985, 1212 Owen Stanley Mts, 209, 210, 216, 218, 219, 242–243, 489, 573, 992 Owlet-nightjar (Aegothelidae), 658, 662, 674, 1389 Pacific Plate, 4, 137, 143, 148, 150, 156, 157, 201, 647, 774, 1151, 1199 Padaido Islands (Cenderawasih Bay), 7, 499, 692–702, 720, 773, 809, 816, 1433 Palau, Republic of (Angaur), 34, 141, 573, 579, 582, 723, 783, 1348, 1350, 1351, 1360

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index / 1459 Paleozoic, 149, 159, 213, 216, 221, 246, 344, 481, 566 Palm oil. See Oil palm Pamwak Rockshelter, 122 Pandanus, 29, 331, 588, 832, 879, 901, 903, 904, 911, 912, 916, 917, 918, 919–920, 921, 922, 923, 926, 928, 929, 931, 933, 934, 935, 941, 943, 948, 949, 951, 953, 954, 957, 967, 968, 969–972, 974, 975, 982, 985, 989, 990, 998, 1003, 1008, 1009, 1010, 1011, 1012, 1029, 1054, 1056, 1057, 1058, 1089, 1356 cultivated, 1130, 1132, 1134, 1136, 1144 Paniai Lakes, 6, 11, 60, 187, 211, 216, 222, 260, 269, 537, 630, 645, 711, 728, 736, 737, 738, 746, 858, 1012, 1087, 1102, 1109, 1112, 1114, 1119, 1129, 1215 Papuacedrus, 315, 345, 347, 978, 997, 998, 1003, 1008, 1009, 1012, 1018, 1019, 1020, 1021, 1027, 1028, 1029, 1030, 1031, 1036. See also Cupressaceae Paradisaeidae. See Birds of Paradise Parrotfishes (Scaridae), 638, 639, 642, 787, 807 Parrots (Psittacidae), 10, 201, 369, 406, 460, 465, 655, 661, 668, 671, 672, 677, 1353, 1359, 1388, 1405 trade, 678, 1161, 1219, 1285, 1286, 1312– 1313 Peanuts (ground nuts), 1134, 1140, 1144, 1156, 1360 Peat, 165, 167, 169, 249, 260, 261, 309, 441, 917, 978, 1014, 1017, 1038, 1043, 1044, 1087, 1089, 1090 swamp, 759, 879, 892, 910, 917, 920, 923, 933–935, 940, 942, 943, 994, 1005, 1012, 1017, 1033, 1036, 1038, 1040, 1043, 1044, 1045, 1046, 1057 See also Swamps/swamp forest Peoples. See Human groups Peroryctes. See Bandicoots Pertamina, 111, 1117, 1119, 1154, 1155, 1212 Peru, 73, 1312. See also Andes Pesticides, 523, 667, 1208, 1216 Pests. See Agriculture and forestry pests; Diseases, human; Diseases, nonhuman Petauris. See Possums and gliders Petroleum. See Oil and gas extraction Philippine Islands, 137, 141, 146, 160, 277, 280, 289, 306, 312, 313, 316, 324, 325, 326, 357, 359, 384, 400, 420, 424, 443, 466, 495, 498, 501, 503, 541, 542, 546– 548, 573, 575, 577, 584, 585, 586, 594, 639, 720, 773, 777, 782, 785, 790, 793, 794, 800, 817, 1150, 1160, 1350, 1360

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

Philippine Sea Plate, 137, 145, 774 Phragmites, 934, 942, 943, 950, 951, 980, 1013, 1056, 1057, 1058, 1060, 1334 Phreatia. See Orchids Phyllocladus, 9, 248, 346, 347, 978, 992, 996, 997, 998, 1003, 1008, 1018, 1021, 1028, 1036 Pigs, 11, 114, 117, 406, 460, 722, 841, 886, 1061, 1128, 1129, 1134, 1156 Pioneer species, 367, 397, 803, 804, 806, 832, 901, 902, 904, 905, 943, 950, 956, 957, 958, 985, 990, 995, 1014, 1018, 1041, 1042, 1045, 1129. See also Disturbed habitat; Regeneration; Secondary vegetation; Seral vegetation Pipefishes (Syngnathidae), 235, 642, 650, 813, 1406 Piper, 958, 998, 1008, 1218 Pitohuis, 406, 661, 662, 673, 1399 Plantations, 33, 35, 36, 40, 68, 94, 347, 1137, 1141, 1142, 1173, 1206, 1242, 1263 ecological effects, 814, 884, 905, 1178, 1207–1209 economic aspects/concessions, 1103, 1141, 1142, 1145, 1150, 1151, 1156–1157, 1178, 1208–1209, 1215 as habitat, 311, 589, 599, 670 land conversion to, 521, 604, 676, 884, 1145, 1177, 1202, 1207–1209 See also Cocoa; Coconut; Coffee; Land conversion; Oil palm Pleistocene. See Quaternary, Pleistocene Pliocene. See Tertiary, Pliocene Plume trade. See Birds of Paradise, trade; Wildlife trade, birds Podocarpaceae, 8, 9, 346, 347, 947, 980, 981, 982, 996, 998, 1029, 1037, 1042. See also Dacrycarpus; Dacrydium; Phyllocladus; Podocarpus Podocarpus, 9, 315, 959, 969, 971, 972, 981, 997, 998, 999, 1000, 1002, 1008, 1009, 1012, 1018, 1019, 1028, 1030, 1032, 1033, 1034, 1035 Police, 643, 1111, 1115, 1284, 1287, 1288, 1289, 1291, 1292, 1293, 1294, 1295, 1296, 1297, 1298, 1299, 1301, 1302, 1303, 1304, 1362. See also Enforcement Pollination, 381, 465 by bats, 391–392, 432 by bees, 367, 381, 401–402, 425, 459–460, 473 by birds, 375, 391–392, 432, 446–447, 465, 473, 668–669 by insects, 367–368, 375, 391–392, 396, 413, 425, 432, 446–452, 465

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1460 / index Pollination (cont.) by wind, 405, 459, 465, 1004, 1006 See also Seed dispersal Pollution, 1184, 1185, 1203 air, 329 chemical, 523, 902 effects of, 523, 814, 1081, 1207 enforcement, 815 marine, 792, 793, 1216 water, 902 See also Pollution and sedimentation, causes of; Sedimentation Pollution and sedimentation, causes of agriculture, 329, 814, 884 deforestation, 523, 510, 814, 1206 human population/development, 651, 778, 792, 814, 1185, 1214 industry, 814, 1208 mining, 264, 329, 523, 666, 806, 810, 811, 814, 817, 849, 884–885, 1037, 1141, 1209–1212, 1221 pesticides, 523, 1208 Pometia (inc. matoa), 9, 289, 457, 458, 460, 467, 923, 932, 935, 937–939, 940, 947, 948, 949, 950, 952, 953, 954, 955, 959, 983, 984, 990, 1139, 1158, 1356, 1359 Port Administration, 1288, 1290, 1292 Port Moresby, 270, 303, 305, 317, 540, 543– 548, 549, 552, 553, 554, 560, 735, 827, 831, 834, 891 Possums and gliders (Acrobatidae, Burramyidae, Petauridae, Pseudocheiridae), 10, 24, 250, 252, 263, 387, 508, 665, 689, 696, 697, 710, 715–717, 726, 727, 730, 731, 732, 734, 735, 746, 841, 1092, 1359, 1404, 1422 See also Cuscuses Potato (Solanum), 1092, 1139, 1142, 1156 introduction of, 54 Prawns. See Shrimp Prince Alexander Mts, 154, 155, 209, 210, 214, 215, 221, 223, 489, 660, 708, 729 Priority setting. See Conservation priority setting Production Forest (Hutan Produksi). See Forests, protected Protected areas Biosphere Reserve (Cagar Biospher), 1253, 1254 Game Reserve (Taman Buru), 1254, 1257, 1259, 1283 Grand Forest Park (Taman Hutan Raya), 1254, 1257, 1283 Marine National Park (TNL: Taman Nasional Laut), 1262, 1271, 1430

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INDX

Marine protected areas, 501, 816, 1269, 1272 Marine Wildlife Sanctuary (Suaka Margasatwa Laut), 1271, 1429–1430 National Park (Taman Nasional) (see National Parks) Natural Protected Area (Kawasan Suaka Alam), 1253, 1255, 1280, 1282, 1283 Nature Conservation Area (Kawasan Konservasi Alam), 1253–1254, 1255, 1257, 1262, 1279, 1280, 1282, 1283, 1302 Recreational Park (TW: Taman Wisata), 1254, 1257, 1259, 1262, 1265, 1271, 1272, 1273, 1283, 1430 Strict Nature Reserve (Cagar Alam), 1251, 1253, 1254, 1256, 1257, 1259, 1260, 1262, 1263, 1265, 1268, 1269, 1271, 1272, 1283, 1425 Wildlife Sanctuary (Suaka Margasatwa), 1253, 1254, 1256, 1257, 1259, 1260, 1261, 1262, 1265, 1271, 1272, 1283, 1321, 1430 See also Forests, protected; Legislation, Government Regulations (PP); Legislation, Ministry of Forestry Decrees (SK) Protected areas, hunting in, 1254, 1259, 1265, 1337, 1338, 1334 Protected areas, logging in, 644, 674, 1206, 1259, 1283 Protected forests (Hutan Lindung). See Forests, protected Pseudomugilidae. See Blue-eyes Psittacidae. See Parrots Psychotria, 292, 296, 952, 997, 1008, 1009, 1011 Pteridophytes. See Ferns Pterocarpus (inc. rosewood), 9, 903, 883, 953, 983 PT Freeport Indonesia (PTFI). See Freeport McMoRan PT Freeport McMoRan. See Freeport McMoRan Ptilonorhynchidae. See Bowerbirds Public health. See Health Puncak Jaya. See Mt Jaya Purari River/Delta, 202, 216, 217, 218, 489, 648, 827, 831, 832, 838, 841, 844, 845, 846, 847, 848, 1404–1409 Quarantine, 679, 1278, 1332 Quaternary (Cenozoic), 138, 165, 214, 220, 255, 264, 481, 698, 711, 723, 732, 736, 1004

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index / 1461 Pleistocene, 8, 121, 122, 123, 125, 128, 137, 174, 214, 248, 249, 250, 251, 252, 263, 264, 443, 496, 497, 519, 729, 733, 962, 981, 1003, 1025, 1038, 1089, 1090, 1094, 1095 Holocene, 122, 123, 125, 126, 128, 144, 190, 250, 251, 255, 260, 261, 264, 731, 1091, 1092, 1093, 1094 Quolls. See Dasyurids Radiation. See Solar radiation; Species radiations Rainbowfish (Melanotaeniidae), 210, 228– 236, 239–242, 646, 647, 648, 649, 650, 651, 863, 868, 871, 872, 873, 875, 887, 891, 892, 1417 Rainfall, 181, 185–189, 190, 778 distribution/locations, 7, 8, 154, 185–189, 827, 831, 874, 876, 877, 934, 966, 977, 983, 1054, 1327 seasonality, 6, 7, 187–189, 802 variability, 7, 188–189, 287, 946 See also Drought; Water balance Raja Ampat Islands, 3, 5, 10, 11, 22, 23, 24, 46, 47, 48, 58, 63, 72, 73, 75, 80, 83, 112, 113, 114, 117, 119, 197, 198, 199, 209, 212–213, 228–229, 282, 287, 366, 479, 489, 496, 498–502, 503, 504, 505, 574, 599, 629, 637, 638, 639, 643, 644, 645, 646, 648, 652, 656, 659, 707, 719, 720, 733, 772, 773, 781, 782, 789, 790, 792, 793, 794, 808, 809, 813, 816, 818, 831, 878, 881, 958, 959, 1200, 1212, 1216, 1217, 1246, 1261, 1271, 1305, 1312, 1323, 1428 Batanta, 7, 24, 46, 47, 62, 63, 72, 75, 80, 157, 197, 199, 209, 210, 211, 213, 228, 257, 487, 579, 599, 601, 624, 629, 638, 644, 646, 648, 659, 692–702, 713, 715, 717, 733, 749, 792, 808, 879, 1244, 1269, 1270 Kofiau, 7, 24, 46, 75, 199, 200, 204, 705, 733, 808, 1429 Mansuar, 881 Misool, 7, 22, 24, 46, 47, 54, 63, 72, 73, 75, 181, 186, 189, 197, 199, 209, 210, 211, 213, 229, 257, 287, 289, 488, 533, 534, 579, 600, 626, 629, 638, 646, 648, 661, 690, 692–702, 705, 714, 716, 717, 733, 749, 808, 881, 991, 1071, 1074, 1075, 1077, 1244, 1269, 1270, 1271, 1426, 1429, 1430, 1431 Salawati, 7, 22, 24, 46, 47, 62, 63, 72, 75, 80, 157, 181, 197, 199, 213, 257, 533,

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534, 619, 620, 621, 624, 627, 628, 638, 646, 648, 659, 661, 692–702, 711, 713, 714, 716, 717, 719, 720, 733, 808, 1154, 1206, 1212, 1270, 1426, 1431 Waigeo, 7, 18, 19, 22, 24, 46, 47, 49, 54, 58, 62, 63, 72, 75, 80, 146, 147, 148, 158, 159, 197, 199, 204, 209, 210, 211, 212, 228, 288, 340, 465, 487, 533, 534, 567, 575, 579, 599, 601, 620, 621, 622, 629, 638, 644, 646, 648, 659, 672, 690, 692– 702, 705, 713, 715, 717, 719, 720, 728, 730, 731, 733, 749, 773, 808, 890, 904, 945, 949, 953, 958, 959, 1217, 1244, 1269, 1270, 1426, 1429, 1431 (see also Mayalibit Bay) Ramsar Convention/Site, 815, 1330 Ramu River. See Sepik River Rapid Assessment Program (RAP), 75, 79, 569, 643, 789, 793, 808, 813, 818, 1241, 1311, 1312, 1323 format, 270 Marine, 75, 498, 502, 505, 643, 1216, 1312, 1323 needed, 1343 RACE, 1171–1172, 1173, 1215 See also Conservation International, surveys Rats. See Murids, rats Rattan, 361, 366, 367 Recreational Park (TW: Taman Wisata). See Protected areas Reef fish blennies (Blenniidae), 638, 642, 1419 butterflyfishes (Chaetodontidae), 638, 639, 642, 781 cardinalfishes (Apogonidae), 239, 241, 638, 642, 644, 650, 882, 1419 damselfishes (Pomacentridae), 638, 639, 640, 642, 643, 644, 882 glassfishes (Ambassis), 649, 868, 878, 882, 1164, 1407, 1418 gobies (Gobiidae), 229–242, 638, 642, 644, 646, 647, 648, 650, 781, 860, 862, 863, 864, 865, 875, 882, 1164, 1408–1409, 1419, 1420 (see also Gudgeons) groupers (Serranidae), 638, 643, 793, 794, 1216 moray eels (Muraenidae), 639, 1406 nurseryfish (Kurtus), 650, 1407, 1421 parrotfishes (Scaridae), 638, 639, 642, 787, 807 pipefishes (Syngnathidae), 235, 642, 650, 813, 1406 snappers (Lutjanidae), 638–639, 643, 644, 813, 882, 1419

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1462 / index Reef fish (cont.) surgeonfishes (Acanthuridae), 638, 639, 807 wrasses (Labridae), 638, 642 wrasses, Napoleon/Humphead (Cheilinus undulatus), 643, 787, 788, 793, 794, 1216 See also Fishing Regeneration, 8, 9, 353, 367, 506, 521, 677, 829, 904, 920, 952, 956, 957, 980, 991, 993–994, 1004–1009, 1013–1014, 1018, 1060, 1061, 1093, 1203. See also Fire; Pioneer species; Secondary vegetation Regional Autonomy. See Autonomy, Regional Religion Christianity, 78, 112, 113, 1099, 1116, 1117 Hinduism, 113 indigenous, 112–113, 115, 1141 Islam, 11, 112, 113, 1112, 1117 See also Christian missionaries REPELITA. See Development planning RePPProT. See Transmigration Resins, 849, 1161. See also Copal Rhizophora, 9, 264, 507, 557, 824, 826–827, 828, 830, 831, 832, 833, 836, 880 Rhododendron, 9, 36, 296, 389–392, 447, 449, 963, 967, 990, 992, 999, 1000, 1001, 1008, 1012, 1013, 1021, 1028, 1029, 1030, 1031, 1032, 1033, 1034, 1035, 1036, 1037, 1038, 1044, 1046, 1047 Rice, 171, 666, 667, 757, 874, 878, 1138, 1141, 1142, 1143, 1144, 1156 Ringtail possum. See Possums and gliders Roads, 31, 56, 76, 77, 1151, 1177–1181, 1182, 1184, 1185, 1214–1215, 1331, 1332, 1334, 1352 effects of, 872, 883, 885, 1177–1181, 1210, 1214, 1215 roadside as habitat, 308, 309, 335, 458, 868–869, 877, 958, 1025 See also Disturbed habitat Rodents. See Murids Rosewood, 9, 883. See also Pterocarpus Rossel Island. See Louisiade Islands Rouffaer River. See Tariku River Rubber, 41, 42, 1103, 1156, 1157 Rubiaceae (Hydnophytum, Myrmecodia), 841, 919, 949, 967, 991, 1008, 1028, 1036. See also Ants, and plants Rumberpon Island, 69, 1429 Rusa Deer (Cervus timoriensis), 723, 886, 1056–1061, 1161, 1330, 1333–1334, 1338

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Saccharum. See Sugar cane Sago (Metroxylon sagu), 367, 810, 879, 901, 910, 917, 920, 923, 925–928, 933, 934, 935, 1094, 1351–1352, 935 cultivated, 114, 118, 119, 367, 1125, 1129– 1130, 1137–1140, 1142, 1143, 1156, 1161, 1332, 1351–1352, 1356 moths (pest), 1365 plantations, 1145 Sahul, 121, 122, 246, 391, 813, 1096 Salawati Island. See Raja Ampat Islands Samoa, 32, 33, 140, 547, 582. See also American Samoa Sand mining, 814, 1333, 1334 Saratoga. See Bony tongues Sarawak, 352, 790, 934, 966, 967, 1354 Sarmi, 109, 110, 773, 810, 1151 Saruwaged Mts (Huon Peninsula), 154, 209, 210, 215, 234, 325 Savanna, 6, 8, 9, 167, 202, 251, 263, 264, 276, 384, 429, 432, 440, 442, 458, 468, 473, 487, 489, 490, 521, 572, 575, 577, 579, 587, 598, 599, 603, 617, 631, 661, 664, 733, 754, 757, 763, 886, 892, 902, 919, 946, 965, 979, 1018, 1025, 1033, 1036, 1038, 1054–1063, 1087, 1089, 1091, 1136, 1231, 1246, 1314, 1327, 1328, 1329, 1330, 1335, 1365, 1431, 1432. See also Grasslands Schefflera, 50, 293, 296, 982, 990, 998, 999, 1012, 1029, 1030, 1031, 1032, 1033, 1034, 1035, 1036 Schouten Islands, 141, 142, 567, 599 Seagrass, 499, 641, 759, 761, 800–818, 846, 901–902, 1208, 1216 Sea level, changes in, 121, 126, 180, 249, 250, 255, 256, 264, 481, 496, 497, 566, 640, 648, 729, 774–775, 814, 817, 825, 1223 Sea snakes, 479, 507, 564, 580, 585, 586, 598, 1415 Secondary vegetation, 264, 317, 350, 385, 387, 394, 399, 406, 412, 414, 457, 458, 468, 470, 506, 619, 720, 761, 951, 952, 954, 955–956, 957, 958, 959, 965, 977, 980, 985, 989, 990, 992–995, 1003, 1008, 1012–1016, 1040, 1087, 1094, 1095, 1351–1352, 1358, 1360. See also Fire; Pioneer species; Regeneration; Seral vegetation; Succession Sedimentary rock, 150, 167, 168, 220, 266 Sedimentation, 125, 264, 501, 523, 777–778, 792, 804, 806, 810, 814, 847, 848, 902, 925, 933. See also Pollution and sedimentation, causes of; Siltation

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index / 1463 Seed dispersal, 357, 377, 392, 465–466 by birds, 352, 369, 392, 396–397, 402, 406, 413, 460, 466, 667–668 by mammals, 352, 369, 392, 397, 402, 406, 413, 432, 460, 466 by reptiles, 402 by water, 352, 357, 413, 460 by wind, 357, 377, 392, 402, 432, 460 See also Pollination Sentani, 182, 186, 189, 232, 261, 264, 1089, 1093, 1110, 1120, 1121, 1152, 1209, 1218, 1354, 1365, 1366, 1367. See also Lake Sentani Sentani (people), 108, 110, 11, 112, 117, 1142 Sepik River/Basin, 32, 35, 43, 150, 209, 210, 215, 216, 220, 234–235, 236, 404, 417, 489, 648, 656, 660, 661, 663, 827, 889, 890, 967, 984, 1089, 1223 Seral vegetation, 956, 957, 995, 1014, 1021, 1039, 1040, 1041–1042, 1048. See also Climax species/communities; Fire; Pioneer species; Regeneration; Secondary vegetation; Succession Seram. See Maluku Sharks, 638, 642, 643, 644, 788, 793, 794, 1216, 1405 freshwater (Carcharinus), 6, 888, 1405, 1416 Shifting cultivation, 172, 329, 521, 883–884, 956, 958, 995, 1055, 1126, 1127, 1129– 1130, 1132, 1134, 1136, 1137, 1329 Shrimp (prawns), 409, 506, 511, 555, 556, 589, 590, 591, 621, 781, 803, 807, 810, 812, 813, 838, 841, 842, 843, 844, 845, 846, 850, 869, 880, 881, 885, 902, 1074, 1150, 1163, 1164, 1207, 1213. See also Crustaceans Siltation, 264, 641, 759, 793, 883, 884, 886, 1208, 1212, 1216. See also Sedimentation Sirunki, 260, 261 Skink, 489, 575, 577, 593, 595, 600, 604, 619 Sleumer, H. O., 76, 80, 89, 90, 389, 390, 416, 423 Sloanea, 381, 386–387, 915, 932, 935, 936, 937, 940, 949, 952, 953, 963, 964, 968, 973, 974, 975, 982, 985, 986, 988, 1012 Snakehead (Channa), 651, 878, 889, 890–891, 1217, 1419, 1421 Snakes biology/ecology, 508, 586, 594–599, 1356, 1359 endemism, 479, 600, 602 introduced, 678, 1218, 1366 as prey, 592, 595, 597, 619

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taxa/distribution, 479, 480, 482, 483, 485, 564, 565, 566, 567, 580–586, 1405, 1414–1415 trade, 604 See also Sea snakes Snappers (Lutjanidae), 638–639, 643, 644, 813, 882, 1419 Snow Mts. See Central Cordillera (Merauke Range) Society Islands, 400, 639, 784 Soils, 165–176, 966 cloud forest, 978 in conservation targets areas, 1234, 1241, 1242 and extractive industries, 1179, 1207, 1213 fertility/nutrients in, 142, 170–174, 329, 760, 828, 947, 964, 965, 978, 983–984, 993, 1005, 1037 heath forest, 962, 964, 965, 966 and human habitation/agriculture, 130, 182, 188, 189, 194, 760, 946, 995, 1126, 1129, 1135 and plant growth, 8–9, 142, 182, 308, 309, 327, 328, 329, 346, 962, 982, 997, 1025, 1013, 1014, 1038, 1048, 1055, 1327 swamps/peat, 910, 917, 920, 933, 934, 942, 1012, 1017, 1033, 1036, 1043, 1046 See also Ultramafics (ultrabasic); Water balance Solar radiation, 182, 776, 829, 964, 1003, 1005, 1052, 1222 Solomon Islands, 122, 137, 158, 207, 269, 313, 322, 324, 339, 357, 371, 373, 374, 375, 376, 381, 382, 383, 384, 385, 386, 387, 411, 417, 423, 467, 515, 534, 564, 571, 573–574, 577, 579, 580, 582, 584, 594, 605, 619, 628, 710, 720, 782, 783, 784, 958, 983, 1200, 1365 Bougainville, 424, 548, 564, 565 Buka, 122, 546 Solomons Arc/Plate/Trench, 137, 142–147, 154–156 Solomon Sea, 144, 145, 147, 158, 221, 541– 545, 552, 1314 Sonneratia, 429, 430, 826, 827, 830, 831, 832, 833, 836, 879, 880, 905, 906, 910 Sorong (Vogelkop), 257, 366, 385, 520, 547, 604, 628, 629, 630, 646, 730, 808, 809, 887, 1103, 1107, 1109, 1111, 1114, 1119, 1120, 1151, 1154, 1157, 1158, 1160, 1161, 1164, 1209, 1212, 1213, 1261, 1319, 1320, 1365, 1433 Sorong Fault, 148, 157, 159, 198, 199, 200, 213

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PAGE 1463

1464 / index South America, 180. See also Amazonia; Andes; Brazil; Colombia; Ecuador; Peru Special Autonomy. See Autonomy, Special Speciation, 203–204, 205, 252, 264, 286, 322, 371, 404, 496, 520, 532, 647, 656, 657, 738. See also Species radiations; Vicariance Species radiations, 208, 252, 436, 445, 566, 574, 577, 579, 580, 586, 599, 655, 668, 673, 689, 718. See also Speciation; Vicariance Sri Lanka, 312, 339, 382, 408, 411, 473, 547, 548, 1251 Star Mountains, 260, 325, 459, 521, 537, 569, 599, 602, 660, 692–702, 708, 716, 717, 719, 735, 736, 744, 745, 748, 949, 957, 966, 1004, 1018, 1066, 1070, 1078 State University of Papua (UNIPA), 270, 276, 516, 1108, 1190, 1245, 1312, 1366 Stick insects (Phasmatodea), 10, 518, 624 Stomatopods (mantis shrimp), 499–502 Strickland River, 714, 734, 1416 Strict Nature Reserve (Cagar Alam). See Protected areas Subalpine ecosystems/vegetation. See Alpine and subalpine ecosystems/vegetation Succession, 288, 381, 385, 631, 785, 806, 940, 950, 951, 956, 990, 993–994, 995, 1007, 1013, 1014, 1040, 1130. See also Fire; Pioneer species; Regeneration; Secondary vegetation Sudirman Range, 4, 8, 197, 201, 488, 1246. See also Central Cordillera (Merauke Range); Mt Jaya; Mt Trikora Sugar cane (Saccharum), 67, 80, 522, 943, 950, 951, 1130, 1134, 1135, 1136 Sugar glider. See Possums and gliders Sukarnopura. See Jayapura Sula Islands. See Banggai/Togian/Sula Islands Sulawesi, 114, 148, 157, 158, 208, 274, 277, 280, 282, 285, 289, 312, 382, 383, 385, 387, 417, 496, 501, 521, 534, 573, 579, 582, 630, 640, 644, 646, 672, 689, 720, 773, 784, 790, 794, 800, 881, 901, 933, 948, 962, 977, 1076, 1101, 1121, 1140. See also Banggai/Togian/Sula Islands; Makassar Sumatra, 138, 272, 274, 277, 280, 282, 285, 289, 312, 394, 400, 542, 640, 689, 760, 766, 784, 793, 794, 933, 948, 962, 965, 977, 1074, 1205, 1207, 1208, 1239, 1286, 1318, 1322, 1350, 1356, 1357 Sunda Arc, 138, 153, 158

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INDX

Sundaland, 138, 145, 246, 367, 1239. See also Bali; Borneo; Java, Malaysia; Nusa Tenggara; Sumatra Sunda Shelf, 359, 566, 585, 689 Supiori. See Biak/Supiori Supreme Court, 1288, 1291 Supulah Hill, 174, 260, 1089, 1090, 1094 Surgeonfishes (Acanthuridae), 638, 639, 807 Swamps/swamp forest, 6, 9, 114, 116, 121, 255, 258, 261, 264, 756, 757, 763, 873– 874, 877–880, 884, 892, 901, 910–943, 946, 950, 1173–1176, 1180, 1223, 1231, 1246, 1431, 1432 animals in, 574, 582, 583, 588, 590, 591, 596, 598, 710, 841 plants in, 350, 367, 368, 416, 440, 441, 468, 470, 833, 951, 960, 980, 990, 993, 1009–1013, 1054–1057, 1058–1061, 1329, 1334, 1351 See also Archeological sites and agriculture, swamps; Mangroves; Melaleuca; Nypa; Peat, swamps; Sago Swart Valley, 56, 736, 746 Sweet potatoes, 11, 185, 1092, 1093, 1094, 1127, 1128, 1129, 1130, 1132–1137, 1139, 1142, 1143, 1144, 1360 moths (pest), 1365 Swidden agriculture. See Agriculture, traditional, swidden Symbiosis on coral reefs, 781 mixed-species bird flocks, 672–673 wasps and figs, 404, 405–406 See also Ants, and plants; Pollination; Seed dispersal Syngnathidae. See Pipefish Syzygium, 292, 296, 429, 431, 432, 879, 903, 911, 912, 915, 919, 920, 923, 926, 929, 932, 933, 935, 937, 938, 939, 940, 949, 950, 952, 954, 955, 963, 968, 969, 970– 973, 974, 975, 981, 984, 985, 986–988, 990, 991, 995, 1008, 1009 Tachyglossidae. See Echidnas Tagula Island. See Louisiade Islands Taiwan, 312, 313, 339, 346, 384, 577, 783 Taman Buru, 1257, 1259, 1430 Tami Islands (Huon Gulf), 35, 40 Tami River/Basin, 49, 50, 63, 214, 467, 1244 Tamrau Mts, 5, 153, 157, 159, 198, 199, 200, 213, 487, 599, 601, 602, 659, 711, 714, 728, 732, 1244, 1246, 1268, 1270, 1427, 1431 Tamrau Terrane, 149, 198

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index / 1465 Tangguh LNG project, 1111, 1117, 1119, 1155, 1167, 1168, 1170, 1212–1214 Tapa cloth, 406 Tari Basin, 249, 261, 1090 Tariku (Rouffaer) River, 5, 173, 198, 215, 261, 1246, 1271 Taritatu (Idenburg) River, 5, 173, 215, 261, 269, 385, 387, 534, 724, 729, 736, 861, 979, 981, 1246 Taro (Colocasia esculenta), 189, 878, 890, 1091, 1092, 1125, 1129, 1130, 1135, 1136, 1137, 1138, 1139, 1144 Tasmania, 257, 391, 1044 Tasman Line, 144, 151 Tasman Sea, 145, 151, 152 Tectonic history, and endemism, 158, 160, 196, 201, 202, 220–223, 288–289, 320, 566. See also Biogeographic history; Biogeography; Endemism; Geological formations Tectonic plates Australian Plate, 4, 143, 196, 201, 481, 532, 640, 646, 658, 662, 825 Caroline Plate, 137, 145–148, 154, 159, 774 Indo-Australian Plate, 137, 774, 1151 Pacific Plate, 4, 137, 143, 148, 150, 156, 157, 201, 647, 774, 1151, 1199 Philippine Sea Plate, 137, 145, 774 Solomons Arc/Plate/Trench, 137, 142–147, 154–156 See also Australian craton; Geological formations Teijsmanniodendron, 929, 933, 937, 939, 952, 954, 955, 968, 969, 970 Telefomin, 993, 1004, 1076, 1079, 1080, 1094 Telepak, 1115, 1205, 1206, 1207 Tembagapura, 7, 186, 193, 222, 736, 977, 998, 1009, 1014, 1017, 1018 Teminabuan, 645, 917, 920, 962, 1151, 1434 Terminalia, 903, 904, 905, 906, 907, 910, 911, 912, 913, 915, 916, 917, 918, 920, 921, 923, 924–927, 929, 932, 933, 935, 948, 952, 954, 957, 984 Termites (Isoptera), 167, 518, 539, 592, 594, 710 Ternate. See Maluku Terranes Adelbert-Finisterre Terrane, 143, 148, 155, 215, 221 Bowutu Terrane, 153 East Papua Composite Terrane, 144, 218, 487 Misool Terrane, 149, 151, 288 Tamrau Terrane, 149, 198 See also Geological formations

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Tertiary (Cenozoic), 100, 138, 142, 144, 146, 149, 150, 152, 153, 154, 156, 157, 159, 193, 219, 220, 221, 222, 246, 247, 248, 306, 481, 483, 1025 Eocene, 143, 144, 145, 152, 153, 154, 158, 159, 220, 248, 473, 481, 566, 574, 1025 Miocene, 138, 141–144, 146, 147, 149, 152, 153, 155, 156, 157, 158, 159, 198, 215, 220, 222, 246, 247, 248, 249, 252, 320, 566, 571, 574, 575, 580, 658, 800, 872, 1025 Oligocene, 138, 143, 144, 145, 146, 147, 151–154, 158, 159, 201, 218, 220, 222, 247, 580, 584 Pliocene, 138, 143, 144, 148, 154, 155, 159, 201, 214, 215, 222, 248, 249, 252, 519 Thailand, 277, 313, 339, 352, 400, 547, 548, 639, 1074, 1350 Theobroma. See Cocoa Threatened species. See Endangered and threatened species Thrips (Thysanoptera), 352, 396, 465, 518, 1384 Thylogale. See Kangaroos and wallabies Tidore, 16, 17, 23 Tilapia (Oreochromis), 651, 871, 873, 878, 887, 888, 889, 890, 1217, 1421 Timika, 5, 6, 8, 9, 186, 264, 366, 384, 385, 387, 645, 652, 810, 814, 861, 865, 866, 869, 885, 889, 890, 892, 957, 958, 1111, 1112, 1114, 1117, 1140, 1142, 1209, 1212, 1217, 1218 Timor, 138, 384, 386, 571, 574, 578, 584, 646, 849, 873, 881 Tiom, 171, 247, 1092, 1128 Tirawiwa River, 154, 867 Tobacco, 35, 37, 1134 Toe´ Cave, 123, 125, 126, 128 Togian Islands. See Banggai/Togian/Sula Islands Tokay gecko, 586, 1367–1368 Tonda Wildlife Management Area (PNG), 1059, 1060, 1261, 1330–1337, 1339– 1342, 1344–1345 Tonga Arc, 138, 141 Tonga Islands, 138, 141, 313 Torassi River. See Bensbach River Torres Strait, 137, 307, 578, 580, 628, 646, 802, 811, 817, 846 Torricelli Mts, 154, 155, 209, 210, 214, 215, 221, 233, 468, 489, 660, 663, 708, 716, 717, 729, 730, 732 Tourism, 1150, 1203, 1254, 1323, 1340 dive, 644, 1216, 1323

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1466 / index Trade, 41, 112, 113, 114 early, 11, 16, 17, 19, 25, 33, 34, 112 See also Wildlife trade Trade Winds. See Winds, Trade Traditional community associations (LMAs), 1337, 1339–1340, 1345 ‘‘Traditional Community Reserve’’ designation, 1323 Traditional medicine, 89, 91, 849, 116, 407, 824, 1203, 1214, 1280 Trans-Fly, 6, 8, 59, 186, 197, 202, 209, 210, 217, 218, 239, 341, 432, 488, 489, 490, 648, 661, 663, 690, 707–709, 710, 711, 713, 717, 719, 723, 733, 734, 746, 747, 878, 1054–1063, 1231, 1246, 1247, 1314, 1327–1346, 1365, 1416–1421, 1422– 1424 Transmigration, 12, 68, 79, 794, 888, 1112– 1113, 1115, 1118, 1140, 1178, 1191, 1213, 1234, 1243, 1286, 1329, 1333 and introduced species, 586, 889 Tree kangaroo (Dendrolagus). See Kangaroos and wallabies Triassic, 144, 150, 151, 335, 574 Tri-National Wetlands Program (TNWP), 1335–1336, 1339, 1342–1343, 1345 Triton Bay, 7, 200, 488, 730 Triton Lakes, 645, 649, 652 Trobriand Arc, 142, 143, 145 Tubers. See Cassava; Sweet potato; Taro; Yams Turtles biology/ecology, 402, 507, 590–591, 757, 803, 804, 805, 808, 813, 815, 901, 1242 conservation, 1242, 1315 endemism, 482, 603 fossil, 30 taxa/distribution, 480, 565, 567, 574, 1366, 1405, 1411–1412 trade/shells/eggs, 11, 603, 604, 1216, 1219, 1276, 1285, 1286, 1366 See also Green Sea Turtle; Hawksbill Turtle; Olive Ridley Turtle Typhoons. See Cyclones Ultramafics (ultrabasic), 141, 142, 143, 168, 169, 173, 220, 282, 287, 288, 412, 465, 470, 949, 958, 959, 979, 984, 994, 997, 1005, 1093, 1384. See also Soils UNCEN. See Cenderawasih University Undisturbed habitat, 310, 314, 521, 619, 627, 631, 643, 716, 788, 861, 885, 966, 1003, 1004, 1199, 1205, 1237, 1254. See also Disturbed habitat

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INDX

UNIPA. See State University of Papua University of Papua New Guinea (UPNG), 87, 94 University of Technology (UniTech, Lae), 87 Urbanization, 1111, 1113, 1170 Vaccinium, 296, 389, 390, 391, 392, 963, 987, 990, 992, 1008, 1012, 1013, 1021, 1027, 1029, 1036, 1037, 1039, 1040, 1043, 1044, 1047 Van Rees Mts, 5, 154, 157, 159, 198, 200, 209, 210, 214, 216, 232, 487, 599, 601, 602, 659, 690, 711, 728, 729, 1200, 1268 van Royen, P., 70, 76, 80, 86, 88, 89, 90, 93, 389, 464, 465, 942, 949, 1055 Vanuatu, 16, 142, 424, 678, 783, 1200 Varanids (monitor lizards), 575, 579–580, 592, 594, 602, 617–634, 1414 Vicariance, 199, 203, 204, 647, 662, 664, 1235. See also Habitat loss, fragmentation; Speciation; Species radiations Vogelkop, 3, 5, 6, 7, 9, 11, 14, 16, 108, 114, 123, 124, 127, 143, 144, 146, 147, 148, 149, 150, 151, 153, 156, 157, 158, 159, 160, 197, 198, 199, 200, 202, 203, 204, 209, 210, 212, 213, 216, 218, 221, 229– 231, 246, 248, 250, 251, 257, 260, 264, 270, 282, 288, 336, 339, 382, 387, 411, 417, 442, 459, 468, 487, 488, 489, 490, 499, 501, 520, 533, 534, 546, 574, 575, 591, 599, 600, 601, 602, 645, 648, 649, 656, 658, 659, 660, 661, 662, 690, 692– 698, 699, 703, 706, 710, 711, 714, 715, 717, 718, 719, 723, 728, 729, 730–733, 736, 737, 772, 773, 784, 792, 794, 808, 861, 871, 880, 885, 887, 890, 892, 945, 950, 992, 1066, 1068, 1070, 1078, 1102, 1103, 1116, 1120, 1121, 1137, 1199, 1200, 1212, 1218, 1243, 1244, 1246, 1261, 1268 Volcanics, 141, 154, 156, 159, 165, 172, 214, 220, 966, 1005, 1065 Volcanoes/volcanism, 8, 143, 147, 566, 647 volcanic activity, 62, 63, 466, 946, 1203, 1221 volcanic ash, 167, 171, 946, 951, 1005 volcano chains/arcs/islands, 138, 140, 141, 153, 220, 662, 771–772 See also Geology; Island arcs; Ophiolites; Volcanics Waigeo Island. See Raja Ampat Islands Walking Catfish (Clarias), 889, 892, 1217, 1421. See also Catfishes Wallabies. See Kangaroos and wallabies

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index / 1467 Wallace, Alfred Russel, 11, 14, 22, 23, 37, 567, 655–656, 659, 689 Wallacea, 204, 1239. See also Maluku; Nusa Tenggara; Sulawesi Wallacea Hotspot, 1200 Wallace’s Line, 307, 411, 480, 496, 497, 646, 659, 689 Wamena (Baliem Valley), 168, 170, 174, 247, 260, 604, 1038, 1066, 1070, 1076, 1089, 1151, 1178, 1179, 1180, 1215 Wandammen (people of), 108, 110, 111, 112, 113, 114 Wandammen Mts, 5, 659, 663, 690, 718, 732, 1268. See also Mt Wondiwoi Wandammen Peninsula, 3, 149–150, 159, 197, 200, 213, 366, 487, 488, 534, 569, 599, 601, 602, 603, 809, 1246 Wapoga River, 153, 154, 216, 485, 589, 601, 645, 652, 861, 864, 866, 867, 1312, 1323 Waropen (Cenderawasih Bay), 9, 630, 809, 1158, 1160, 1209, 1312 Waropen (people of), 108, 109, 110, 111, 112, 113, 114, 118, 119 Wasmannia, 678 Wasps, 451, 519 fig wasps (Agoninae), 404, 405–406 Wasur National Park, 713, 1055, 1056, 1059– 1062, 1262, 1271, 1317, 1328, 1330, 1331–1342, 1344, 1345, 1433 Water balance, 166, 167, 170, 181, 182, 189, 933, 940, 965, 966, 978, 1007, 1017, 1041, 1058. See also Rainfall; Soils Wau Ecology Institute (PNG), 15, 94, 98, 217 Bishop Museum Field Station, 79, 84, 94, 97 Wau Valley/Wau-Bulolo district, 248, 252, 270, 885, 980 Weber, Max, 37, 48, 638, 645, 646 Weber’s Line, 646, 890 Weyland Mts, 156, 159, 168, 187, 201, 204, 209, 210, 216, 236, 246, 257, 487, 489, 521, 599, 601, 602, 712, 713, 714, 716, 717, 728, 732, 735–738, 1269, 1270, 1426 Wildlife Sanctuary (Suaka Margasatwa). See Protected Areas Wildlife trade, 1161, 1162, 1219, 1276–1277, 1278, 1283–1288, 1299–1300, 1302, 1303, 1304, 1309, 1333, 1336, 1365–1368 birds, 14, 23, 31, 41, 46, 55, 654, 655, 675, 678 (see also Birds of Paradise; Cassowaries; Lories; Parrots) fish, 643, 788, 793, 794, 886–887, 1164,

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1216 (see also Bony tongues; Rainbowfish; Sharks) herpetofauna, 569, 580, 603–605, 623, 624, 626, 629, 632–634, 1161 plants, 47, 454, 1219 (see also Orchids) as threat to biodiversity, 603, 1214, 1219 See also Enforcement; Legislation, Laws (UU) Wilson, Edward O., 97 theory of island biogeography, 785 Winds, 190, 978, 982 local, 181, 182, 190, 446, 845, 882, 902, 959, 977, 991, 1004, 1005, 1025, 1221, 1237 Trade, 7, 178, 179, 187, 184, 188 See also Cyclones; Dispersal, by wind; Monsoon Wissel Lakes. See Paniai Lakes Woodlark Island, 208, 209, 220, 245, 386 Worldwide Fund for Nature (WWF), 1055, 1231, 1240, 1241, 1246, 1253, 1260, 1330, 1331, 1333, 1335, 1337–1345 Wosi market, 1286, 1287 Wrasses (Labridae), 638, 642 Humphead/Napoleon Wrasse (Cheilinus undulatus), 643, 787, 788, 793, 794, 1216 Wuvulu. See Admiralty Islands WWF. See Worldwide Fund for Nature Xylocarpus, 827, 830, 831, 832, 833 YALI (environmental foundation) 1312, 1436 Yali (people), 82, 962, 1128, 1131, 1132, 1134, 1143, 1144 Yam (Dioscorea sp.), 11, 1129, 1130, 1134, 1137, 1138, 1139, 1144, 1332 Yapen Island (Cenderawasih Bay), 4, 5, 7, 154, 157, 197, 198, 200, 201, 209, 210, 214, 232, 257, 366, 387, 466, 487, 488, 489, 537, 569, 584, 591, 599, 601, 602, 603, 621, 630, 644, 645, 659, 692–696, 699–702, 705–706, 713, 714, 716, 717, 720, 724, 727, 729, 730, 733, 749, 792, 809, 813, 946, 991, 994, 1077, 1079, 1120, 1158, 1160, 1200, 1209, 1246, 1269, 1270, 1426, 1432 Yei, 1331, 1332, 1339 Ylang-ylang oil, 352 Yos Sudarso Bay (people of), 109, 112, 117 Yos Sudarso Island. See Dolok Island YPMD, 1112, 1113, 1219 Zaglossus. See Echidnas Zingiberaceae. See Gingers

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

INDX

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PS

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