Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate [1st ed. 2020] 978-3-030-25520-6, 978-3-030-25521-3

Table of contents :
Front Matter ....Pages i-xiv
Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina): Subtropical Mountain Forest (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 1-8
General Features of Serranías de Zapla Multiple Use Ecology Reserve (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 9-22
Bioclimatology (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 23-41
Geobotany of Serranías de Zapla Multiple Use Ecology Reserve: Flora and Vegetation (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 43-108
Biodiversity Analysis: A Geobotanic Interpretation (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 109-156
Final Remarks (Gabriela S. Entrocassi, Rosario G. Gavilán, Daniel Sánchez-Mata)....Pages 157-162
Back Matter ....Pages 163-191

Citation preview

Geobotany Studies Basics, Methods and Case Studies

Gabriela S. Entrocassi Rosario G. Gavilán Daniel Sánchez-Mata

Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate

Geobotany Studies Basics, Methods and Case Studies

Series Editor Franco Pedrotti, Department of Botany and Ecology, University of Camerino, Camerino, Italy Editorial Board Members S. Bartha, Vácrátót, Vácrátót, Hungary F. Bioret, University of Brest, Brest, France E. O. Box, University of Georgia, Athens, GA, USA A. Čarni, Slovenian Academy of Sciences, Ljubljana, Slovenia K. Fujiwara, Yokohama City University, Yokohama, Japan D. Gafta, “Babes-Bolyai” University, Cluj Napoca, Romania J. Loidi, University of Bilbao, Bilbao, Spain L. Mucina, University of Perth, Perth, Australia S. Pignatti, Università degli Studi di Roma “La Sapienza”, Roma, Italy R. Pott, University of Hannover, Hannover, Germany D. Sánchez Mata, Universidad Complutense Madrid, Madrid, Spain A. Velázquez, Centro de Investigación en Sciéncias Ambientales, Morelia, Mexico R. Venanzoni, University of Perugia, Perugia, Italy

The series includes outstanding monographs and collections of papers on a range of topics in the following fields: Phytogeography, Phytosociology, Plant Community Ecology, Biocoenology, Vegetation Science, Eco-informatics, Landscape Ecology, Vegetation Mapping, Plant Conservation Biology, and Plant Diversity. Contributions should reflect the latest theoretical and methodological developments or present new applications on large spatial or temporal scales that will reinforce our understanding of ecological processes acting at the phytocoenosis and vegetation landscape level. Case studies based on large data sets are also considered, provided they support habitat classification refinement, plant diversity conservation or vegetation change prediction. Geobotany Studies: Basics, Methods and Case Studies is the successor to BraunBlanquetia, a journal published by the University of Camerino from 1984 to 2011 in cooperation with the Station Internationale de Phytosociologie (Bailleul, France) and the Dipartimento di Botanica ed Ecologia (Università di Camerino, Italy) and under the aegis of the Société Amicale Francophone de Phytosociologie, the Société Française de Phytosociologie, the Rheinold-Tüxen-Gesellschaft and the Eastern Alpine and Dinaric Society for Vegetation Ecology. This series promotes the expansion, evolution, and application of the invaluable scientific legacy of the Braun-Blanquet school.

More information about this series at http://www.springer.com/series/10526

Gabriela S. Entrocassi • Rosario G. Gavilán • Daniel Sánchez-Mata

Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate

Gabriela S. Entrocassi Facultad de Ciencias Agrarias Universidad Nacional de Jujuy San Salvador de Jujuy, Argentina

Rosario G. Gavilán Departamento de Farmacología, Farmacognosia y Botánica Universidad Complutense Madrid, Spain

Daniel Sánchez-Mata Departamento de Farmacología, Farmacognosia y Botánica Universidad Complutense Madrid, Spain

ISSN 2198-2562 ISSN 2198-2570 (electronic) Geobotany Studies ISBN 978-3-030-25520-6 ISBN 978-3-030-25521-3 (eBook) https://doi.org/10.1007/978-3-030-25521-3 © Springer Nature Switzerland AG 2020 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Ahora digo limpio de corazón, los ojos puros, el nombre de los árboles de la tierra que habito, su alta serenidad, su lenta sombra y su resina cristalina y triste. Yo voy a la madera y de ella vengo. Doblado en la luz, quemado en arenales, con una sombra más entre los brazos como quien se recuerda con el alma del aire. Vengo desde las vigas cenicientas, caídas, asoléandose, con la baba brillosa todavía de los bueyes y desde las semillas de los naranjos viejos sembradas por carreros en Orán y por loros sobre un camino solo y sin regresos. Desde allí, desde el yuchán panzudo donde los peces miran su memoria de limo cuando los sapos rezan a la tierra, desde los urundeles serenísimos quema la voz alzada de chahuancos y tobas en el baile que muele maíces y dolores.

¡Oh pura levedad de los chañares! ¡Oh doliente algarrobo, sobre tu pensamiento los hermanos siguen muriendo para hacerse pájaros! Si es que digo quebracho y digo brea viene la sangre con sus polvaredas y vienen los abuelos pensativos doblados en la sal, juntando leña sobre la costra ardida que le crece a Santiago del Estero. Vengo desde el laurel que huele como el hombre, desde el fondo del cedro donde dormita la rosa su amanecer de greda y de guayacanes donde comienza el ébano. Vengo de allí, desde sus hojas vivas, vengo desde el incendio en paz de los lapachos cuando los tarcos se pierden en un tierno olvido lila. Yo sé de sus raíces por donde Dios camina lleno de barro y savia ciego y doliente, pero jubiloso. Yo sé de sus veranos interiores y de los vendavales cuajados en sus vetas cuando el hombre era apenas un blando mineral sobre la tierra, una tierna memoria enamorada. Voy a sus huesos verdes bajo el solazo que tritura cañas, me pierdo por la sombra rota de las papayas de cuyos frutos pende el semen de todas las primaveras venideras; me entierro entre bambúes

y por los molles lloro y en la orejas negras del pacará que trepo oigo los pasos de agua de los que están viniendo. Desde la aun callada certitud de la vida voy a sus huesos verdes con un iluminado destino de semilla. Entonces mi alegría se arrodilla en el fruto donde se cumplen dulces agonías. ‘LOS ÁRBOLES’ (Manuel J. Castilla, 1918–1980) ‘Our gratitude to the Yungas, we admire their lush greenness, spicy humidity, orange and brave rivers, impressive ravines, biblical storms, magical fauna and wonderful trees. They were and will continue to be the inspiration but also the clay to model life’ (G. Entrocassi).

Preface

The subtropical forest of northern Argentina is included within the buffer zone of the Reserva de la Biosfera de las Yungas and is part of a biological corridor connected with other international and national protected areas. It is included as forest maintenance area under conservation for sustainable management plan. Moreover, the territory protects the upper watercourses of the Bermejo River (Alta Cuenca del río Bermejo) and houses a high environmental and biological diversity with characteristic taxa of the Selva Montana and other floristic taxa from the lower vegetation belts (Selva Pedemontana) and even from the Chaquean biogeographical region where they have their upper distribution boundary. It also has a high forest and biological value as refuge and food source for many resident and migratory animals. We have studied Las Yungas forest throughout an environmental gradient framed within the Serranías de Zapla Multiple Use Ecology Reserve (‘Reserva Ecológica de uso múltiple Serranías de Zapla’, Jujuy province, Argentina). For this purpose, we follow the Braun-Blanquet phytosociological methodology and the updated bioclimatic proposals compiled into the Worldwide Bioclimatic Classification System. The methodological research design includes 120 vegetation transects where we recorded the phytosociological relevés. These relevés include all the taxa appearing in such selected plots grouped into different strata (trees, shrubs and herbs) and their abundance-dominance index values. From this data richness, life form and frequency of taxa were also analysed. We have identified 257 taxa framed into 194 genera and 66 families. The study included the bioclimatic characterisation of the selected territory; each vegetation transect was bioclimatically characterised by indices, mainly thermicity index (It) and ombrothermic index (Io) using standard temperature and precipitation records from the available climatic stations that were extrapolated by standard methods to any transect. Floristic and vegetation data have been used to describe forest communities of Las Yungas. A vegetation matrix was analysed by multivariate methods to identify such plant communities and also to frame them in the South America vegetation context. These analyses include hierarchical classifications which allow us to identify similarities among relevés separating homogeneous groups according to their ix

x

Preface

floristic composition. Canonical correspondence analysis showed how the environmental gradient influenced both the floristic composition and the distribution pattern of the recognised communities. Both analyses also served to delimit plant species groups whose abundance and distribution were associated with altitudinal and bioclimatic differences supporting their adscription as ‘characteristic’ or ‘differential’ of forest types. We have recognised thirteen different plant communities distributed along an altitudinal gradient of 600 m asl. They are micro- and mesoforests, mainly semideciduous and seasonal evergreen forests belonging to Selva Montana and Bosque Montano biogeographical districts within the Yungas biogeographical province (Amazonian domain, Neotropical region) correlated with the Subandean and Montane Pluviseasonal Bolivian-Tucuman vegetation of the Bolivian-Tucuman biogeographical province (South Andean region, Neotropical Subkingdom). Bioclimatically, they are found within the lower and upper mesotropical, lower and upper subhumid and lower humid bioclimates (Tropical macrobioclimate). Nine recognised plant communities are distributed within the lower mesotropical, lower and upper subhumid bioclimates (1015-1275 m asl; It ¼ 399-429; Io ¼ 4,7-5,5) as lowland forests developed mainly on warm and subhumid territories of the Selva Montana (selva basal) in the south and western study area. Four plant communities are distributed within the upper mesotropical, upper humid bioclimate (1260-1620 m asl; It¼352-395; Io¼7,7-8,7) as highland forests developed on more temperate and wet territories of the Selva Montana (selva basal) and Bosque Montano, at the central part of the study area. As conclusion, the following phytosociological associations are proposed as new for South America: Enterolobio contortisilici-Anadenantheretum cebilis; Schino bumeloidis-Allophyletum edulis; Xylosmo pubescentis-Blepharocalycetum salicifolii; Jacarando mimosifoliae-Vassobietum breviflorae; Erythrino falcataeTipuanetum tipi; Schinetum myrtifolio-gracilipedis; Juglandi australisBlepharocalycetum salicifolii; Zanthoxylo cocoi-Blepharocalycetum salicifolii; Tecomo stantis-Anadenantheretum cebilis; Myrciantho pseudomatoiBlepharocalycetum salicifolii; Cinnamomo porphyrium-Blepharocalycetum salicifolii; Pruno tucumanensis-Podocarpetum parlatorei; Salici humboldtianaeAcacietum aromae. The composition and distribution of the plant communities of the study area are determined by the environmental gradient accounted for by the altitude and climate (mainly expressed by the thermicity, It, and ombrothermic indices, Io). In certain areas the topographic aspect of slopes together with the local geomorphology or particular edaphic characteristics also explains some differences in floristic composition of communities. The northern part of the study area is located within the buffer zone of the Yungas Biosphere Reserve and is part of a biological corridor that connects not only Argentinian but also Bolivian protected areas. The reserve is included as a forest maintenance area under conservation and/or sustainable management plans (Category II-Yellow, Territorial Planning Plan for Native Forests of the Jujuy Province). It is also part of the territory declared as Model Forest, framed in the National Model Forest Program. River courses are also included in this forest

Preface

xi

model, mainly the Bermejo River upper basin, with a high environmental and biological diversity, including species from the Selva Montana and Selva Pedemontana and even from the Chaqueña Biogeographic Region, which find their upper limit of distribution there. This research contributes to the knowledge of the subtropical mountain forests of South America. It opens a great opportunity to conduct other ecological and phytosociological research works in unexplored territories from a geobotanical perspective. San Salvador de Jujuy, Argentina Madrid, Spain Madrid, Spain

Gabriela S. Entrocassi Rosario G. Gavilán Daniel Sánchez-Mata

Contents

1

2

3

4

Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina): Subtropical Mountain Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2 Las Yungas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3 Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4 Las Yungas Biosphere Reserve (RBYungas) and the Jujuy Model Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

.. .. ..

1 1 2

..

4

..

7

General Features of Serranías de Zapla Multiple Use Ecology Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1 Geomorphology, Geology, Hydrology and Soils . . . . . . . . . . . . . 2.2 Vegetation and Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Management: Situation of Serranías de Zapla in the Context of Protected Areas and the Territorial Ordination Plan of Jujuy Province . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bioclimatology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 General Bioclimatic Features from Las Yungas . . . . . . . . . . . . . 3.3 Bioclimatic Features of Serranías de Zapala Multiple Use Ecological Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 Bioclimatic Features from Jujuy Province . . . . . . . . . . . . . . . . . 3.5 Final Remarks on Jujuy Bioclimates . . . . . . . . . . . . . . . . . . . .

9 9 14

19

. . .

23 30 30

. . .

31 33 38

Geobotany of Serranías de Zapla Multiple Use Ecology Reserve: Flora and Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1 Field Data Collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.1 Identification, Species Richness and Life Forms of Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.2 Analysis of Floristic-Phytosociological and Bioclimatic Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

43 43 44 45 xiii

xiv

Contents

4.1.3 Synthesis of the Floristic-Phytosociological Data . . . . . . . 4.1.4 Floristic Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2.1 Vegetation Structure (Stratification) and Floristic Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2.2 Richness in the Relevés . . . . . . . . . . . . . . . . . . . . . . . . . 4.2.3 Frequency of the Recorded Species and Biological Spectrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.1 Floristic-Phytosociological Data. Hierarchical Cluster Analysis (HCA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.2 Floristic-Phytosociological and Bioclimatic Data: Analysis of the Gradient. Canonical Correspondence Analysis (CCA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.3 Relationships Between Species and Environmental Variables Defined by Axis 1 on the CCA Ordination Diagram . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.4 General Synthetic Comparative Table . . . . . . . . . . . . . . . 4.3.5 Delimitation of the Plant Communities . . . . . . . . . . . . . . 4.3.6 Nomenclature of the Communities . . . . . . . . . . . . . . . . . 4.3.7 List of Associations Proposed for the Study Area . . . . . . .

47 48 49

Biodiversity Analysis: A Geobotanic Interpretation . . . . . . . . . . . . . 5.1 Floristic Composition in Detail . . . . . . . . . . . . . . . . . . . . . . . . . 5.2 Species Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3 Species Frequency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4 Biological Spectrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.5 Analysis, Interpretation and Characterisation of the Plant Communities in the Study Area . . . . . . . . . . . . . . . . . . . . . . . . . 5.6 Influence of the Environmental Gradient on the Composition and Distribution of the Plant Communities in the Study Area . . . . 5.7 General Diagnosis of the Plant Communities in the Study Area . . . 5.8 Terrestrial Vegetation in the Lower Mesotropical Bioclimatic Belt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.9 Terrestrial Vegetation in the Upper Mesotropical Bioclimatic Belt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.10 Final Considerations on Terrestrial Plant Communities in the Lower and Upper Mesotropical Bioclimatic Belt . . . . . . . . 5.11 Riparian Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

109 109 113 115 116

4.2

4.3

5

6

49 49 51 51 51

54

57 60 62 66 81

116 124 127 128 143 153 153

Final Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

Appendix A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 Appendix B . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 Appendix C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Chapter 1

Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina): Subtropical Mountain Forest

1.1

Introduction

Vegetation is the most easily recognisable component of the ecosystem, and is the result of the joint action of environmental factors: it reflects climate, the nature of the soil, the availability of water and nutrients, and biotic and anthropic factors. It therefore acts as an indicator of the characteristics and status of ecosystems (Whittaker 1975; Matteucci and Colma 1982). The study of vegetation is of exceptional importance as it is a vital part of the ecological system: it captures, transforms and stores solar energy, protects the soil, regulates the local climate and serves as sustenance and refuge for fauna; it is the origin of raw materials for humans and a source of spiritual and cultural well-being thanks to its aesthetic, recreational and educational value. It is the essential basis for obtaining information on the composition, structure, dynamic and distribution of plant communities within a specific plant formation (Braun-Blanquet 1964; Whittaker 1975; Matteucci and Colma 1982). Vegetation studies serve numerous purposes, basically for research and development, and are closely linked to the conservation of the environment. The information obtained from these studies can be used to assess the vegetation for the purposes of conservation and to reach a diagnosis on its current state, with a view to designing strategies and programmes for conservation, ordination and territorial management and for planning the use of the territory (Whittaker 1973; MuellerDombois and Ellenberg 1974; Meaza Rodriguez 2000). These studies serve as the basis for creating maps of vegetation distribution within a given area, delimiting homogeneous ecological units and determining the degree of priority of its conservation. They also provide indicators on the environment, the effects of management (livestock, forestry production and so on) and different treatments (fertilisation, irrigation, reforestation and others), and supply valuable information for studies on environmental impact and the sustainable management of natural resources (forestry, management of wild fauna, etc.), among others. The detailed study of the composition, structure, dynamic and distribution of vegetation is therefore the © Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_1

1

1 Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina). . .

2

cornerstone of a greater understanding and a balanced management of ecosystems, of which humans also form part. The increasing anthropic pressure to which natural ecosystems are subject today makes this type of study absolutely essential (Matteucci and Colma 1982; Barbour et al. 1998).

1.2

Las Yungas

In South America, mountain cloud forests form a very broad type of vegetation that extends as rainforests and seasonal rainforests across the eastern and western slopes of the Andean mountain range from Venezuela to northwest Argentina. This plant formation has a notable originality and affinity from the phytogeographical and zoogeographical point of view, and contains great species richness and a large number of endemics, making it possibly one of the most biodiverse areas on Earth (Myers et al. 2000; Soutullo et al. 2008). These forests also constitute a genetic repository, contribute to the protection and stabilisation of the ecosystems in the region and play a key role in regulating climate and maintaining water resources (Brown and Kappelle 2001). Within this broad formation of South American forests, and at around 18
S in central Bolivia (Cochabamba), a dramatic shift occurs in the direction of the Andean mountain range from north to south, giving rise to a change in the climate and causing the disappearance of the rainforest and the establishment of seasonal rainforests on the eastern slopes of the Andean range (Navarro and Maldonado 2002). These seasonal rainforests belong to what is known as the Tucuman-Bolivian forest, in the phytogeographical province of Las Yungas (Hauman 1931; Cabrera and Willink 1980; Cabrera 1994), and constitute one of the characteristic vegetation types in the Bolivian-Tucuman province (tropical south-Andean biogeographic region; Rivas Martínez et al. 2011), an area that includes the eastern and western flanks of the tropical Andes and containing several ecological belts and ecosystems, of which some of the most significant are the valley, sub-Andean, montane and plain environments. Within these ecological belts, and depending on the latitude, altitudinal gradient and relief, there are a range of plant formations including particularly the aforementioned seasonal rainforests, which—unlike rainforests—have a semideciduous and seasonal evergreen character and undergo a period of drought coinciding with the coldest season. These seasonal rainforests are known in Argentina by the name of subtropical mountain forests as they lie south of the Tropic of Capricorn, or “mountain cloud or rain forests”, or “Tucuman-Bolivian forest”, and are popularly called “Las Yungas”, or simply “el monte” (meaning “uncultivated area”) by the local inhabitants. The distribution area of Las Yungas comprises the northwest of Argentina and they extend over the eastern slopes of the Sub-Andean and Pampean mountain ranges and the pre-mountain range, from the limit with Bolivia (22
S) to the northeast of the

1.2 Las Yungas

3

province of Catamarca (29
S), crossing the provinces of Salta, Jujuy and Tucumán. Altitudinally, they are distributed between 300 and 3000 m asl along a length of 700 km in a north-south direction and with a width of less than 100 km (Brown et al. 2002). Las Yungas represent one of the most important forest formations in the country, and the most extensive subtropical forest ecosystem in Argentina. They occupy 5.2 million ha (only 2% of Argentinian territory) and have the greatest biodiversity in Argentina along with the Paraná forest, as they contain approximately 50% of the country’s flora and fauna (Pacheco and Brown 2006). They have been identified as among the richest and most diverse biodiversity hotspots since it forms part of the ‘Tropical Andes Biodiversity Hotspot’ (Myers et al. 2000; Burkart 2006). Las Yungas are home to over 200 tree species, many of which are of significant interest for forestry, 80 varieties of fern, over 100 species of mammals, 500 species of birds and some 30 species of frogs and toads. Las Yungas also serve as a natural refuge for many vulnerable or endangered species such as the yaguareté or jaguar, the tapir, the green wing macaw, the alder amazon and the Muscovy duck, among others. Las Yungas have been inhabited by man for at least 8000 years, and yet in the last half century human activity has caused a sweeping transformation, leading to the disappearance of large swathes of native forest (www.proyungas.org.ar). The environmental diversity that characterises the mountain regions in northwest Argentina where the subtropical mountain forests grow is determined by the varying influence of an array of geophysical factors such as altitude, slope exposure, bioclimate, geomorphology, and soils, with altitude being the key to controlling the vegetation and the factor that primarily determines the bioclimate in these mountain regions. In turn, the bioclimate is the main determinant of the vegetation, given that temperature and precipitation are the parameters that exert the greatest effect (Navarro and Maldonado 2002). The joint action of all these factors is responsible for the high biodiversity and the composition and distribution of the plant communities that characterise the subtropical mountain forests, particularly those growing in the province of Jujuy, where the present study is focused. There are several studies at the regional level on these forests in northwest Argentina (Brown et al. 1985; Mármol 1992; Brown 1995; Brown and Grau 1995; Morales et al. 1995; Prado 1995; Núñez and Mármol, 1997; Giusti et al. 1997; Arturi et al. 1998; Brown and Kappelle 2001; Brown and Malizia 2004; Malizia et al. 2006; Brown et al. 2007, 2009; Entrocassi and Vignale, 2009; Entrocassi and Lambaré 2011; Bulacio and Ayarde 2009; Blundo et al. 2012; Malizia et al. 2012). Studies have been done on diversity, composition and structure, particularly in the locality of Yala in the province of Jujuy (Carranza 2003) and in a sector of the Serranías de Zapla (Cuyckens 2005). The first phytosociological and bioclimatic studies of these forests focused on the basin of the Chijra river (Departamento Manuel Belgrano; Martín 2014) and the site of El Caulario (Departamento Libertador General San Martín; Haagen Entrocassi 2014).

4

1.3

1 Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina). . .

Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina)

This study of the floristic composition, vegetation distribution and bioclimatic characteristics of the subtropical mountain forests in the Serranías de Zapla Multiple Use Ecology Reserve is approached from a phytosociological and bioclimatic standpoint, and is the first work of these characteristics in this territory. The use of the phytosociological approach requires a method that allows the classification, analysis and description of the existing plant communities and the explanation of their spatial distribution and their relationship with the environment. The phytosociological method is one of the most precise and detailed procedures for the analysis of vegetation, as it defines the different types of vegetation existing in a specific territory with a high degree of methodological and conceptual rigour. This procedure is based on the comparative study of phytosociological relevés, and consigns and quantifies the characteristics of each community or plant association, which constitute the basic unit of phytosociology (Braun-Blanquet 1979; Rivas-Martínez 1976, 1994; Rivas-Martínez et al. 1999; Arozena Concepción 2000; Alcaraz Ariza 2013). In these mountain areas, the bioclimate, which is itself conditioned by altitude and relief, is the main factor determining the composition and distribution of the vegetation. Geomorphology and soils are the second determining factor. Together they constitute the most important environmental factors for the control of the vegetation in these environments (Navarro and Maldonado 2002). The relationship between this set of geophysical variables and their direct influence on the vegetation forms the basis of the study of bioclimatology, an ecological science that links the distribution areas of plants, communities and plant formations with specific numerical climate values (temperature and rainfall; Rivas-Martínez 2001, 2008). This study therefore also examines the bioclimatic typology of the province of Jujuy in order to obtain environmental and particularly climatic information at the local scale. It is based on the Worldwide Bioclimatic Classification System and other works developed under this system (Rivas-Martínez 2001, 2008; Rivas-Martínez et al. 1999, 2011; Entrocassi 2011; Entrocassi et al. 2014). This classification is currently being applied successfully in different parts of the world, as it satisfactorily explains the correspondence between climate and vegetation (Sayre et al. 2009, 2013). It is an adjusted predictive model capable of analysing the spatial distribution of the communities and their composition along a particular environmental gradient. This classification and its typology are based on the use of different latitudinal and altitudinal bioclimatic units that have a specific territorial and vegetational expression (macrobioclimates, bioclimates, thermotypes and ombrotypes), and are delimited by a set of bioclimatic indices that weight the intensity of the cold and the amount of moisture supplied by the rainfall, both limiting factors for many plants and plant communities. These indices are calculated according to a variety of climatic parameters, mainly temperature and precipitation. This classification enables a particular locality to be characterised from a bioclimatic point of view, by assigning it a macrobioclimate

1.3 Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina)

5

and a bioclimate, and different bioclimatic temperature and moisture belts based on the latitude and altitude, and which in turn correspond to different types of vegetation. The environmental characteristics of the study area are determined by the general environmental framework where it is located: the province of Jujuy. This province is crossed through its middle part by the Tropic of Capricorn; it has an area of 53,219 km2 and is located in the northwest part of Argentina, between 21
460 -24
360 S and 64
100 -67
110 W. It borders Bolivia towards the north, Chile to the west, and the province of Salta to the south and east. Most of the province is dominated by broken relief with different geomorphologies, climate, hydrographic systems and plant formations (Nicola et al. 2012). The environmental complexity of the province of Jujuy corresponds to the four geological units that cross the territory in a north-south direction: la Puna, Cordillera Oriental, the sub-Andean ranges and the Santa Bárbara mountain system (Ramos 1999). In general terms the whole province can be said to lie on the western edge of the Brasilia massif, formed by igneous, crystalline and metamorphic rocks that have withstood strong endogenous movements. The mountain systems that became established here in pre-Cambrian times were subsequently eroded and transformed into peneplains, such as the La Puna block. The Andean orogeny in the Tertiary Period raised the western edge of the La Puna block to a great height and subjected it to dislocations that formed interior basins and mountain ranges, where blocks of rock and sedimentary mantles on the eastern border were fractured and overelevated to conform the current Cordillera Oriental. Meanwhile the sediments located towards the east of the province in the Chacoan-Pampean fill trench, containing the so-called “petroliferous formation”, among others, also suffered the pressure of Andean orogeny, producing low irregular folds that gave rise to the sub-Andean ranges. All these systems are found not only in the province of Jujuy, but continue throughout other Argentinian provinces and in neighbouring countries such as the republics of Chile and Bolivia (Nicola et al. 2012). The various large-scale atmospheric processes and the influence of the relief are the factors determining the climatic characteristics in the province of Jujuy: it has extensive very cold dry raised areas such as La Puna and the Cordillera Oriental, other warm or temperate and humid areas such as the sub-Andean mountain ranges and valleys, through to very warm and dry lower areas such as those located near the Chacoan plain, all reflecting a wide variability in terms of rainfall and temperature. The rainfall distribution in the province responds to a summer regime. 80% of the precipitation is concentrated in the warm season (from November to March) as a result of the wind regime, which is affected by the interaction of the Atlantic and Pacific subtropical anticyclonic centres and by the polar anticyclone. In the summer months a low pressure centre is created over the Chacoan plain, allowing the arrival of air masses carrying moisture from the Atlantic and producing abundant rainfall. When these air masses meet the mountain ranges they under the well-known process of ascent, adiabatic cooling and condensation, which concludes with the production of abundant “orographic precipitations” on the sub-Andean ranges and in the easternmost sectors of the Cordillera Oriental. The precipitations generally increase

6

1 Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina). . .

with height until a certain level located at between 900–2500 m asl, after which they decrease, regardless of the continued increase in altitude. In winter, the temperature drops and creates a high pressure centre, the thrust of the moist winds from the Atlantic abates, and as a result the rains become very scarce. This produces a prevalence of the fine weather conditions and the clear dry days that are so characteristic of this province in winter. The temperature regime in the province is mainly determined by the relief, latitude and altitude. The highest annual temperatures occur in the low valleys and decline as the altitude rises from the sub-Andean ranges, across the Cordillera Oriental and as far as La Puna. Here the climate conditions are extremely harsh, characterised by marked variations in temperature and daily frosts, with the start of the lower limit of the permasnow at 5500 m asl until 6200 msn on the highest peak in the province (Cerro Chañi; Buitrago 2000; Entrocassi et al. 2014). These environmental features determine the environmental and biological diversity of Jujuy, and particularly the development of different ecosystems and plant formations such as the subtropical mountain forests described in this study. However, traditional human activity has been developed in parallel in the low warm valleys in the primitive forests, primarily for the purposes of timber production, which in some sectors has been replaced by the cultivation of sugar cane, citrus fruits and vegetables. In the somewhat higher and temperate-warm valleys there is a predominance of tobacco cultivation, and this is also the site of the iron deposits in the Serranía de Zapla. The inter-Andean pre-Punean valleys in the Cordillera Oriental (Quebrada de Humahuaca) are the site of small-scale agriculture, particularly microthermal Andean cultivation, and are subject to intense tourist activity. Finally, La Puna is the sector least suited to human settlement, although it contains important mining resources (Nicola et al. 2012). The Serranías de Zapla Multiple Use Ecology Reserve is a protected territory located in the south of Jujuy with an area of 37,139 ha, created in 2003 by the city of Palpalá, due to the fact that this is a site of scientific, economic and cultural interest. Its main function is to preserve the balance between its natural resources and the human presence through the sustainable use of its environments, thus amalgamating the productive activity that has historically existed in the area with the conservation of its natural environments and its native resources. The designation of this conservation category enables the development of experiences in harmonious coexistence between production and conservation activities in the reserve, all under the control and technical auditing of the municipality, which is responsible for regulating and guaranteeing the development and conservation of its productive potential, its wildlife and its landscape. It should be noted that one sector of the reserve is home to the 9 de Octubre mine (at 1450 m asl), where the rich iron deposits in the Sierra de Zapla were previously exploited for the purpose of promoting the national iron and steel industry, constituting a mainstay of the country’s economy. The mine began operating in 1943 and remained active until its definitive closure in 1997. At its height, it had a residential civic centre with the basic services and spaces for social, educational, recreational and cultural development, and a stable population of approximately 600 inhabitants. This historic circumstance, together with the fact that the reserve comprises municipal and private lands where some productive

1.4 Las Yungas Biosphere Reserve (RBYungas) and the Jujuy Model Forest

7

activities are undertaken today, makes this an area with a certain degree of intervention and transformation, conforming a mosaic of sectors affected by human activity and other areas with very little human impact. In spite of this, it still continues to be a very diverse environment where the geophysical factors that typically act in these mountain regions favour the permanence of a significant extension of subtropical montane forests. Some of these formations are fairly well conserved due to the fact that they are located in sites that are difficult to access, and in others that are firmly on a course to recovery, covered for a large part of the year by mists that settle on the slopes, with exuberant flora and diverse plant communities that warrant a detailed study.

1.4

Las Yungas Biosphere Reserve (RBYungas) and the Jujuy Model Forest

It should be noted that 7730 ha (21%) of the Serranías de Zapla Multiple Use Ecology Reserve are also included within the large Las Yungas biosphere reserve (RBYungas), a transprovincial protected area that covers the provinces of Jujuy and Salta, and was created within the framework of the UNESCO’s Man and Biosphere Programme in 2002, with several goals: to conserve and protect the subtropical mountain ecosystems in northwest Argentina, as this is an area of outstanding biodiversity in Argentina; to prevent and resolve problems such as the fragmentation of the natural landscape, the impoverishment and loss of native forests, the erosion of slopes, flooding and pollution; to promote integration between protected natural areas and those that have been transformed by human activity through territorial ordination, for the purpose of encouraging sustainable development and the improved management of natural resources; and to provide support for education, training and research projects. The Las Yungas Biosphere Reserve (RBYungas) is located in the northwest of the Republic of Argentina; it covers an area of approximately 1,350,000 ha and is the second largest reserve in Argentina. There are four core areas in the RBYungas: the Baritú National Park, the Pintascayo Provincial Park, the Calilegua National Park and the Potrero de Yala Provincial Park. It also includes two protected areas that have been created since its official designation, and which were therefore originally not included as core areas, although they currently function as such. These are the El Nogalar de Los Toldos National Reserve and the Serranías de Zapla Multiple Use Ecology Reserve, the subject of the present study. The Las Yungas Biosphere Reserve, along with the Tariquía de Tarija National Flora and Fauna Reserve (Bolivia) form part of the biological corridor known as the “Las Yungas Corridor”, whose aim is to protect the ecoregion of the Las Yungas forest, and thus safeguard the hotspots in the native forest that are so important for maintaining the connectivity between the different habitats, most of which are in a situation of vulnerability due to the increasing trend towards deforestation at the expense of the native forest. It should be noted that in Argentina this corridor extends

8

1 Vegetation of Las Yungas (Serranías de Zapla, Jujuy, Argentina). . .

over areas that contain over 300,000 ha of soil suitable for farming and which are still under forest cover. In 1973 it was reported that there was 5.5% of agricultural use at the cost of the native forest, and this area doubled to 11% in 2000 (Burkart 2006; Anon 2010a). RBYungas is located within the territory designated as the “Jujuy Model Forest”, a part of the National Model Forest Programme, aimed at promoting the recovery and highlighting the value of traditional skills and culture, improving the quality of life in the communities and encouraging participative management in the use of natural resources, while offering a space to carry out research and education tasks. This reserve is included in Category II (Yellow) of the Territorial Ordination Plan of the Native Forests in the Province of Jujuy, which promotes the preservation of native forest resources and divides the province into zones with different conservation values. It is what is known as an “area of forest maintenance”, administered by conservation and/or sustainable development plans that also allow the transformation of limited sectors in duly justified situations. The present study is an important addition to the knowledge of the flora, vegetation, phytosociology and bioclimate of the subtropical mountain forests of Las Yungas at the provincial and regional scale, and in the general setting of South America. The data obtained in this study also represent a valuable contribution to the conservation of these forests, given that they offer an assessment of the current state of the vegetation in terms of its attributes and potential, and can be used as an argument for incorporating the entire territory within the Las Yungas Biosphere Reserve.

Chapter 2

General Features of Serranías de Zapla Multiple Use Ecology Reserve

Serranías de Zapla Multiple Use Ecology Reserve is located in the Department of Palpalá, in Jujuy province (Fig. 2.1). It is delimited by the geographic coordinates 24
090 –24
21’S and 65
10 –65
14’W (Fig. 2.2), and has an area of 37,139 hectares, with altitudes ranging from 834 to 2183 m asl. Within this altitudinal range, the forest extends along an altitudinal gradient of 600 m at between 1015 and 1620 m asl, and this is the area chosen for our study.

2.1

Geomorphology, Geology, Hydrology and Soils

The environmental features of the Serranías de Zapla Multiple Use Ecology Reserve correspond to the general context of the sub-Andean mountain ranges, which form the southern end of the chain of the same name that descends from Peru, crosses Bolivia and penetrates into Argentina as a narrow belt running north to south and covering part of Jujuy and Salta provinces. This belt is approximately 100 km wide and 500 km long, increasing in altitude from east to west. The sub-Andean mountain ranges limit to the west with the Cordillera Oriental range, to the east with the Chacoan plain, and to the south with the Pampean ranges. They constitute the first orographic barrier that condenses the moist air masses from the Atlantic, and condition the particular climate conditions responsible for the diverse drainage network that characterises the region, and the dense tree coverage growing on their eastern slopes. These represent approximately 23% of the total area of Jujuy province (Anon 2010b; Santamans and Franco 2010). Specifically, the orographic layout of the Reserve is structured in belts of sierras and mountain ranges, and oriented uniformly in a sub-southern direction. Its relief is predominantly concordant in nature, with anticlinal mountains and synclinal valleys; between the mountain ranges there are small valleys and intermontane gorges with a network of rivers and streams that form part of the sub-basin of the San Francisco river, a tributary of the upper basin of the Bermejo river. © Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_2

9

Fig. 2.1 Geographic location of the study area (yellow circle) in Jujuy province (Argentina). Source: Google Earth and National Geographic Institute of Argentina (NGI)

10 2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

2.1 Geomorphology, Geology, Hydrology and Soils

11

Fig. 2.2 Satellite image of the Serranías de Zapla Multiple Use Ecology Reserve in the provincial context (limits in yellow line). Source: Google Earth Table 2.1 Soil associations and soil types present in the study area, following the FAO and USDA soil classification Association Abra de Caña-Río Jordán Sevenguial-Río Jordán Palpalá

Taxonomic classification FAO Lithosols-Eutric Regosols Calcaric Phaeozems-Eutric Regosols Haplic Phaeozems

USDA (subgroup) Lithic Ustorthents-Udic Ustorthents Typic Hapludolls-Udic Ustorthents Aquertic Hapludolls

Source: INTA-UNSa (2009)

From the geological point of view, the study area is located on ancient bedrock dating from the lower Palaeozoic (Ordovician) era which appears on the surface in the Sierras de Zapla as an anticlinal structure containing a rich iron deposit. This base supports younger rocks such as Mesozoic and Cenozoic sedimentites that form a substantial sedimentary covering (Paoli et al. 2009; Anon 2010b). The main basins belong to the Zapla and Las Capillas rivers. The hydrology is typical of mountain regions; most of the water courses have very marked seasonal variations, with their maximum flows recorded in the period of heavy rainfall (between December and March), and the minimums in the period of maximum drought (between September and October). The intense flooding produces landslides and mass removals (Anon 2010b). According to the FAO’s taxonomic classification, the following main soil types are recognised within these associations: Lithosols, Eutric Regosols, Calacaric

12

2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

Phaeozems and Haplic Phaeozems (Table 2.1; Fig. 2.3a) (Nadir and Chafatinos 2008; Paoli et al. 2009; Rosas and Chayle 2011). The Lithosols are distributed from north to south in narrow bands on the crests and flanks of the Sierras de Zapla, where the relief is steep to very steep, with abrupt slopes of over 40%. These are soils with incipient development and an A-R profile, and a sequence of A, C, IIC2, R or A, C, R horizons. The dominant bedrocks are sedimentary (sandstone, arcillite, limonite and conglomerate) or metamorphic (quartzite, schist and slate), while igneous rocks occupy a reduced area. These soils are characterised by fast and excessive drainage and runoff, and moderate to severe erosion determined by the water agents. The surface horizon corresponds to an ochric epipedon. Eutric Regosols are found in the foothills and intermontane valleys in the Reserve. These are poorly developed soils with an A, AC, R profile with a predominance of medium coarse textures. They are neutral to slightly acid and moderately well-drained, with slopes of 6–13% and moderate erosion.

Fig. 2.3 Main soil types (a) and associations (b) in Serranías de Zapla Multiple Use Ecology Reserve. Soil types: 1 Lithosols-Eutric Regosols, 2 Calcaric Phaeozems- Eutric Regosols, 3 Haplic Phaeozems. Soil associations: Sev-Rj Sevenguial-Río Jordán association, Ac-Rj Abra de Caña-Río Jordán association, Pp Palpalá association. Black line: limit of the Serranías de Zapla Multiple Use Ecology Reserve. Source: INTA-UNSa (2009)

2.1 Geomorphology, Geology, Hydrology and Soils

13

Fig. 2.3 (continued)

Calcaric Phaeozems are present in the terminal part of the foothills of the mountain ranges and on terraced levels, adjoining the mountain ranges, or on their summits. These are soils with a well-developed profile, whose sequence of horizons is generally A1, B2t, B3, C. They are well structured, with fine to medium textures and slightly acid. They have a mollic epipedon and abundant organic matter. Their distinguishing feature is their accumulations of calcium carbonate in the upper 20 and 50 cm. Finally, the Haplic Phaeozems are uniformly distributed within the same environments as indicated for Calcaric Phaeozems. They have the same sequence of horizons, physical and chemical properties, but no accumulation of carbonates. The soil series in the Reserve are distributed longitudinally from north to south; in the intermontane valleys and gorges the soils are more developed than at overelevated levels. There are three soil associations, namely: the Abra de Caña-Río Jordán association at the northern end; the Sevenguial-Río Jordán association, which

14

2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

occupies the greatest extension in the study area; and the Palpalá association, which extends along a narrow strip at the southern end of the Reserve (Table 2.1; Fig. 2.3b).

2.2

Vegetation and Biogeography

As mentioned in the introduction, the dominant plant formation in the study area is the subtropical montane forest or Yungas. According to Navarro and Maldonado (2002), these forests correspond to the designation Pluviseasonal BolivianoTucuman Sub-Andean Montane Vegetation, characterised by sub-humid and humid forests growing on east-facing slopes in the Cordillera Oriental in the Andes, from central Bolivia (Cochabamba) through to the La Rioja mountain range in Argentina. According to Cabrera and Willink (1980) and Cabrera (1994) there are three altitudinal vegetation belts in the Yungas in northwest Argentina that in turn delimit three phytogeographical districts: the Transitional Rainforest or Foothills, which occupy the plains and low-lying mountains adjacent to the first slopes of the mountain range between 350 and 550 m asl; the classic Montane Rainforest between 550 and 1600 m asl; and the Montane Forest, which grows on inclines and high ridges at between 1600 and 2300 (2500) m asl. This is the coldest belt, and has frequent precipitation in the form of snow in winter. Other authors fix the limits at between 400 and 900 m asl for the first, between 700 and 1500 m asl for the second, and between 1500 and 3000 m asl for the last (Brown et al. 2002; Malizia et al. 2012). As is to be expected, these limits are approximate, given that they are determined by local environmental factors that affect the distribution of the different vegetation belts. The altitudinal vegetation belts containing the forests in the study area (1015–1600 m asl) correspond to Montane Rainforest and Montane Forest, according to Cabrera (1994). Structurally, these are dense, semi-deciduous and seasonally perennial broadleaved laurel micro- and mesoforests with an average canopy height of 10–20 m and an arboreal, shrubby and herbaceous understory of different heights or levels; creepers, climbers and epiphytes are fairly abundant, particularly in the highest sectors of the forest (1500–1600 m asl), and in sectors where they grow on south- and east-facing slopes where the moisture input is higher due to the effect of orographic precipitation and the frequent mist banks that settle on these areas, mainly in summer and early autumn. The species present in the forests in the study area include particularly some that are considered characteristic and/or endemic species of the sub-Andean and montane vegetation in the Boliviano-Tucuman province, according to Navarro and Maldonado (2002). For example, in the forest canopy there is a presence of Cinnamomum porphyrium (laurel), Erythrina falcata (ceibo), Juglans australis (nogal criollo), Myroxylon peruiferum (quina), Parapiptadenia excelsa (cebil blanco) and Tipuana tipu (tipa blanca), among others. The characteristic and/or endemic species in the lowest tree layer are Escallonia millegrana, Gleditsia amorphoides (espina corona),

2.2 Vegetation and Biogeography

15

Ilex argentina (palo yerba), Jacaranda mimosifolia (jacarandá), Myrcianthes pseudomato (mato arrayán), Pisonia zapallo (zapallo caspi), Podocarpus parlatorei (pino del cerro), Prunus tucumanensis (palo luz), Randia micrantha, Schinus gracilipes and Xylosma pubescens (coronillo); whereas those in the shrub layer are Barnadesia odorata, Budleja iresinoides, Croton saltensis and Justicia kuntzei, while Cortaderia hieronymi is notable in the herb layer. The Montane Rainforest layer has the greatest plant diversity. Above it, in the Montane Forest, the richness of species, genera and families declines with the increase in altitude, following the general pattern observed for the altitudinal gradients in the Neotropic. This reveals an impoverishment in species richness with increasing altitude, a higher rate of species exchange in the different altitudinal layers, and significant changes in the forest structure in terms of decreasing tree height or greater basal area and density at higher elevations (Gentry 1988; Lieberman et al. 1996; Vásquez and Givnish 1998). The same pattern has also been observed in studies on the tree stratum in the Yungas in northwest Argentina, where the altitudinal interval between 600 (700 m asl) and 1600 (1700) m asl has the greatest species richness. Beyond 1700 m asl this richness declines significantly, particularly in the southernmost sector of the Yungas (Grau et al. 1995; Brown et al. 2001; Brown and Malizia 2004). For example, in the Yungas in the upper basin of the Bermejo river (northwest Argentina and southern Bolivia), the number of tree species reported in an altitudinal range between 600 and 1600 m asl (with DAP 10 cm) was 78 species in the basal part of the Montane Rainforest (600 m asl), 95 species in the intermediate rainforest (1100 m asl) and 63 species at the upper limit (1600 m asl). This is evidence that the diversity of tree species in this sector of the Yungas tends to decrease with altitude, but presents a peak of richness at intermediate levels, at approximately 1100 m asl (Malizia et al. 2006). Gentry (1988, 1995) also noted the existence of peaks of maximum species richness between 1000 and 1500 m asl for Andean forests, suggesting that this is linked to historic factors that produce the meeting or convergence of species of different biogeographical origins, causing a superimposition of species at intermediate altitudes. According to Malizia et al. (2006), the confluence of species with different biogeographical origins could explain the richness found at this intermediate altitudinal level in the Montane Rainforest in northwest Argentina and southern Bolivia (between 1000 and 1500 m asl), as this is the point of convergence of floristic elements belonging to holartic families such as Cedrela angustifolia (Meliaceae), Juglans australis (Juglandaceae), and species with a Gondwanan (Roupala meisneri) and tropical origin (Clethra scabra, Cinnamomun porphyrium, Styrax subargenteus, Miconia molybdea, Trichilia hieronymi). According to this author, this diversity of origins could be a response to the extraordinary complex of environmental gradients characterising the Andes. Particularly in regard to the study area, the same pattern of decreasing species richness with increasing altitude was observed in a previous study carried out in a sector of the Sierras de Zapla (Cuyckens 2005), where 31 tree species were reported in the intermediate sector of the Montane Rainforest (at 1150 m asl), 20 species in the highest sector (1600 m asl) and 15 species in the Montane Forest.

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2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

From the biogeographic point of view, the forests in the study belong to the Boliviano-Tucuman province, which falls within the Tropical South-Andean Region (Neotropical Kingdom) (Rivas-Martínez et al. 2011). This biogeographic province extends from central Bolivia, at 18
S (in Cochabamba), as far as the La Rioja mountain range in Argentina at 29
S, occupying the eastern slopes of the Andean ranges. Within this biogeographic province there are several ecological belts containing numerous plant formations, and including particularly the subtropical montane forests or Yungas, which can be assimilated to the “Pluviseasonal BolivianoTucuman Sub-Andean Montane Vegetation” according to Navarro and Maldonado (2002). It should be noted that there are currently very few studies on the Jujuy province to support the delimitation of lower biogeographic units within the Boliviano-Tucuman province (for example, at the sector and district level), only the “Biogeographical Map of South America” by Rivas-Martínez and Navarro (1994) (in Navarro and Maldonado 2002) and its updated version by Rivas-Martínez et al. (2011). According to the typologies in this work, the study area can be located within the South American biogeographic context at the level of kingdom, region and province (Fig. 2.4). According to Cabrera and Willink (1980) and Cabrera (1994), the forests in the study area belong to the Montane Rainforest and Montane Forest districts in the phytogeographical province of the Yungas (Amazonian Domain, Neotropical Region). (Fig. 2.5), whose equivalents are the Yungas province (Morrone 2001), the classic “Tucumano-Bolivian Rainforest” (Hauman 1931; Castellanos y Pérez Moreau 1941; Parodi 1945; Tortorelli 1956) or the “Tucumano-Oranensian Rainforest” (Ragonese 1967), or earlier, the “Subtropical Formation” (Holmberg 1898) or “Sub-tropical mountain forests of the northwest” (Kühn 1930), among other denominations. There is a biogeographic relation between the vegetation of the subtropical mountain forests and the vegetation of the Cerrado and Paranaensian biogeographic provinces, as they share some genera such as Allophyllus, Blepharocalyx, Jacarandá, Ocotea, Nectandra, Handrohantus, Cedrela, etc.; while others are exclusive to the Yungas, such as Tipuana, Cascaronia, Phoebe, Myroxylon, Amburana, Juglans, Alnus, Phoebe, Cnicothamnus, etc. (Cabrera 1994; Navarro and Maldonado 2002). In the subtropical montane forests there is a high number of neotropical (81) and pantropical (29) genera, and a few genera with an Andean (3), holarctic (7) and southern (3) origin. No endemics have been detected at the level of genera, although there is a high percentage of endemics at the species level (Quiroga 2010); this could be linked to the climate changes occurring during the Quaternary era (Pleistocene) in the Amazonian subregion, which led to a reduction and fragmentation of the forests, the isolation of several species, and the establishment of forest patches or biodiversity refugia. This is the case of the subtropical montane forests of the “upper basin of the Bermejo river” in northwest Argentina and Bolivia (Morales et al. 1995; Brown et al. 2001; Morrone 2001; Quiroga and Premoli 2007). Particularly in the Montane Rainforest belt there is a presence of dominant species with a tropical origin, for which this region constitutes the southernmost

2.2 Vegetation and Biogeography

17

Fig. 2.4 Biogeographic map of South America (Rivas-Martínez and Navarro 1998; Navarro and Maldonado 2002; Rivas-Martínez et al. 2011). The units in the map are: NEOTROPICALAUSTROAMERICAN Kingdom; NEOTROPICAL Subkingdom; CARIBBEAN-

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2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

Fig. 2.5 Phytogeographic territories in Jujuy province (Cabrera 1994), adapted by Burgos (2013). Approximate location of the study area within the Montane Rainforest district, in the Yungas phytogeographical province (circle). Black and green lines indicate provincial and department limits; light blue, Altoandina quechoa (Quechuan upland); light green, Bosque montano (Montane forest); dark brown, Chaco seco (dry Chaco); light brown, Chaco serrano (sierran Chaco); orange, Prepuna; very light brown, Puna; dark green, Selva montana (montane rain forest); light green, Selva de transición (transitional rain forest)

geographic limit of their distribution (Brown and Malizia 2007). According to Quiroga (2010), most of the genera present in the study area come from elements with a neotropical origin, namely: Blepharocalyx, Schinus, Ilex, Cnicothamnus, Handroanthus, Jacaranda, Tecoma, Ceiba, SCCAellium, Carica, Cnidoscolus,

Fig. 2.4 (continued) NEOGRANADIAN Superegion; 9. Caribbean-Mesoamerican Region; 10. Neogranadian Region; AMAZONIAN-GUYANAN Superegion; 11. Guyanan-Orinoouian Region; 12. Amazonian Region; 13. Brazilian-Paranense Region; 14. Chacoan Region; TROPICAL SOUTH ANDEAN Superegion; 15. Tropical South Andean Region; 15.1. Desertic PeruvianEcuadorean Province; 15.2. Mesophytic Punenian Province; 15.3. Xerophytic Punenian Province; 15.4. Bolivian-Tucuman Province; 15.5. Yungenian Province; 16. Hyperdesertic Tropical Pacific Region; AUSTROAMERICAN Subkingdom; 17. Pampean Region; 18. Middle Chilean-Patagonian Region; 19. Valdivean-Magellanian Region; CIRCUMANTARCTIC Subkingdom; 20. Insular Antarctic Region; 21. Continental Antarctic Region. Approximate location of the study area within the Bolivian-Tucuman province (red circle)

2.3 Management: Situation of Serranías de Zapla in the Context of. . .

19

Saccellium, Stillingia, Anadenanthera, Enterolobium, Mimosa, Parapiptadenia, Myroxylon, Tipuana, Cedrela, Myrsine, Myrcianthes, Bougainvillea, Condalia, Coutarea, Scutia, Coutarea, Pisonia, Vassobia, Zanthoxylum and Duranta. The pantropical genera present are: Eupatorium, Trema, Sapium, Sebastiania, Senna, Acacia, Prosopis, Erythrina, Cinnamomum, Zanthoxylum, Xylosma, Allophylus and Chrysophyllum, whereas Celtis, Juglans, Prunus and Salix are holartic genera and Podocarpus is southern.

2.3

Management: Situation of Serranías de Zapla in the Context of Protected Areas and the Territorial Ordination Plan of Jujuy Province

Most of the area of the Serranías de Zapla Multiple Use Ecology Reserve is private land, and in a lesser proportion, municipal land that previously belonged to the Mina 9 de Octubre, from which iron was extracted then processed in the Altos Hornos Zapla blast furnaces (in the city of Palpalá). When the mine closed in 1997 the premises became the property of the city’s municipal authority. Most of the Reserve has coverage of native forest (Yungas); however there are areas of grazing and forest plantations. Approximately 11,000 hectares of the Reserve were transformed due to forestry activity with exotic species, mainly Eucalyptus grandis, E. camaldulensis, E. viminalis, E. tereticornis, Pinus elliottii and P. taeda, which are currently used for sawn wood, ground wood and as energy. Specifically, the different species of eucalyptus were introduced in 1948 by the state-run Altos Hornos Zapla to obtain charcoal to feed their iron and steel plant in the city of Palpalá. There are small rural settlements associated to forestry activity and dedicated to subsistence livestock and family farming. There are also temporary grazing lands in the high areas of the Reserve above the tree line, where there are natural mist grasslands (Montane Grassland). The rest of the territory in the Reserve is covered by the Yungas. One part of these forests is in a good state of conservation, particularly in areas that are difficult to access, whereas other sectors are in a clear process of regeneration and recovery as a result of having been degraded or fragmented by extraction activities in the past. This is particularly the case of valuable lumber species for forestry such as Cedrela angustifolia (cedro coya), Juglans australis (nogal criollo) and Myroxylon peruiferum (quina), in addition to other species used for fuel, posts, building roofs and others, such as Anadenabnthera colubrina var. cebil (cebil colorado), Parapiptadenia excelsa (cebil blanco), Prosopis alba (algarrobo blanco), Allophyllus edulis (chal-chal), Alnus acuminata (aliso del cerro) and more. The intense past exploitation of the native forests has declined significantly in recent years, aided by the creation of the Reserve in 2003 and the enactment of laws for territorial ordination and protection of the native forest. The native forests in the Reserve therefore coexist with areas in which the human presence has had varying degrees of intensity, conforming a mosaic of characteristic

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2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

landscapes with some well-conserved and practically unaltered sectors, along with others that have been transformed by human impact. Tourist activities take place in the areas under municipal jurisdiction to exploit the imposing scenic landscape and the infrastructure of the old residential complex of the 9 de Octubre mine, including excursions to the pit and adventure sports in the surrounding rainforest. The northern end of the Serranías de Zapla Multiple Use Ecology Reserve is within the buffer zone of the Las Yungas Biosphere Reserve (Fig. 2.6) and forms

Fig. 2.6 Inclusion of the extreme north of the Serranías de Zapla Multiple Use Ecology Reserve (in yellow) within the Las Yungas Biosphere Reserve (in green). Source: SIGA Proyungas

2.3 Management: Situation of Serranías de Zapla in the Context of. . .

21

part of a biological corridor that connects several protected national areas, such as the Potrero de Yala Provincial Park, Calilegua National Park, Laguna Pintascayo Provincial Park, El Nogalar de Los Toldos National Reserve and the Baritú National Park, and foreign areas such as the Tariquía Flora and Fauna Reserve in Tarija, Bolivia. As mentioned before, the Las Yungas Biosphere Reserve is a protected area created in 2002 within the framework of the UNESCO’s Man and the Biosphere Programme. Its purpose is to conserve and protect the subtropical montane ecosystems in northwest Argentina, one of the most biodiverse regions in the country, to implement strategies that combine conservation with sustainable development, and to promote activities of research, training and education. The Reserve is managed by the governments of Jujuy and Salta provinces and by the National Parks Administration, which depends on the office of the presidency in Argentina. The study area is also included in Category II of the Territorial Ordination Plan of Native Forests in Jujuy province (Decree 2187/08, Provincial Act 5676/2011). This legislation promotes centres for the conservation of native forest resources in their provincial territory, and establishes the zoning of the province based on a series of criteria according to National Act 26.331 on Minimum Budgets for the Environmental Protection of Native Forests (Fig. 2.7). According to the legal framework of this territorial ordination plan, Category II includes territories with a medium to high

Fig. 2.7 Situation of the Serranías de Zapla Multiple Use Ecology Reserve (in blue) within the legal framework of the Territorial Ordination Plan for the Native Forests in Jujuy province. Source: SIGA Proyungas

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2 General Features of Serranías de Zapla Multiple Use Ecology Reserve

conservation value which have a slope of over 5%. This category includes so-called ‘forest maintenance areas’, where the decisions and activities are governed by conservation and/or sustainable management plans that authorise the transformation of limited sectors of the forest when the situation warrants, and after an environmental impact assessment (such as for example the location of infrastructure works and traditional and non-traditional productive uses). This category may also include private protection areas in which the protection and/or the sustainable development of their natural resources is deemed necessary. It also part belongs to the territory known as the Jujuy Model Forest, a part of the National Model Forest Programme which affects 433,500 ha of native forest and one of whose objectives is to administer the use of natural resources in a participative way. This management category was established through an agreement between the National Department of the Environment and Sustainable Development, and the Department of Environmental Management of the Government of Jujuy province.

Chapter 3

Bioclimatology

The relationship between climate and plants was noted as long ago as the third century BC by Theophrastus (Hort 1916), who highlighted the importance of climate in plant distribution through direct and experimental observation. The ideas of this thinker were developed no further until the late eighteenth and early nineteenth century—aided by the invention of the thermometer and the barometer in the seventeenth century—in the works of de Willdenow (1792), von Humboldt (1807), Wahlenberg (1811) and Grisebach (1838), when it became evident that climate was the main factor driving the distribution of plants and the communities they form, giving rise to a new science called Bioclimatology. Bioclimatology studies the relationship between climate and the biological environment, and was extensively developed in the twentieth century through the close interrelation between climate and vegetation; that is, phytoclimatology. Today there is an extensive bibliography on the subject, including several reviews (Emberger 1930; Thornthwaite 1931, 1948; Philippis 1937; Bagnouls and Gaussen 1954; Daget 1977a; Tuhkanen 1980). The study of the climate in a particular area may be a useful tool to obtain information on the type of vegetation contained within it, although naturally certain factors such as the typical floristic composition of the territory and its origins, topography and soil type, also have a significant influence on vegetation. The climate in a territory is a complex whole formed of numerous elements such as rainfall, temperature, radiation, moisture and wind regime, among others, whose combination, superimposed on a number of other environmental factors (topography, soils), plays an important role in the development of plants, and consequently on their distribution. These are combined in the form of bioclimatic indices and used in addition to the gross data mentioned above—insufficient in the opinion of certain authors (de Martonne 1926; Tuhkanen 1980)—to quantify and determine this influence on a large scale. This formulation led to significant development in global climate classifications which aimed to integrate the whole variability of climate and vegetation present on Earth. Due to the large volume of information found in the

© Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_3

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literature, we have attempted to summarise the various trends in the works published in the twentieth century or subsequently. The first studies of vegetation distribution in relation to climate took place in Europe; however, we will only refer in this section to works published in the present century. Due to their scientific training, European researchers used a similar initial starting point in their methodology, and in most cases it was botanists or phytogeographers who explained the influence exercised by climate on vegetation. These studies were based on the knowledge of the vegetation types and their distribution; the use of elementary climate variables such as precipitation and temperature and the indices that combine them served to explain this distribution. One of the first approaches to the study of bioclimatology was by the Russian botanist Köppen in 1900, with the publication of a climate classification system that was subsequently developed and improved in successive years (1918, 1930–1936). In this work the author sought to establish an objective numerical classification for use in phytogeography based on different climate variables for mean and limit temperature in climate areas that coincided as closely as possible with the major vegetation zones. To describe this phenomenon he coined terms such as poplar climate, beech climate, olive climate and so on, in one of his first approaches. He subsequently used the mean temperatures of the warmest and coldest months and summer and winter rainfall and its duration as discriminant climate parameters, and distinguished five types of climate, of which four are based on temperature parameters (tropical, temperate, continental and polar), and one on moisture parameters (dry climate). The first subdivision of these climate types was made according to precipitation factors, while another second-order subdivision was based on temperature factors. The Mediterranean climate, with its acute summer drought, was included in temperate and not dry climates, which were reserved for areas on Earth with constant drought throughout the year. The last years of French colonialism in Northern Africa and the development of very close diplomatic ties between France and the Arab countries in the western Mediterranean—mainly Algeria and Morocco—presented an opportunity to conduct important phytoclimatic studies. These were used as a basis for formulating a variety of climate indices and classifications that were subsequently widely applied throughout the Mediterranean world, along with several global classifications, although they had far less repercussion outside Mediterranean countries than the Russian or German systems. The first works focused on the study of precipitation, either as a factor exerting a major influence on continentality (Angot 1906, 1918), or else as part of a study of the characteristics of global aridity (de Martonne 1926). Martonne introduced the P/T ratio to demarcate a dry and a humid month (de Martonne 1942). Gaussen (1921) used the climate studies on rainfall conducted by Angot (1918) to study the relation between summer precipitation and Mediterranean vegetation distribution, which has adapted to withstand long periods of drought. Years later Gaussen proposed the xerothermic index, or the number of days in the driest season of the year (summer) with no recorded precipitation (Gaussen 1949; Gaussen and

3 Bioclimatology

25

Bagnouls 1952), derived from the previous studies on summer aridity. This culminated in a thorough revision of the concept of the dry season (Bagnouls and Gaussen 1954), determining its geographic distribution (limits) in addition to its duration (expressed by the P/T ratio), and further developing the pluviothermal diagrams previously adopted by Gaussen (1949). Gaussen was a highly prolific author who conducted bioclimatic studies both all around his own country (1935) and in EuroSiberian areas (the Paris basin 1936; Basque Country 1941); and even in other disciplines such as his study of the relation between climate and soils (1941). Finally, Bagnouls and Gaussen (1957) proposed a global climate classification based on annual precipitation and temperature regimes, considering the monthly mean of these variables and the duration of the cold and warm, dry and humid periods. Another French botanist who had a significant impact on climate studies was Emberger, whose first works focused on the study of the Mediterranean climate (Emberger 1930, 1932). He considered that the most important climate factors for the development of vegetation were precipitation, temperature and evaporation. Based on these parameters he formulated an ombrothermic index from which he derived a pluviothermic climate diagram of Mediterranean bioclimatic belts which has been widely used (see Tuhkanen 1980; Nahal 1981; Guara et al. 1986; Defaut 1989). Emberger subsequently published a series of works (1938, 1942) exploring a variety of general climate notions that would culminate (Emberger 1954) in a biogeographical climate classification on a global scale, although without abandoning the study of the Mediterranean climate (Emberger 1943, 1959, 1971). In parallel with the publication of the works of Martonne, Emberger, Gaussen and others, further equally important works appeared such as the review of climate indices and classifications by Philippis in Italy (1937), and the works of Giacobbe (1938, 1958, 1959, 1967) in which the concept of summer aridity was thoroughly revised and new aridity indices formulated. These works, in addition to addressing the delimitation of the duration of the dry season—questioning the T/P ratio as an indicator of its limits—, proposed the limits of the Mediterranean region, assessing the degree of mediterraneaneity of a territory according to: (1) type of pluviothermic regime, (2) number of days of summer precipitation, (3) pluviothermic percentage range between the rainiest and the driest season, and (4) degree of ombric irregularity in summer. From the 1960s onwards the scientific output of the French school declined significantly. Notable among its works were that of Vernet and Ph (1966) on summer drought, with the combination of the relative seasonal pluviometric range and a summer thermopluviometric quotient along a continentality or oceanity index which the authors themselves also defined as an index of mediterraneity/ atlanticity. The studies on the Mediterranean climate by Gaussen, Emberger and others culminated years later in the proposal of a classification of bioclimatic vegetation belts (Ozenda 1975; Quézel 1979) which had a strong repercussion on the works of Rivas-Martínez on the Iberian Peninsula in the early 1980s, and which are discussed

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below. Finally, in the late 1970s it is worth noting the review by Daget (1977a, b), a continuation of Emberger’s work, on the concept of the Mediterranean climate. Walter’s genetic classification (1973) divides the biosphere into climate zones corresponding to large biomes, called zonobiomes, which respond to the latitudinal zoning of vegetation and are related to the general circulation of the atmosphere. Because this is a hierarchical type of classification, the zonobiomes are subdivided into subzonobiomes, and these in turn into biomes, natural geographic units. Zonobiomes are delimited from each other by contact zones called zonoecotones. Climate types are obtained from the relation between precipitation and temperature, factors that are represented graphically by climate diagrams (Walter and Lieth 1964; Walter et al. 1975). The zonobiomes were subsequently further subdivided into pedobiomes, a term applied to azonal vegetation types growing in extreme soil conditions, and orobiome, based on the altitudinal zonation of vegetation (Walter and Box 1976). In North America, the first works published in the twentieth century set out to study the climate from a range of approaches, either by applying Köppen’s worldwide classification to the climates of California (Russel 1926), or seeking to relate the physiological parameters of the plants to mean temperatures (Livinstong and Livinstong 1913). However, the most important American contributions to bioclimatology were the work of two authors: Thornthwaite and Holdridge. The first was also the closest competitor to Köppen’s system, whereas the second focused his studies on the Neotropical biogeographic kingdom. Thornthwaite devised two independent climate classification systems. The first (Thornthwaite 1931) was based on two indices that express thermal efficiency and precipitation effectiveness, the latter calculated from evaporation (in turn calculated from temperature), obtaining a very similar index to Martonne’s aridity index (Tuhkanen 1980). Thermal efficiency was calculated from the sum of the temperatures above 0
C. The author defined six thermal provinces which are dependent on precipitation effectiveness, finally producing a total of eight main climate types, named with letters as in the Kóppen system. He was the foremost critic of Köppen’s method, because of the use of a large number of climate elements derived from mean temperature and precipitation to define the limits of the vegetation, being considered an excessively empirical procedure (Thornthwaite 1943). Thornthwaite’s second classification (1948) is also based on two indices, potential evapotranspiration (ETP), calculated based on the mean monthly temperature and duration of daily insolation (inferred from the latitude); and the humidity index. Based on these indices—of which the first has greater importance in the classification—, eight types of thermal efficiency can be deduced, from the least efficient or coldest—called tundra—to the most effective or warmest—called megathermal. Based on the humidity index, the climate types range from arid to hyperhumid. The ETP index is calculated empirically by extrapolation from direct measurements of evaporation and transpiration and their subsequent correlation with temperature in different types of crops in the United States, where it has shown the best results

3 Bioclimatology

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in relation with vegetation distribution. However, he never published a global classification, only partial maps showing some or other indices, due to the doubts expressed by the author (Thornthwaite 1954) on the usefulness of his system outside North America. This last classification has subsequently been widely used, above all in North America, due to the experimental crop measures on which it is based and to its relationship with the water requirements and availability of plants (Sanderson 1948). Its assessment as an empirical method was positive for periods of over 1 month, when there was only a small variation between ETP and temperature, meaning that the estimative errors were not very large (Pelton et al. 1959). In other cases the formula has been modified (Wilcock 1950). Outside the scope of North America it has been used at the global or local level, as an index on its own (Elías Castillo and Ruiz Beltran 1978) or combined with other indices (Box 1981a; Rivas-Martínez 1987, 1990, 1993). Its practical utility derives on the one hand from the manifest need for bioclimatology studies to operate with some estimate of the water availability conditions for plants, and on the other, from its relative ease of calculation based exclusively on mean monthly temperatures, whereas other formulas using ETP require different types of climate parameters that are not always available from meteorological stations. Thornthwaite’s ETP has also had many detractors, both in regard to the delimitation of vegetation types (see Tuhkanen 1980), and to the assessment of the formula itself (Gentilli 1953). Holdridge (1947) proposed a predictive climate system based on temperature and precipitation. The climate spaces for different types of plant formations were determined by projecting isolines from these data (from tundra—rainy or desert—to tropical rainforest). He also proposed five extratropical thermoclimatic belts (alvar, alpine, subalpine, montane and lower montane). The author subsequently established a climate classification based on “life zones” (lifezone system; Holdridge 1959, 1966, 1967). The model used, which was originally two-dimensional (1947)—as explained earlier—was gradually modified around this new concept. Mean temperature was replaced (Holdridge 1959) by what was called mean temperature of comparative plant growth, defined as the sum of the positive mean temperatures—over 0
C—in a particular period of time (year, month, week and so on), a term that would subsequently become known as biotemperature (Holdridge 1966). The author also devised a formula to calculate evapotranspiration based on biotemperature (Holdridge 1959). Finally, the model was built using data for annual precipitation, potential evapotranspiration, altitude and mean annual biotemperature (Holdridge et al. 1971). The division is represented in a logarithmic system and produces areas of equal climatic significance corresponding to different vegetation types. A new extratropical bioclimatic belt also emerges, called premontane (Fig. 3.1). Although this system may appear very comprehensive it has not been widely used, except in tropical areas of America and in some areas in Asia (Boonpragob and Santisirisomboon 1996; Pan et al. 2003).

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Fig. 3.1 Tridimensional representation of climate classification based on “life zones” (lifezone system) by Holdridge (1959, 1966, 1967)

A large number of works have been published in the last 20 years of the XX century on the relations between plants (vegetation) and climate, featuring numerous approaches to this complex phenomenon. Some authors have attempted to interpret these relationships by studying the ecophysiological responses of plants to climate as a way of predicting their distribution (Woodward and Williams 1987; Woodward 1987). The works of Box (1981a, b, 1982, 1987) are based on the idea that the biological forms (biotypes) of plants to some degree signify adaptations to the climate in the territory and therefore constitute a preliminary tool for climatically characterising the main types of plant formations at both the global and local scale. Other authors have set out to establish the criteria for a climate classification according to the phytogeographical zonation of the vegetation. In this regard the works by Daget (1977a, b) and Daget and David (1982) focused, as has been mentioned, on the characterisation and delimitation of the Mediterranean region, revising and comparing indices and classifications from the French school, particularly those of Emberger. Elsewhere, Tuhkanen (1980) made an exhaustive bibliographic review of the proposed indices and classifications, particularly those applicable to northern Holarctic territories, with comparative analyses in the Scandinavian Peninsula. This type of study was subsequently developed by the same author, also in other Holarctic areas on the Earth (Tuhkanen 1984).

3 Bioclimatology

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The altitudinal zonation of vegetation on the Iberian Peninsula and in Western Europe as a whole has been reviewed by Rivas-Martínez (1981a, b, 1983). Unlike other authors, the systems of Rivas-Martínez are based on a highly detailed knowledge of the plant communities, to which is fitted a model of bioclimate belts, defined independently for the Mediterranean and Euro-Siberian regions, which initially follows the line taken by other authors from the French school (Ozenda 1954, 1975; Quézel 1979) although with a more explicit conceptual separation between the climatic continent (defined strictly by climate parameters) and its vegetation content (plant communities). These belts are in principle defined based on data for mean annual temperature and the minimums of the coldest month (RivasMartínez 1981a). This definition was subsequently extended with the formulation of the thermicity index (Rivas-Martínez 1983, 1984), which combines the information on the maximum and minimum temperatures of the coldest month as a way of offsetting the effects of the increase in thermal continentality in certain areas of the interior of the Iberian Peninsula. In addition, and unlike the previous systems, this climate classification considers the thermal parameters—which delimit the thermotypes—independently from the ombric parameters—which define the ombrotypes—: a bioclimatic belt is defined by the combination of a particular thermotype and ombrotype (Rivas-Martínez 1981a). Continentality was subsequently the subject of more detailed studies with the incorporation of the continentality index into the classification (Rivas-Martínez et al. 1991). The influence of precipitation on vegetation distribution (annual precipitation) also defined each biogeographical region (Rivas-Martínez 1993). Summer aridity also led him to delimit Mediterranean climate based on the values of the quotient between ETP (Thornthwaite) and precipitation for the summer period. In consequence, a set of monthly, bimonthly or seasonal indices were created (mediterraneity indices, RivasMartínez 1987). The attempt to extrapolate a system with these characteristics, initially prepared on the Iberian Peninsula, prompted the modification of some indices and the formulation of other new with a more general application in a Worldwide Bioclimatic Classification (Rivas-Martínez 1993). The main modifications introduced, compared to the first published works, refer to the differentiation of different types of Mediterranean bioclimates based on annual ombrothermic (IOT) and continentality indices, the adaptation of the thermicity index (compensated thermicity index (ITC)) based on continentality, the use of the mean annual temperature range as the simple attenuated continentality index, and a new definition of the ombrotypes based on the ITC values. Some of these indices were then used in other studies to compare vegetation and climate in large areas of the Iberian Peninsula (Gavilán and Fernández-González 1997; Gavilán et al. 1998; Gavilán 2005; Sánchez-Mata et al. 2017).

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3.1

3 Bioclimatology

South America

In 1994 Rivas-Martínez & Navarro proposed a Bioclimatic essay for South America based on the ‘Worldwide Bioclimatic Classification System’ (Rivas-Martínez 1997, 2001, 2008, 2010; Rivas-Martínez et al. 1999, 2011). It is a bioclimatic study at continental scale using bioclimatic indices. The classification makes possible to determine different climate types and their areas of geographic distribution. It also analysis quantitatively the relationship between climate and vegetation or any distribution of biodiversity using bioclimatic indices as commented above. It tries to be as comprehensive as others, together to an eclectic, predictive and reliable bioclimatic model that can be applied to any geographical point. Navarro and Maldonado (2002) have used this climatic model for Bolivia describing it as tropical at any altitude because it has a maximum rainfall in summer and at higher elevations the period of plant activity is continuous along the year. This two characteristics are typical of a tropical climate. Thus it covers latitudes up to 23
S in South America. In the present study we therefore have chosen this type of bioclimatic modelling to establish a detailed bioclimatic typology of Jujuy province that accurately reflects the climate types and their distribution in the territory, in addition to the correspondence between these types and the distribution of the main vegetation units.

3.2

General Bioclimatic Features from Las Yungas

According to Rivas-Martínez (2001, 2008), Rivas-Martínez et al. 1999, 2011) and Navarro and Maldonado (2002), subtropical mountain forests are found in the Tropical Pluviseasonal bioclimate with lower subhumid, upper subhumid and lower humid ombrotypes and lower and upper mesotropical thermotypes (Entrocassi et al. 2014). The highest precipitation in the study area occurs in the summer months, whereas the dry season extends from May to September or October. Precipitation follows the general pattern for the region and is mainly of an orographic type; that is, it is caused by the presence of mountain ranges that force the moist winds from the East to rise up their flanks. As they rise the water vapour they carry cools and precipitates in the form of abundant rainfall (Foëhn effect). Another important moisture contribution comes from the banks of mist that accumulate on the mountainsides (Buitrago 2000). The available climate data for the localities in the study area (Algarrobal, El Cucho, Las Capillas, Socavón, Arroyo Pacará and Mina 9 de octubre) indicate that the mean annual temperatures vary within a thermal gradient between 14.9
C and 17.5
C, while the mean annual precipitation ranges from 991 to 1472 mm. However, within the intervals of these gradients, the values of these bioclimatic parameters showed variations in specific sampling sites (transects), as discussed later in the

3.3 Bioclimatic Features of Serranías de Zapala Multiple Use Ecological Reserve

31

text (Entrocassi et al. 2014; Entrocassi 2016). As a result of the variations in temperature and precipitation as the elevation increases, forest sectors that are at a lower altitude grow in a warmer and less humid belt than forests at a higher altitude, which are cooler and more humid. Finally, we also include the bioclimatic typology of the province of Jujuy (Argentina), where the six localities are characterised bioclimatically with 121 meteorological stations within the study area. This typology satisfactorily represents the set of climate conditions prevailing in the territory in the province; in addition, the geographic distribution of the established climate types corresponds very closely to the distribution of the large vegetation formations in the area. It also outlines the foundations, concepts and methodological tools that were used for the bioclimatic characterisation of the 121 reference localities, leading to the determination of the different bioclimate types for the Province of Jujuy, represented on three maps.

3.3

Bioclimatic Features of Serranías de Zapala Multiple Use Ecological Reserve

In the bioclimatic characterisation of a given territory it is essential to use certain bioclimate indices to express quantitatively the influence of climate on the vegetation composition and distribution. A general bioclimatic characterisation was made before embarking on the bioclimatic characterisation of the study area, using the information from the six reference localities within the Reserve: Algarrobal, El Cucho, Las Capillas, Socavón, Arroyo Pacará and Mina 9 de Octubre (Table 3.1). To obtain a specific bioclimatic analysis to detect and analyse the correspondence between the bioclimatic indices and the vegetation composition and distribution in the study area, a bioclimatic characterisation was made of the 120 relevés taken in the transects created within the reserve. Given that obviously there were no specific climate data for each transect, they were extrapolated from the altitude, exposure, physiognomy and geomorphology of the climate data in the reference sites closest to the sampling site. For each transect, the values were estimated for the most important bioclimatic parameters and indices to define the thermotype and ombrotype (Tables 3.2 and 3.3). As temperature is a variable whose linear response is only correlated with altitude, the extrapolations considered a drop in temperature of 0.6
C for every 100 metres of altitude according to the adiabatic gradient of saturated air from altitudes over 600 m asl (Buitrago 2000; Fernández-González 1997). With regard to the mean annual precipitation (P) and the annual positive precipitation (Pp), it should be noted that the exact amount could not be calculated in each transect, as precipitation has more irregular behaviour based on altitude. Unlike temperature, precipitation tends to present a relatively complex pattern that is statistically independent of a linear change with altitude, which explains the fact that precipitation in the sub-tropical forests has a unimodal response (Lomolino 2001; Blundo et al. 2012). It is well known that normally, and in a generalised

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Table 3.1 Bioclimatic characterisation of the reference meteorological stations within the Serranías de Zapla multiple use ecology reserve Meteorological station Bioclimate/Ombrotype Algarrobal Tropical Pluviseasonal Upper Subhumid El Cucho Tropical Pluviseasonal Upper Subhumid Las Capillas Tropical Pluviseasonal Upper Subhumid Socavón Tropical Pluviseasonal Lower Humid Arroyo Pacará Tropical Pluviseasonal Lower-Upper Subhumid Mina 9 de Tropical Pluviseasonal octubre Lower Humid

Thermotype Lower Mesotropical Lower Mesotropical Lower Mesotropical Lower Mesotropical Lower Mesotropical Upper Mesotropical

Altitude (m asl) T 1200 16.7

P 991

It 408

Io 4.9

1230

16.5

1079

403

5.4

1185

16.7

1010

409

5

1150

16.4

1294

408

6.6

1015

17.5

992

429

4.78

1450

14.9

1472

372

8.2

T Mean annual temperature (
C), P Mean annual precipitation (mm), It Thermicity index, Io Annual ombrothermic index Source: Entrocassi et al. (2014) Table 3.2 Bioclimatic parameters and indices used to determine the thermotypes and ombrotypes of the vegetation plots (Rivas-Martínez 2001, 2008; Rivas-Martínez et al. 1999, 2011) Bioclimatic parameters and indices Mean annual temperature Mean maximum temperature of the coldest month of the year Mean minimum temperature of the coldest month of the year Annual positive temperature

Abbrev./formula T M

Definition Total mean monthly temperature Mean maximum temperature of the coldest month of the year

m

Mean of the minimum temperatures of the coldest month of the year

Tp

Total mean monthly temperature of the months with a mean temperature higher than 0
C. If all the months of the year have a mean temperature higher than 0
C, then Tp ¼ T  12 Total mean monthly precipitation Total mean precipitation of the months whose mean temperature is higher than 0
C. If all the months of the year have a mean temperature higher than 0
C, then Pp ¼ P Total mean annual temperature, mean maximum temperature of coldest month and mean minimum temperature of the coldest month, multiplied by 10 Quotient between the total mean precipitation of the months whose mean temperature is higher than 0
C (Pp) and the total mean monthly temperatures also higher than 0
C (Tp), multiplied by 10

Mean annual precipitation Annual positive precipitation

P Pp

Thermicity index

It ¼ (T + M + m).10

Annual ombrothermic index

Io ¼ (Pp/Tp).10

3.4 Bioclimatic Features from Jujuy Province

33

Table 3.3 Range of bioclimatic indices (Rivas-Martínez 2001) used to determine the thermotypes Index Thermicity index It ¼ (T + m + M).10 Ombrothermic index Io ¼ Pp/Tp.10

Range 395–490 320–395 3.6–4.8 4.8–6 6–9

Thermotype Lower mesotropical Upper mesotropical Lower subhumid Upper subhumid Lower humid

manner, precipitation in mountain areas tends to increase until a certain altitude, after which it begins to decrease (Buitrago 2000; Navarro and Maldonado 2002). However, these variations based on altitude are difficult to estimate numerically. Due to the difficulties inherent in this situation, we worked with the precipitation values for the reference sites without making corrections for altitude, in order to obtain at least an approximate precipitation value for each transect. After calculating the bioclimatic parameters and indices, we determined the bioclimate, thermotype and ombrotype for the 120 transects, obtaining as a result a precise and specific bioclimatic characterisation for each vegetation plot. These data were used to prepare a bioclimatic matrix which shows the variations in the bioclimatic parameters and indices along the altitudinal gradient of the forests in the study (see Appendix A).

3.4

Bioclimatic Features from Jujuy Province

There are relatively few climate studies on Jujuy province. Buitrago (2000) analyzed the climatic information on the province using the Köppen classification (1936) to describe the climate types present in the area. There are other kind of classifications that expressed the quantitative relationship between climate and organisms distribution, mainly vegetation because it is the most obvious component of ecosystems. In 1994 Rivas-Martínez & Navarro proposed a Bioclimatic Essay for South America based on the Worldwide Bioclimatic Classification System using a range of bioclimatic indices (Rivas-Martínez 1990, 2008; Rivas-Martínez et al. 1999). Following this classification we have described the Bioclimatic typology of Jujuy province (Entrocassi et al. 2014). We have used simple climate parameters such as Mean annual temperature (T), Mean monthly temperature (Ti), Mean maximum temperature of the coldest month (M), Mean minimum temperature of the coldest month (m), Annual positive temperature (Tp), Positive temperature of the driest 6 months of the year (Tpd2); Mean annual precipitation (P), Mean monthly precipitation (Pi), Annual positive precipitation (Pp), Positive precipitation of the driest 6 months of the year (Ppd2), Precipitation in the warmest month of the year (Pss) and Precipitation in the coldest 6 months of the year (Psw). Some parameters such as mean maximum temperature of the coldest month (M) or the mean minimum temperature of the coldest month

34

3 Bioclimatology

(m) were estimated by first calculating the altitudinal thermal gradient, and then adding or subtracting this value from the known values for M and m for one of the localities considered in the extrapolation since there were no records for such parameters in some localities. Bioclimatic indices from those parameters were also calculated such as Thermicity index (It), Annual ombrothermic index (Io), and Ombrothermic index of the driest 2 months of the year (Iod2). Based on the values of the simple climate parameters and bioclimatic indices and using specific keys we determined the macrobioclimate and the bioclimates for Jujuy province. Different bioclimatic belts (thermotypes and ombrotypes) in each bioclimate, and their bioclimatic horizons (upper and lower) were also determined. For the determination of the macrobioclimate the following parameters were considered: Mean annual temperature (T), Mean maximum temperature of the coldest month (M), Thermicity index (It), Precipitation in the warmest 6 months of the year (Pss) and Precipitation in the coldest 6 months of the year (Psw). We considered the values of the annual ombrothermic index (Io) and the ombrothermic index of the driest 6 months of the year (Iod2) to determine the predominant bioclimate in each locality. To determine the thermotypes and ombrotypes and their horizons (upper and lower) we considered the values of the thermicity index (It) and the annual ombrothermic index (Io); when necessary—as a consequence of the influence of altitude—we used the Annual positive temperature (Tp). Finally, we compiled three bioclimatic maps showing, respectively, the bioclimates, the thermotypes and the ombrotypes. Geographic Information System tools were used to compile all the data and information extracted by generating vectorial layers to define the bioclimatic unit limits and its geographic boundaries throughout the province, and then we could compile the bioclimatic maps of Jujuy province (see Entrocassi et al. 2014). Province of Jujuy falls within the Tropical Macrobioclimate at any altitude (Fig. 3.2). It also presents two different bioclimates: Tropical Pluviseasonal and Tropical Xeric. The province is located in the called Eutropical- Subtropical latitudinal belt (21
460 -24
360 S) and is intersected by the Tropic of Capricorn, receiving large amounts of solar radiation that determines the existence of continuous vegetative activity throughout the whole year. The region of the lower valleys and Sierras Subandinas has a Tropical Pluviseasonal bioclimate with thermo-, meso- and supratropical thermotypes (Fig. 3.3) and humid and subhumid ombrotypes (Fig. 3.4). It is an environment of great topographic complexity and marked altitudinal gradient, between 300 and 4150 m asl. It appears in the south of Jujuy, covering a vast region exposed to warm air masses from the Atlantic Ocean. It comprises the sub-Andean mountain ranges, the Santa Bárbara mountains, and the San Francisco river valley, following an altitudinal gradient of 320–2120 m asl approximately. However, warmer valleys located at the threshold to the Chaco Plain mainly show a dry Tropical Xeric bioclimate. It is situated in the far south-southeast of the province, occupying dry low-lying valleys and mountain ranges between 400 and 1100 m asl approximately. In the Puna region and the Eastern Cordillera (Eastern Andes mountain range), above 2500 m asl, dominates the Tropical Xeric bioclimate. It covers the greatest

3.4 Bioclimatic Features from Jujuy Province

Fig. 3.2 Bioclimates of Jujuy province. From Entrocassi et al. 2014

35

36

Fig. 3.3 Thermotypes of Jujuy province. From Entrocassi et al. 2014

3 Bioclimatology

3.4 Bioclimatic Features from Jujuy Province

Fig. 3.4 Ombrotypes of Jujuy province. From Entrocassi et al. 2014

37

38

3 Bioclimatology

area of the province extending to areas exposed to moist air masses, in general ranging between 2000 and 4100 m asl approximately. They occupy high valleys, high plains, high mountains and the Quebrada de Humahuaca mountain valley. Respect to thermothypes, supra-, oro- and criorotropical belts appear in the area and ombrotypes ranging from dry to semiarid are the most characteristics. In this territory there are some high, cold and subhumid localities between 2450 and 4150 m asl with Tropical subhumid Pluviseasonal bioclimate.

3.5

Final Remarks on Jujuy Bioclimates

– The most extreme situations in the lower supratropical belt—both, the Tropical Pluviseasonal and Tropical Xeric bioclimates—appeared between 2000 and 2500 m asl (approx.), including the most humid and the driest areas in Jujuy province. – The most humid areas are found between 1950 and 2450 m asl within the Tropical Pluviseasonal bioclimate; this altitudinal range is where the effect of orographic precipitation is intensified, a situation that is borne out by the high values of the ombrothermic indices (Table 3.4) and by the type of vegetation, characterized by large expanses of rainforest and cloud mountain grassland. – The driest areas are found between 2078 and 2461 m asl within the Tropical Xeric bioclimate, and are located in the region of Quebrada de Humahuaca. – In the territories below 1950 m asl with a Tropical Pluviseasonal bioclimate the temperatures rise sharply and precipitation decreases gradually, a situation that favours the establishment of warmer and less humid bioclimatic belts. – Above 2500 m asl, in the supratropical and orotropical localities of the Puna and Cordillera Oriental (between 3000 and 4100 m asl) in localities oriented to moisture winds, there is a decrease in temperature and precipitation. Those colder and less humid bioclimatic belts allow to get at these altitudes a Tropical Pluviseasonal bioclimate (Table 3.4). – The territories with the Tropical Xeric bioclimate show two different bioclimatic patterns according to their altitude. In the low, warm dry valleys in the southsoutheast of the province located below 1950 m asl, temperature and precipitation increase at the same time, a situation which allows the establishment of warmer and less dry bioclimatic belts (Table 3.5). In the Puna and Cordillera Oriental region above 2500 m asl (between 2500 and 4100 m asl) temperature decreases and precipitation increases slightly, allowing the presence of colder and less dry bioclimatic belts, but slightly more humid (Table 3.5). – This research also reflects the relationship between climate and the distribution of vegetation at the province level. On the other hand, it will provide valuable information that may be used in future research and applied projects planning, management and conservation of natural resources of this wide territory.

1950–2450

3000–3520

Lower Supratropical

Upper Supratropical

Tropical Pluviseasonal Upper Subhumid to Upper Humid Tropical Pluviseasonal Lower Subhumid to Upper Subhumid

1345–1790

Upper Mesotropical

Tropical Pluviseasonal Lower Subhumid to Lower Humid

500–1300

Altitude (m. asl) 320–500

Lower Mesotropical

Thermotype (Map 2) Upper Thermotropical

Tropical Pluviseasonal Lower Subhumid to Lower Humid

Bioclimate and ombrotype (Maps 1 and 3) Tropical Pluviseasonal Lower Subhumid

8.4–9.9

11.9–12.9

13.8–16.1

16.1–20

T(
C) 20–21.3

Table 3.4 Tropical Pluviseasonal bioclimate in Jujuy Province

405–578

805–1470

646–1472

726–1385

P (mm) 800–1150

161–211

269–297

329–390

403–489

It 491–512

3.9–4.8

5.6–9.7

3.7–8.6

3.6–7.1 

Io 3.6–4.8 

Cianzo, El Durazno, Tafna

(continued)

Localities studied Caimancito, Calilegua, El Piquete, El Talar, Florencia, Fraile Pintado, La Lucrecia-Las Moras, La Toma-Chalicán, Ledesma, Lote Sora, La Paulina, Puesto Viejo, Río Piedra-Bananal, Yuto Afatal, Algarrobal, Arrayanal (río Candelaria), Arroyo Pacará, Las Capillas, El Cucho, El Palmar, Las Lajitas, Los Alisos (Policía), Los Perales, Madrejón, Palpalá, Peña Alta (Valle Grande), Puerta de Sala-Carahunco, Río Blanco, San Salvador de Jujuy, Santa Bárbara, Sauzal, Siete Aguas, Socavón Corral de Piedras, El Fuerte, El Morado, El Paño, Guerrero, juntas del Morado-negro, León, los Alisos (Arriba), los Nogales, Mina 9 de Octubre, Ocloyas, Pampichuela, San Antonio, san Lucas (Valle Grande), Termas de Reyes, Tilquiza, Tiraxi, Valle Grande, Villa María, Yala El Molulo, Laguna El Rodeo, Repetidora Canal 13 II, San Bernardo

3.5 Final Remarks on Jujuy Bioclimates 39

Altitude (m. asl) 3510–3950

4150

4500?

Thermotype (Map 2) Orotropical Inferior

Upper Orotropical

Cryorotropical

310

–

–

P (mm) 375–454

5.7

T(
C) 5.7–7.7

–

91

It 112–149

–

4.5

Io 4.3–5.7

No localities studied

Lulluchayoc

Localities studied Cangrejillos, Cieneguillas, Cóndor, Mina Pan de Azúcar, Rinconada, Santa Catalina

T Mean annual temperature, P Mean annual precipitation, It Rivas-Martínez thermicity index, Io Rivas-Martínez annual ombrothermic index From Entrocassi et al. 2014

Bioclimate and ombrotype (Maps 1 and 3) Tropical Pluviseasonal Lower Subhumid to Upper Subhumid Tropical Pluviseasonal Lower Subhumid Tropical Pluviseasonal Subhumid?

Table 3.4 (continued)

40 3 Bioclimatology

254 –

5.9 –

3560–3920

4100

4300?

Upper Orotropical Cryorotropical

6.6–8.6

156–357

152–315

Lower Orotropical

8.1–12

2980–3573

112–392

431–737

Upper Supratropical

11.5–14.3

17.7–20.2

P (mm) 546–704

2078–2500

520–1090

Lower Mesotropical

T (
C) 19.8–20.7

Lower Supratropical

Altitude (m. asl) 433–578

Thermotype (Map 2) Upper Thermotropical

From Entrocassi et al. 2014 a A locality with Io ¼ 1.8 b Three localities with Io < 1

Tropical Xeric Lower Semi-arid to Lower Dry (with an Upper Arid enclave) Tropical Xeric Lower Semi-arid to Upper Dry Tropical Xeric Upper Semi-arid to Upper Dry Tropical Xeric Upper Dry Tropical Xeric Dry?

Bioclimate and ombrotypes (Maps 1 and 3) Tropical Xeric Lower Dry to Upper Dry Tropical Xeric Upper Semi-arid to Upper Dry

Table 3.5 Tropical Xeric bioclimate in Jujuy Province

–

96

121–158

160–239

245–323

425–490

It 491–527

–

3.6

2–3.6

1.2–3.2

No localities studied

Abdón Castro Tolay, Abra Pampa, Coctaca, El Moreno, Humahuaca, Iturbe, La Quiaca, Palca de Aparzo, Puesto del Marques, Yavi Abra Laite, Barrios, Cochinoca, Inti Cancha— Suripugio, Oratorio, Pumahuasi, Sey, Susques, Tres Cruces Coranzulí

Arroyo Colorado, Bajada de Pinto (Lavayén), Barro Negro, El Milagro, Fuensanta, La Escuela, La Mendieta, Las Maderas, Las PeñasLas Delicias, Los Lapachos, Miraflores, Palos Blancos, Pampa Blanca, Perico, San Antonio (La Esperanza), San Juan de Dios, San Juancito, San Juancito (La Mendieta), Santa Clara Caspalá, Huacalera, Maimará, Purmamarca, Tilcara, Tumbaya, Uquía, Volcán

1.8–3.1a

1–2.3b

Localities studied Chalicán, El Quemado, La Esperanza, San Pedro, Villa Las Rosas

Io 2.2–2.9

3.5 Final Remarks on Jujuy Bioclimates 41

Chapter 4

Geobotany of Serranías de Zapla Multiple Use Ecology Reserve: Flora and Vegetation

The composition and distribution of the vegetation in the subtropical montane forests of the Serranías de Zapla Multiple Use Ecology Reserve was determined using the phytosociological methodology of the Zurich-Montpellier school (Braun-Blanquet 1979) adapted to the study area, which allowed the identification and delimitation of the different plant communities along the altitudinal gradient in the Reserve.

4.1

Field Data Collection

After prior explorations in the study area, preferential samplings were made to select physiognomically, floristically and ecologically homogeneous areas, in order to ensure the existing plant communities were adequately represented (Matteucci and Colma 1982; Ramírez et al. 1997; Navarro and Maldonado 2002; Navarro and Ferreira 2009, 2011; Navarro 2011; Alcaraz 2013). 120 sampling plots were made (transects) in these areas at different altitudes and topographic exposures, covering a total of 117,500 m2; 115 transects had an area of 1000 m2 (each one 10 m wide by 100 m long), which is a sufficient and representative size for sampling in tropical and subtropical woody areas (Gentry 1982). Only five transects had a lesser area (500 m2) due to their particular situation based on the forest type and the geomorphology of the site. The samplings were carried out during numerous campaigns in spring and summer of 2007 and 2008. A phytosociological relevé was recorded in each transect following the methodology of Braun-Blanquet (1979), which weighs the abundance and dominance (or coverage) of a species through phytosociological indices on a scale of 1 to 5 (Table 4.1); the two lower indices (+ and r) register the abundance while the others

© Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_4

43

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

44

Table 4.1 Braun-Blanquet abundance-dominance scale (1979). See text for additional explanations Significance r + 1 2 3 4 5

Index A single individual, negligible coverage More individuals, very low coverage Abundant but with low coverage, or not very abundant with a coverage of less than 5% Very abundant and with less than 5% coverage, or not very abundant with a coverage of 5–25% 25–50% coverage, regardless of the number of individuals 50–75% coverage, regardless of the number of individuals Coverage equal to or over 75%, regardless of the number of individuals

(1–5) take account of the dominance (Alcaraz 2013). The absence of a species in a particular relevé was noted with a dot (.). A total of 120 phytosociological relevés were made, and the tree, shrub and herb species present were recorded in each one. Their phytosociological indices were consigned according to the aforementioned abundance-dominance scale (Table 4.1). In each relevé, data were also noted for height above sea level, geographic coordinates, exposure of the slopes and other ecological data that we considered significant (canopy height, open or closed forests, perennial or deciduous foliage, poorly developed soils, etc.) and whether the relevé was located in foothills other type of ecological information was also recorded such as slopes, gorges, ridges, terraces or river beaches. The number of species present in each relevé was used to determine the variation in species richness of the forests studied along the environmental gradient and to obtain the bioclimatic belts for the study area (Fig. 3.3). All the plants were collected and many identified in the field and subsequently corroborated in laboratory-based studies.

4.1.1

Identification, Species Richness and Life Forms of Species

The taxonomic identification of species in the study area was done by consulting the specific bibliography in order to build a reliable floristic list of all the species recorded (Lourteig and O’Donnell 1942, 1943a, b; Burkart 1952, 1974, 1979, 1987; Font Quer 1953; Lourteig 1955; Barkley 1957; Digilio and Legname 1966; Parodi 1972; Sorarú 1972; Orsi 1976; Cabrera 1978, 1983, 1993; Cabrera and Zardini 1978; Giberti 1979; Hutchinson 1982; Legname 1982; Hunziker 1984, 1995/2007; Guaglianone 1987; Nicora and Rúgolo de Agrasar 1987; Biloni 1990; Novara 1991/2007, 1994; Boelcke 1992; Dottori and Hunziker 1994; Zuloaga and Morrone 1996, 1999a, b; Zapater et al. 2004; Seo and Xifreda 2008; Zuloaga et al.

4.1 Field Data Collection Table 4.2 Presence index used in synthetic phytosociological tables based on species frequency

45 Presence index I II III IV V

Frequency (%) 0–20 20.1–40 40.1–60 60.1–80 80.1–100

2008; Zuloaga 2014). The scientific nomenclature was updated and the status assigned to each species (native, endemic, exotic, etc.) according to the catalogue of vascular plants of the Southern Cone (Zuloaga 2014). The total species richness (number of species) was determined for each vegetation stratum and for all the botanical families recorded, in addition to the number of trees, shrubs and herb species per botanical family. The frequency (rate of presence) was also calculated for each of the 257 species registered in the total plots collected, and expressed as a percentage, which was used to determine the presence indices in the synthetic tables of relevés (Table 4.2). Finally, the number of species in each physiographic group was counted and represented in a histogram. In order to obtain a characterisation that corresponded more closely to the set of communities and plant species comprising the forest formation in the study area, the biological types of each species were identified according to the classification of Raunkiaer (1934). The percentages of each biological type were then calculated, and the corresponding spectrum was built. The use of this classification offers important support for the characterisation of a particular plant formation, as the spatial organisation of the biological types of the dominant plants is the factor that determines its physiognomy (Arozena 2000; Rivas Martínez 2001).

4.1.2

Analysis of Floristic-Phytosociological and Bioclimatic Data

The floristic-phytosociological data contained in the relevés were transferred to a primary matrix (raw table) comprising 120 columns (relevés) and 257 rows (species) including the abundance-dominance value of each species recorded according to the Braun-Blanquet scale. Such value must be firstly transformed to an ordinal scale in order to be able to apply any kind of numerical analyses. There are different methods to do it depending if the data are of presence/absence (Williams and Lambert 1959) or uses other type of data. We have followed the Westhoff and Van Der Maarel (Van der Maarel 1979) ordinal scale, one of the most used transformations. Phytosociological values are transformed to an ordinal 1–9 scale, while the value ‘2’ is splitted into three parts attending the cover percentage criteria used in Barkman et al. (1964). The techniques used for the statistical treatment of the floristic-phytosociological and bioclimatic information were numerical classification analysis (hierarchical

46

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

classification analysis) and direct gradient analysis (canonical correspondence analysis). These multivariate analysis methods were selected because they reduce the dimensionality of the data and detect the variables that condition their structure, conserving the greatest variability and producing the least possible distortion (Pielou 1984; Fariñas 1996). Numerical classification analysis is a more objective way to obtain homogeneous groups of relevés based on attributes. There are two types of classification: hierarchical and non-hierarchical. The first obtains homogeneous groups that are successively merged according to a hierarchy; in this case the homogeneity decreases as the groups become larger. It makes possible to determine and understand the relationships between them. In contrast, the application of non-hierarchical techniques forms homogeneous groups without establishing relations between them (Escudero et al. 1994). Numerical classification techniques can be divided into agglomerative and divisive. With agglomerative techniques, the individuals or objects (in this case relevés) become progressively merged and form groups. That is, each relevé is processed individually and so on until they are all merged. Divisive techniques start with the whole group of data as a conglomerate and successively divide it into smaller groups. There is a diversity of opinion with regard to the efficacy of these techniques. Some authors are in favour of agglomerative techniques, whereas others maintain that divisive techniques are more effective when working with ecological data. Agglomerative techniques are polythetic, whereas divisive techniques may be monothetic or polythetic. Monothetic classification is based on a single very relevant characteristic and uses as its classificatory criterion the presence or absence of a species; whereas polythetic classification is based on a large number of characteristic species and the usual procedure in this case is agglomerative. In general, polythetic classifications are more frequently used due to the fact that the random presences or absences of some species can produce significant deviations, mainly when working with a reduced number of species or relevés (Escudero et al. 1994). The gradient analysis encapsulates a set of techniques that explain and describe the relations between the vegetation data and the environmental variable data. The communities can therefore be interpreted based on the species’ response to the environmental gradients. Notable among the different gradient analysis techniques is canonical or constrained ordination, which distinguishes the set of environmental variables that best explains the variations in species abundance (Ter Braak 1986; Ter Braak and Prentice 1988). The present study uses canonical correspondence analysis as it is a direct gradient analysis technique that includes two types of variables, one biotic and another abiotic, and which directly relates the patterns in variation in the communities’ composition with the environmental variations (Escudero et al. 1994; Lozada Dávila 2010). A hierarchical classification analysis was applied to the primary data matrix (hereinafter HCA) in order to organise the floristic-phytosociological data obtained, delimit groups of homogeneous relevés and determine the relations existing between them. This was done using the statistical package Tilia 1.7.14 (Grimm 1992),

4.1 Field Data Collection

47

initially created for pollen data, but widely used with stratigraphic, geological and ecological data, particularly for vegetation studies that involve natural gradients and use transects as sampling plots; this package produced excellent results compared with other tests made with other computer programs. The grouping analysis was done by divisive classification with the CONISS program (Constrained Incremental Sum of Squares) (Grimm 1987) in the Tilia package. This program uses Euclidian distance as a measure of similarity between the samples (relevés), with the possibility of applying restrictions to the classification or not. No restrictions were used in this case, as it was not important to maintain the order in which the relevés had been made. An ordered phytosociological matrix of relevés in homogeneous groups was obtained from the classification analysis, easier to interprete the relationships between the groups. This matrix had to be divided into three parts because of the number of species and relevés. The groups were arranged in increasing order of altitude, except for the group of riverine relevés, located last of all; data on altitude, area and species richness were also included, for each relevé. The relationship between the floristic-phytosociological data and the environmental variables was analysed using canonical correspondence analysis (hereinafter CCA) implemented in the CANOCO program (Ter Braak 2002). This analysis verifies whether there is a statistically significant relation between the communities’ composition and these variables. The ordered phytosociological matrix was therefore processed jointly with the bioclimatic matrix (Appendix A), and the results were displayed in two ordination diagrams (“triplots”) showing the species, the entries and the environmental variables studied, and offering a general quantitative description of the species habitat.

4.1.3

Synthesis of the Floristic-Phytosociological Data

Given that plant communities are vegetation types that are identifiable due to a particular floristic combination, and that within the set of species integrating them there are some that are better indicators of interspecies relations and relations with the environmental variables, the species with a diagnostic value can be used to identify, delimit and characterise plant communities (Alcaraz 2013). In order to determine these diagnostic species, a general comparative synthetic table was prepared to summarise the information on the groups obtained through the presence indices in the hierarchical classification analysis (HCA) (Appendix C). Based on a direct comparison of the presence indices and the phytosociological values of the recorded species, the degree of restriction of each species to the groups obtained in the HCA was determined along with the characteristic or indicator species and the combination of characteristic taxa in each group. The criteria for this selection primarily valued the fidelity (Table 4.3), physiognomy and stenoicity

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

48

Table 4.3 Degrees of fidelity (Gf) Gf 5 4 3 2 1

Character Exclusive species, confined to a single plant community or group of communities Selective species, with a clear preference for a particular plant community Preferential species, which although present in several communities, is more abundant or has greater vitality in the one that is the object of the study Companion species, indifferent, without a marked preference for any plant community Accidental species, which clearly has its optimum in another community

Source: Alcaraz Ariza (2013)

Table 4.4 Criteria used to select the characteristic or indicator species (based in Navarro and Maldonado 2002) Criteria Phytosociological fidelity Physiognomy Ecological stenoicity

Endemics Dynamic-successional aspects Biogeographical aspects

Description Species with high or significant phytosociological values and presence indices in a particular group Dominant species that contribute to the appearance of the vegetation Species most restricted to certain environments characterised by their specific ecological features (altitude, bioclimate, soil, geomorphology, etc.), with high or significant phytosociological values and presence indices Endemic species in subtropical montane forests Species relevant to the substitution stages of the forest Relevant species that do not belong to the biogeographic unit to which the sub-tropical montane forests are ascribed

of the species, and in some cases also considered endemics and dynamicsuccessional and biogeographic aspects. These criteria were based on the proposals of Navarro and Maldonado (2002) for the vegetation in Bolivia (Table 4.4) and were adapted for the purposes of the present study. The selection of these species was subsequently corrected, adjusted and defined based on the results of the canonical correspondence analysis (CCA), analysing the relationship between species and environmental variables.

4.1.4

Floristic Composition

The total sample area was determined in the subtropical mountain forests of the Serranías de Zapla Multiple Use Ecology Reserve. 257 species were recorded belonging to 194 genera and to 66 botanical families (Appendix B). Of the total species recorded (257), 216 are native (84%), 25 are endemic (10%), 13 are exotic (5%) and 3 are cosmopolitan (1%). The native species are

4.2 Species Richness

49

autochthonous to Las Yungas and their distribution may be restricted to the adjacent countries in the same biogeographical unit, and at most to contiguous units. The exotic species include adventitious and cultivated plants from other regions, many of which have become naturalised in Las Yungas. This is the case of Anagallis arvensis, Duchesnea indica, Eleusine indica, Leonurus japonicus, Morus alba, Mirabilis jalapa, Primula malacoides, Verbascum virgatum, Veronica arvensis and Veronica persica. Of the 66 botanical families recorded, 31 belong to the tree layer, 25 to the shrub layer and 37 to the herbaceous layer.

4.2 4.2.1

Species Richness Vegetation Structure (Stratification) and Floristic Composition

In terms of the total number of species in each vegetation layer, 107 species were recorded in the herbaceous layer, 85 in the shrub layer and 65 in the tree layer. This shows that the herbaceous layer had the greatest richness at the level of species and botanical families. The analysis of species richness in each botanical family revealed that of the 66 families recorded, five have the greatest number of species, namely: Asteraceae (54), Fabaceae (20), Poaceae (18), Solanaceae (16) and Euphorbiaceae (15). The rest of the families have between one and seven species (see Appendix B). With regard to the species richness of the families in each vegetation layer, it was observed that the family with the greatest number of tree species was Fabaceae (with 12 species), followed by Euphorbiaceae (5), Anacardiaceae and Myrtaceae (4), Asteraceae, Celtidaceae and Rutaceae (3); the rest of the families have one or two tree species (Fig. 4.1). With regard to the shrub layer, the best represented family was Asteraceae (with 28 species), followed by Solanaceae (8), Euphorbiaceae (7), Fabaceae (5), Amaranthaceae and Malvaceae (4); the rest of the families have between one and three shrub species (Fig. 4.1). Finally, in the herbaceous layer the families with the greatest number of species were Asteraceae (23) and Poaceae (18), followed by Plantaginaceae and Solanaceae (6), Acanthaceae and Rubiaceae (4); the rest of the families have between one and three herbaceous species (Fig. 4.1).

4.2.2

Richness in the Relevés

Of the 120 relevés made, 52 had between 20 and 40 species, 39 between 40 and 60 species, and 20 between 60 and 80 species; only five relevés had fewer than 20 species and four were considerably richer, with over 80 species. (Fig. 4.2). A relation was also observed between species richness and the size of the sampling

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

50

30 25 20 15 10 5 Acanthaceae Alstroemeriaceae Anacardiaceae Apocinaceae Araceae Asteraceae Begoniaceae Betulaceae Boraginaceae Budlejaceae Caricaceae Combretaceae Escalloniaceae Fabaceae Juglandaceae Lauraceae Lythraceae Malvaceae Meliaceae Myrsinaceae Nyctaginaceae Orobanchaceae Piperaceae Poaceae Polygonaceae Ranunculaceae Rosaceae Rutaceae Samolaceae Sapotaceae Smilacaceae Turneraceae Valerianaceae

0

Trees

Shrubs

Herbs

Fig. 4.1 Bar chart showing the total number of trees, shrubs and herbs by family

120

Número de especies

100 80 60 40 20

1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85 88 91 94 97 100 103 106 109 112 115 118

0 Número de orden de los Inventarios

Fig. 4.2 Species richness of the relevés compiled in the study area (Relevé N vs N species)

plots (transects), as most of the smaller relevés (less than 500 m2) were poorer in species (relevés 40, 42, 52, 90 and 91). It should be noted that their size was limited by the geomorphological characteristics of the sampling sites in the field as they were taken in gorges, ravines, moderately or steeply sloping foothills and riparian environments. Except for these relevés, the others (with 1000 m2) had greater species richness, most between 20 and 60 species, whereas the high richness of some relevés may also be linked to their location in sites that are difficult to access and very little altered, where the vegetation is very well conserved.

4.3 Vegetation

4.2.3

51

Frequency of the Recorded Species and Biological Spectrum

The analysis of the species frequency in all the relevés (Appendix B) revealed that a large number of species (174) are included in less than 20% of the relevés, 62 species are in 20–40%, 16 species are in 40–60%, four species are in 60–80%, whereas one single species is in over 80% of the relevés. The determination of the lifeforms (biotypes) of all the species recorded in the study area, based on the Raunkiaer classification (1934), allowed us to compile a biological spectrum showing the five biological types or biotypes. The analysis of this spectrum established that Phanerophytes represent the most abundant biotype, corresponding to the type of plant formation studied. This biotype had 147 species (57%) and included nanophanerophytes, microphanerophytes, mesophanerophytes and creepers, according to the criterion indicated by Braun-Blanquet (1979). These were followed by hemicryptophytes with 67 species (26%), therophytes with 21 species (8%), geophytes with 17 species (7%) and chamaephytes with six species (2%).

4.3 4.3.1

Vegetation Floristic-Phytosociological Data. Hierarchical Cluster Analysis (HCA)

The results of the Hierarchical Cluster Analysis (HCA) applied to the data matrix were represented graphically on a dendrogram (Fig. 4.3) that delimits two main branches: one upper (A), divided into three large clusters containing a total of 77 relevés (Clusters 1–3); and one lower (B) divided into three clusters with a total of 43 relevés (Clusters 4–6). The upper branch (A) is in turn divided into two branches that separate Clusters 1 and 2 from Cluster 3. All the relevés in these clusters are in the lower Mesotropical belt in the Tropical Pluviseasonal bioclimate, with lower and upper Subhumid ombrotypes (except relevé no. 106, which has an upper Humid ombrotype). Cluster 1 has 35 relevés in total and is subdivided into three smaller clusters (1A, 1B and 1C) (Fig. 4.3, Appendix A). Cluster 1A has eight relevés (53–60, according to the order established in the dendrogram), from the southern part of the study area; these relevés are located in the interval with lowest altitude (1032–1037 m asl), in the area around the locality of “Arroyo Pacará”. Cluster 1B has 15 relevés (23–26, 10–19 and 25; 1233–1275 m asl), and Cluster 1C has 12 relevés (1–8, 20–22 and 106; 1115–1258 m asl). Both clusters contain the relevés located mainly in the western part of the study area, in the zone of influence of the localities “Las Capillas”, “Algarrobal” and “El Cucho” and the site “Abra de Tunalito”, in addition to some located in the central zone.

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

52

UPPER MESOTROPICAL-LOWER HUMID

Inventarios

LOWER MESOTROPICAL- LOWER and UPPER SUBHUMID

CONISS 59 60 57 56 58 53 54 55 23 24 10 26 12 15 14 17 19 9 18 25 13 16 11 2 4 20 7 22 1 6 3 8 5 21 106 83 85 82 81 84 86 64 68 62 65 67 61 70 69 63 66 77 79 74 71 72 78 80 73 76 75 36 39 40 42 37 28 32 30 33 35 38 41 31 29 34 27 103 104 96 97 98 99 101 102 94 95 88 91 92 93 87 89 47 51 45 43 117 120 115 118 49 109 111 110 113 107 108 44 48 114 50 116 100 112 46 90 105 52 119

1A

1

1B

1C

SMb

2A

2

A

2B

2C 3A

3 3B

4

BM

B

5A

5 5B

SMa 5C

5D 6A

6 6B 2000

4000

6000

8000

10000

12000

14000

16000

18000

20000

22000

24000

26000

28000

30000

32000

Total sum of squares

Fig. 4.3 Dendrogram of the Hierarchical Cluster Analysis (HCA) based on the phytosociological values of the plant species in the study area. SMb: Basal mountain forest; SMa: High mountain forest; SMrib: Riparian mountain forest. Outlined in red: riparian relevés

4.3 Vegetation

53

Cluster 2 has a total of 26 relevés, all located in the western zone of the study area; this cluster is subdivided into three smaller clusters (2A, 2B and 2C) (Fig. 4.3, Appendix A). Cluster 2A has six relevés (81–86) from the area around the “Los Blancos” site (1192–1200 m asl). Cluster 2B has ten relevés (61–70) located on the “Centro Forestal” site (1085–1102 m asl). Cluster 2C also has ten relevés (71–80) from the area near the “Finca La Norteña” and “Finca Pedetti” zone and the locality of “Las Escaleras” (1105–1127 m asl). Finally, Cluster 3 has a total of 16 relevés located in the northern limit of the western part of the study area, in the area of influence of the site known as “El Nogalar” and “Finca Babni”. This cluster is subdivided into two smaller clusters (3A and 3B) (Fig. 4.3, Appendix A). Cluster 3A has five relevés (36, 37, 39, 40 and 42; 1235–1243 m asl), while Cluster 3B has 11 relevés (27–35, 38, 41; 1239–1253 m asl). The lower branch (B) is in turn divided into three major branches that separate Cluster 4 from Clusters 5 and 6. This branch of the dendrogram clusters the relevés located in the upper Mesotropical-lower Humid belt in the Tropical Pluviseasonal bioclimate and a small cluster of five relevés from the lower Mesotropical and the transition to the upper Mesotropical belt, with the same lower humid ombrotype (relevés 105, 107–110). All the relevés are located in the central part of the study area, except for two that belong to the riparian environments in the central-southern and southern zone of the study area and lie within the lower Mesotropical belt, with lower Subhumid (52) and lower Humid (105) ombrotypes. Cluster 4 has eight relevés (96–99, 101–104; Fig. 4.3, Appendix A), all located in the highest sector of the whole study area (1488–1620 m asl). Cluster 5 has a total of 27 relevés, all from the area around the locality known as “Mina 9 de Octubre”. Four smaller clusters can in turn be distinguished within this cluster (5A–D, Fig. 4.3). Cluster 5A has eight relevés located particularly on steeply sloping hillsides (91–94, 87–89; 1319–1360 m asl). Cluster 5B has nine relevés (43, 45, 47, 49, 51, 115, 117, 118 and 120; 1320–1433 m asl). There are six relevés in Cluster 5C (107–111, 113; 1180–1300 m asl), and four in Cluster 5D (44, 48, 50 and 114; 1310–1360 m asl). It should be noted that when compiling the ordinated phytosociological matrix and the bioclimatic matrix (Appendix A), some adjustments were made in three of these clusters, namely: Clusters 5B and 5D were gathered in a single cluster (hereinafter Cluster 5B–5D), given that both had similar phytosociological characteristics that warranted their combination although they were further apart on the dendrogram. Also, and for the same reasons, relevé no. 114 was included in Cluster 5C (in the dendrogram it appears in Cluster 5D). Finally, Cluster 6 has eight relevés and is subdivided into two smaller clusters (6A and 6B; Fig. 4.3, Appendix A). Cluster 6A is less uniform as it has five relevés (116, 100, 112, 46 and 90) arranged in a dispersed manner and at various altitudes on gorges and ravines with significant differences in topography, in some cases with rocky outcrops where water emerges (1285–1561 m asl). Cluster 6B has three relevés (105, 52 and 119); the first two from riparian environments (1015–1030 m asl), and the last located at a higher altitude (1417 m asl) in an environment with similar characteristics to those described for Cluster 6A. For these reasons, and

54

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

because of its phytosociological characteristics, this relevé was included in Cluster 6A (it appears in the dendrogram in Cluster 6B). Apart from these adjustments, the results obtained in the Hierarchical Cluster Analysis were consistent and highlighted the floristic and phytosociological relationships within and between the different clusters of relevés. Essentially, the dendrogram defines two major areas where the woodlands in the study are located: one lower, warmer subhumid level (lower Mesotropical and upper Subhumid) and another higher, more temperate and humid level (upper Mesotropical and lower Humid) (Fig. 4.3). These results serve as a valuable tool to support the interpretation of the information in the original matrix and contributed substantially to identifying and delimiting the plant communities in the study area.

4.3.2

Floristic-Phytosociological and Bioclimatic Data: Analysis of the Gradient. Canonical Correspondence Analysis (CCA)

The relationship between species abundance and distribution and environmental variables was analysed using Canonical Correspondence Analysis, applied jointly to the ordinated phytosociological matrix (Appendix B) and the bioclimatic matrix (Appendix A). This analysis used 257 species and three environmental variables: altitude, thermicity index (It) and ombrothermic index (Io). A CCA was obtained with 90.2% of the accumulated variance explained by axes 1 and 2. The results of the Montecarlo test conducted to calculate the significance of this relation (499 permutations) indicate that the values obtained with the CCA are statistically significant (p < 0.02), suggesting that the ordination diagram obtained (Fig. 4.4) offers an accurate representation of the contribution and distribution of the plant communities based on the environmental variables studied. The data in Table 4.5 extracted from the correlation matrix as a result of the CCA show the values of the three environmental variables for two species axes and two environmental axes. The correlation between species-environmental variables for axis 1 is 0.945 and 0.867 for axis 2. The eigenvalues of the ordination axes (Table 4.6) indicate that the first is the most important, explaining 61.3% of the variance extracted from the data for species-environmental variables; the second axis explains 28.9%; and the third axis explains 9.8% (Appendix C). As the first two axes contain the greatest percentage of the variance of the species-environment variance (90.2%), indicating that most of the information was concentrated on these two axes, the third axis has not been considered in the present analysis. The results obtained in the Canonical Correspondence Analysis were graphically represented in an ordination diagram with three types of units: species, relevés and environmental variables (triplot) (Fig. 4.4). This graph shows the relation between the three environmental variables analysed and the abundance and distribution of the

4.3 Vegetation

55

Fig. 4.4 Ordination diagram of the two first axes of the Canonical Correspondence Analysis (CCA) for species, relevés and environmental variables (triplot). Species are indicated with codes (see text); relevés with numbers (in red); and environmental variables with vectors: It: Thermicity index; Io: Ombrothermic index; Altitude (masl). Clusters obtained in the Hierarchical Cluster Analysis (CA): 1A, 1B, 1C, 2A, 2B, 2C, 3A, 3B, 4, 5A, 5B–5D; 5C and 6B. SMb: Basal mountain forest; SMa: High mountain forest; SMrib: Riparian mountain forest; BM: Mountain woodland

Table 4.5 Data extracted from the correlation matrix for the Canonical Correspondence Analysis: correlation coefficients between environmental variables, canonical axes of the species, and environmental axes estimated by the CCA Spec Ax1 Spec Ax2 Envi Ax1 Envi Ax2 Altitude It Io

Spec Ax1 1.0000 0.0234 0.9449 0.0000 0.8814
0.9175 0.898

Spec Ax2

Envi Ax1

Envi Ax2

Altitude

It

Io

1.0000 0.0000 0.8675
0.2985 0.1944 0.2417

1.0000 0.0000 0.9327
0.9709 0.9501

1.0000
0.3441 0.2241 0.2786

1.0000
0.9918 0.7752

1.0000
0.8483

1.0000

species in the ordination plan, and the ordination of the relevés by the means of these variables. Also located in the graph were the clusters obtained in the Hierarchical Cluster Analysis (CA), except Cluster 6A, which, as mentioned earlier, represents a more or less homogeneous cluster whose species and relevés are distributed more

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4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.6 Results of the ordination of the Canonical Correspondence Analysis (CCA) for the first two axes. The values of the characteristic roots are shown, along with the correlation between species and environmental variables, and percentage of variance explained by the species data and species-environment relations Eigenvalues Corr. species-env. var. Cum. variance (%) Species Species-env. var.

CCA1 0.384 0.945

CCA2 0.181 0.867

9.1 61.3

13.3 90.2

dispersedly in the ordination plan, and it is not therefore possible to delimit them. On the first two canonical axes on the ordination plan, the position of the species indicates an approximation to their ecological optimum, and the environmental variables are represented by “vectors” pointing in the direction of their variation; the direction and relative length of the vectors constitute their contribution to the ordination with regard to the distribution of the species in the same plan (Ter Braak 1986). Figure 4.4 shows that on axis 1 (61.3% of variance) the positive results are associated with high altitudes (r ¼ 0.9327), high Io (r ¼ 0.9501) and low It values (r ¼
0.9709), whereas the negative results are associated with high It and with lower altitudes and Io (Table 4.5). From these correlation values it can be deduced that axis 1 shows the floristic gradient generated in response to the existing environmental gradient, which is strongly determined by the altitude, ombrothermic index (Io) and thermicity index (It). This axis separates two environments within which the species are distributed according to their ecological optimums: a lower, warmer and subhumid environment, and another higher, more temperate and humid environment (Fig. 4.4). In turn, within these two environments, four sectors were delimited in the triplot that adjust the distribution of species to certain intervals in the environmental variables that are closely associated with axis 1: Sectors I and II were distinguished for lower, warmer and subhumid environments, and Sectors III and IV for higher, temperate and humid environments. Sector I (top left negative quadrant) and Sector IV (lower right positive quadrant) contain the most opposing and pronounced environmental conditions in the study area, whereas Sectors II and III have transitional and more attenuated conditions (Table 4.7 and Fig. 4.4). With regard to axis 2 (28.9% of variance), it was observed that the correlation coefficients did not reveal any significant values with the environmental variables (altitude, It and Io) (Table 4.5). Figure 4.1 shows that the positive results of this axis are associated to low values of It (r ¼ 0.2241), Io (r ¼ 0.2786) and altitude (r ¼
0.3441). This situation showed that the relation of axis 2 with these variables was not very significant and that other environmental variables could be associated with it, such as soil or geomorphological variables, which were not analysed quantitatively in this research. Specifically the only situation that showed a possible association of axis 2 with certain soil variables occurred due to the significant

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Table 4.7 Distribution sectors of species on the ordination plan, arranged from lower, warmer and subhumid to higher, temperate and humid. In blue: sectors with the most extreme environmental conditions Environmental variables Altitude It Io

Canonical axis 1 Sector I (
top left) 1015–1121 417–429 4.7–4.8

Sector II (
lower left) 1085–1275 399–422 4.8–5.5

Sector III (+ top right) 1180–1400 378–404 7.5–8.1

Sector IV (+ lower right) 1417–1620 352–376 8.1–8.7

presence on the positive side of the axis (Sector I) of certain indicator species of riparian environments such as Salix humboldtiana (Sali), Baccharis salicifolius (Bsal), Tessaria integrifolia (Tein), Tessaria dodoneifolia (Tedo), Asclepias curassavica (Ascl) and Paspalum distichum (Pasp) (Fig. 4.4). Additionally, with regard to the characteristics of the soils in the study area (Fig. 2.3a), no clear separation of the relevés was observed on the CCA ordination diagram according to the type of soil on which they were growing (Fig. 4.4). The positive quadrant of axis 2 clustered relevés located on the two soil types existing in the area: the relevés in Sector I (Clusters 1A, 2B, 2C and 6B) and some relevés in Sector III (Cluster 5C) are located on Haplic Phaeozem-type soils in the Palpalá association, whereas other relevés in Sector III (Cluster 5A and part of Cluster 5B– 5D) are located on calcareous Phaeozem-eutric Regosol-type soils in the SevenguialJordán River association. However, the negative quadrant of axis 2 may show a greater association with certain soil variables, as it clustered the relevés in Sector II (Clusters 1B, 1C, 2A, 3A and 3B) and Sector IV (Cluster 4 and another part of Cluster 5B–5D), all also located on calcareous Phaeozem-eutric Regosol-type soils in the Sevenguial-Jordán River association. In consequence, and on a provisional basis, axis 2 could be assumed to be associated to certain soil and geomorphological variables that are not considered quantitatively in the present study, but which could influence the distribution of some of the species studied.

4.3.3

Relationships Between Species and Environmental Variables Defined by Axis 1 on the CCA Ordination Diagram

The location of the species on the ordination diagram defined by axes 1 and 2 (Fig. 4.4) responds mainly to the gradient of altitude, It and Io defined by axis 1; this gradient determines the variations observed in the floristic composition and the distribution pattern of the plant communities in the study area. The distribution of the species and variables in the diagram defined by these axes can be interpreted as follows: the positive part of axis 1 (Sectors III and IV) is related

58

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

to the presence of vegetation types represented by semi-open to closed mainly mesowoodland growing at high altitudes and characterised by less thermophilous species with greater moisture requirements, whose distribution optimum is located under more temperate and humid conditions. As a group, these species showed a positive response to high values of altitude (1260–1620 m asl) and Io (7.7–8.7) and to low values of It (352–395) (Fig. 4.4). Notable among these species for their importance when differentiating plant communities on the ordination diagram are those located at the end of the environmental gradient (Sector IV) and which form part of the mountain woodland (Cabrera 1994), such as Podocarpus parlatorei (Podo), Sambucus nigra ssp. peruviana (Samb), Alnus acumuninata (Alnu), Ilex argentina (Ilex), Prunus tucumanensis (Prun), Calceolaria teucrioides (Calt), Berberis jobii (Berb), Lepchinia vesiculosa (Lepe), Austroeupatorium inulifolium (Aust), Ophryosporus lorentzii (Oplo), Campovassouria cruciata (Camp), Solanum aligerum (Soli) and Sibthorpia conspicua (Sibc). In humid areas at lower altitudes it is common to find other typical species of the mountain forest (“selva alta”) (Cabrera 1994) and of the ecotone areas (Sectors III and IV), such as Cedrela angustifolia, Blepharocalyx salicifolius (Blef), Duranta serratifolia (Dura), Cinnamomum porphyrium (Cinn), Myrcianthes pseudomato (Myps), Myrcianthes pungens (Mypu), Bougainvillea stipitata (Boug), Aralia soratensis (Aral), Cedrela saltensis (Cesa) and Chrysophyllum marginatum (Chry), all species in the tree layer; whereas in the shrub and herbaceous layer there are mainly Stevia yaconensis var. subeglandulosa (Stey), Solanum confusum (Scon), Duchesnea indica (Duch), Tibouchina paratropica (Tibo), Solanum betaceum (Sbet), Festuca hieronymi (Fehi), Clinopodium bolivianum (Clin), Koanophyllon solidaginoides (Koan), Collaea argentina (Coya), Rubus imperialis (Rubu), Heimia montana (Heim), Baccharis latifolia (Blat), Onoseris alata (Onos), Seemannia gymnostoma (Seem), Petunia occidentalis (Petu), Phenax laevigatus (Phen), Piper hieronymi (Pipe), Aphelandra hieronymi (Aphe), Justicia mandonii (Jusm), Cnidoscolus vitifolius (Cniv), Acalypha communis, Acalypha plicata (Apli), Agalinis genistifolia (Agal) and Phytolacca bogotensis (Phyt), among others. The negative side of axis 1 (Sectors I and II) is related to the presence of vegetation types represented by open to semi-open woodland with a significant proportion of micro-woodlands, located in low-lying areas of the mountain forest (“basal forest”) (Cabrera 1994) and characterised by thermophilous species that respond positively to high values of It (399–429) and to low values of Io (4.7–5.5) and altitude (1015–1275 m asl) (Fig. 4.4). The most important species on this side of axis 1 include Terminalia triflora (Term), Myroxylon peruiferum (Myro), Zanthoxylum petiolare (Zanp), Carica quercifolia (Carq), Enterolobium contortisiliquum (Ente), Tipuana tipu (Tipa), Acacia aroma (Acar), Sebastiania brasiliensis (Sebr), Celtis iguanaea (Ceig), Manihot grahami (Mani), Schinus bumeloides (Schb), Schinus fasciculatus (Schf), Xylosma pubescens (Xylo), Vassobia breviflora (Vaso), Jacaranda mimosifolia (Jaca), Prosopis alba (Poso), Condalia buxifolia (Cond), Chloroleucon tenuiflorum (Chlo), Geoffroea decorticans (Geof), Sebastiania commersoniana (Seco), Erythrina falcata (Eryt) and Acacia

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59

caven (Acac), all species in the tree layer; also Capsicum chacoense (Caps), Senna occidentalis (Seoc), Urera baccifera (Urer), Senna pendula var. eriocarpa (Sepe), Acalypha amblyodonta (Acaa) and Barnadesia odorata (Barn) in the shrub layer; and Elephantopus mollis (Elep), Mirabilis jalapa(Myra), Samolus valerandi (Samo), Jungia pauciflora (Jupa), Cantinoa mutabilis (Cant) and Modiolastrum malvifolium (Modi) in the herbaceous layer, among others. In addition, the species that are close to the central coordinates or above the zero value of axis 1 are more indifferent to the environmental variables considered here. This may be due to the fact that they have a broader tolerance range so they can occupy several environments (wide-habitat or eurioic species) or that they respond to the influence of other environmental variables that are not analysed in this study. Most of these species are broadly distributed throughout the Reserve, and include some that behave as dominant and contribute significantly to the physiognomy of the plant communities they integrate. Examples are tree species such as Parapiptadenia excelsa (Para), Anadenanthera colubrina var. cebil (Anad), Allophylus edulis (Allo) and Schinus gracilipes (Schg) (Fig. 4.4). Only a few indifferent species with regard to axis 1 have a distribution which is somewhat more restricted and show a preference for certain environments that tend to be more unstable or fluctuating (gorges, terraces, riparian environments or edges of woodland), as was demonstrated in the field and will be seen in the Discussion section. These include tree species such as Tecoma stans (Teco) and Trema micrantha (Trem). Finally, the species that are more isolated on the triplot are rare or scarce, with low presence indices, and may reflect very specific situations associated to more extreme ecological conditions; for example, the positive quadrant (top) of axis 1 contains Cordia saccelia (Cord), a rare species that is present in a single relevé and with a low phytosociological value (Cluster 5C). The use of Canonical Correspondence Analysis (CCA) allowed a more objective determination of clusters of species associated to particular ranges of environmental variables. The first axis essentially explains the abundance and distribution of the species along a gradient of altitude, Io and It, from lower, warmer and subhumid to higher, temperate and humid environments. However, axis 2 is probably related to soil and geomorphological variables that may affect the species distribution in some sectors. In summary, a close and significant relation was observed between species and environmental variables; however, some species were also recognised that did not have any significant association with them. This is the case of certain dominant species with a wide distribution and the ability to colonise different sites, and others whose distribution is more influenced by other variables, as mentioned earlier. The results show the importance of the environmental gradient as a determining factor in the variations in the floristic composition and the general distribution pattern of the plant communities, while the possible contribution that can be attributed to soil and geomorphological variables would play a secondary role in this pattern rather than determining it.

60

4.3.4

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

General Synthetic Comparative Table

The general synthetic comparative table (Appendix C) allowed a comparison of all the relevés that were summarised or synthesised in the presence columns. The top of the table shows the clusters obtained in the Hierarchical Cluster Analysis, ordered in increasing altitudinal intervals (1A, 2B, 2C, 2A, 1C, 3A, 3B, 1B, 5C, 5A, 5B–5D and 4), except cluster 6B (riparian), which is located at the end of the tables. The columns show the presence indices of the species in each cluster indicated with Roman numerals (except in the clusters with less than six where they appear in Arabic numbers). The plant associations proposed a posteriori are also similarly numbered. According to the criteria in Table 4.4, various species with a diagnostic value were selected under the name of “characteristic or indicator” species (Appendix C; Fig. 4.5), namely:

Fig. 4.5 Canonical Correspondence Analysis (CCA) for the species-environmental variables: Characteristic or indicator species (circled in red) of the clusters obtained in the Hierarchical Cluster Analysis (CA: 1A, 1B, 1C, 2A, 2B, 2C, 3A, 3B, 4, 5A, 5B–5D, 5C and 6B). SMb: Basal mountain forest; SMa: High mountain forest; SMrib: Riparian mountain forest; BM: Mountain woodland

4.3 Vegetation

61

– species with a high degree of fidelity (exclusive species) present in a single cluster with a high presence index (V), e.g. Sambucus nigra subsp. peruviana (Samb) and Manihot grahami (Mani); or species present in several clusters and thus with a less restrictive distribution, but with high or very significant phytosociological values and presence indices in a particular cluster; that is, with a distribution optimum in a specific cluster (at most in two clusters related territorially or ecologically) (selective and preferential species), some of which also behave as dominant, e.g. Juglans australis (Jugl) and Cinnamomum porphyrium (Cinn). In the CCA ordination diagram, these species were located on the cloud formed by the cluster of relevés they characterise, or very near to it. Most of these species showed a significant association with the environmental variables, except for some that were more indifferent, such as Tecoma stans (Teco) and Trema micrantha (Trem), which, as mentioned earlier, also tend to appear in more unstable environments and are probably subject to the influence of other variables (soil or geomorphological) (Appendix C; Fig. 4.5). – species that behaved as dominant in one or several clusters of relevés with high or significant phytosociological values; that is, species with a broad distribution and which contribute significantly to the physiognomy of the vegetation. These species showed two types of behaviour: some had an optimal association with the environmental variables, while others were indifferent. In a very few cases there were dominant species with a very restricted distribution. The best example is Podocarpus parlatorei (Podo) (Appendix C; Fig. 4.5). – differential species that occurred in one single cluster and with only presence indices II, III and IV (not I or V), as they were better able to separate two clusters of relevés (Alcaraz Ariza 2013) (Appendix C; Fig. 4.5). – stenoic species, restricted to certain environments and indicating specific ecological conditions. These species did not significantly respond to the environmental variables and reflect the influence of specific soil and geomorphological factors. For example, Salix humboldtiana (Sali), Tessaria dodoneifolia (Tedo) and Tessaria integrifolia (Tein), among others, are typical species of edaphohygrophilous vegetation associated to riparian environments or with shallow water tables (Appendix C; Fig. 4.5). – endemic species, e.g. Schinus gracilipes (Schg) and Stevia yaconensis var. subeglandulosa (Stey), among others; it is worth mentioning that of the total endemic species recorded (25), ten were characteristic. – species associated to dynamic or fluctuating conditions, e.g. Acacia aroma (Acar); or else relevant in biogeographical terms, e.g. Geoffroea decorticans (Geof) and Prosopis alba (Poso) which belong to the biogeographical region of El Chaco and tend to appear in disjointed enclaves in subhumid areas of Las Yungas; all have significant phytosociological values and presence indices, either dominant or not, and with varying degrees of association with the environmental variables.

62

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

It should be noted that not all the species that were associated to a greater or lesser extent with the environmental variables were selected as characteristic or indicator species, due to the fact that their phytosociological values and presence indices were not significant enough to assign them a diagnostic value for the cluster, regardless of whether they were found in many clusters or merely restricted to one or only a few. Thus the general synthetic comparative table (Table 3.4) offers a vision of the whole of the vegetation studied, grouped into 13 clusters of relevés, each formed by a combination of characteristic or indicator species, and by companion species without any significant diagnostic value (companion species). Specifically, Cluster 6A was not incorporated in this table as it did not have a characteristic combination of species that could be attributed to a community with its own identity.

4.3.5

Delimitation of the Plant Communities

The analysis of the floristic-phytosociological results obtained from the ordinated phytosociological matrix enabled the comparison of the abundance-dominance and frequency of these species in each relevé. In turn, the Hierarchical Cluster Analysis (CA) (Fig. 4.3) allowed the delimitation of clusters of homogeneous relevés to compare them in terms of their presence indices (Appendix C). Finally, the statistical processing of the species data and environmental variables by the Canonical Correspondence Analysis (CCA) provided valuable support when selecting the characteristic or indicator species in each cluster; for each one the combination of taxa was obtained that best expressed its relationship with the environment and revealed the gradient established in the study area (Fig. 4.5). Based on the comparison of all the results it was possible to establish and formally delimit the plant communities growing in the study area, which correspond to the clusters obtained in the Hierarchical Cluster Analysis (CA). These communities adequately reveal the current diversity and the floristic and phytosociological relations between them. As a result of this, 13 communities were recognised and described for the first time for the province of Jujuy (Appendix C; Fig. 4.6); these communities were typified from the phytosociological standpoint as new associations with a provisional character (except Association 12 which has already been described for Bolivia): eight (8) belong to woodlands growing in the lower Mesotropical-lower and higher Subhumid bioclimatic belt (Associations 1–8); four (4) correspond to woodlands in the upper Mesotropical-lower Humid bioclimatic belt (Associations 9–12); and one (1) belongs to humid-subhumid riparian woodlands in the lower Mesotropical belt (Association 13). Each of these associations is supported by the corresponding table, correlatively numbered (Tables 4.8–4.20). Only Cluster 6A could not be attributed to any community; this cluster has six relevés located at

4.3 Vegetation

63

Fig. 4.6 Canonical Correspondence Analysis (CCA) of the species-environmental variables: Vegetation composition and distribution in relation to the environmental gradient. This shows the five groups that cluster the associations in the subtropical mountain woodland in the Serranías de Zapla Multiple Use Ecology Reserve. Associations: 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 and 13. SMb: Basal mountain forest; SMa: High mountain forest; SMrib: Riparian mountain forest; BM: Mountain woodland

different altitudes on gorges and in steep humid shady ravines where there may even be small springs. The cluster reveals a herbaceous and shrubby layer with some species with high presence indices; they are fairly restricted to the cluster and contribute significantly to the physiognomy of the vegetation, giving it the appearance of a grassland, such as Agalinis genistifolia (Agal), Collaea argentina (Coya), Cortaderia selloana (Cors), Cortaderia hieronymi (Corh), Festuca hieronymi (Fehi), Begonia boliviensis var. boliviensis (Begb) and Seemannia gymnostoma (Seem). However, it has a species-rich but very disperse and sparse tree layer, which also plays a role in the physiognomy. These particularities hindered the understanding and interpretation of the cluster, and it was therefore decided not to attribute it to a particular community until more thorough sampling campaigns can be done to enhance the knowledge of the ecology and phytosociology of the species that form it and make it possible to determine whether this is a vegetation type with its own identity and phytosociological rank.

64

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.8 Enterolobio contortisilici-Anadenantheretum cebilis ass. nova (Juglandi australisPhoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper tree layer (canopy) Parapiptadenia excelsa Anadenanthera colubrina var. cebil Tipuana tipu Enterolobium contortisiliquum Myroxylon peruiferum Terminalia triflora Erythrina falcata Cedrela angustifolia Cinnamomum porphyrium Lower tree layer: Sebastiania brasiliensis Acacia aroma Sebastiania commersoniana Vassobia breviflora Celtis iguanaea Xylosma pubescens Schinus bumeloidesa Sapium haematospermum Zanthoxylum petiolare Carica quercifolia Acacia caven Allophylus edulis Scutia buxifolia Myrsine laetevirens Morus alba Schinus fasciculatus Solanum riparium Tecoma stans Bougainvillea stipitata Randia micrantha Trema micrantha Scrub layer Urera baccifera Barnadesia odorata

103 100 66 53 1

103 100 58 54 2

103 100 36 56 3

104 100 39 55 4

104 100 34 57 5

104 100 27 60 6

104 100 33 59 7

104 100 38 58 8

9

3 3 2 2 . . . . .

4 3 2 2 . . 1 . .

3 4 2 . 1 . . . .

4 3 1 2 . . . . 1

4 4 1 1 1 1 1 . .

3 3 1 1 1 1 1 1 .

4 3 1 1 1 1 1 1 .

4 3 1 . 1 1 . 1 .

V V V IV IV III III II I

3 4 2 3 3 2 1 2 1 2 2 . 1 . 1 1 1 1 . 1 1

3 4 3 2 2 2 2 2 . 2 2 1 1 . 1 1 1 1 . . .

4 4 3 3 3 2 2 1 1 2 2 . 1 . . . . . . . .

4 3 3 3 3 2 2 2 . 2 . 1 . . . . . . 1 . .

3 3 3 3 2 3 1 2 2 . . 1 . 1 . . . . . . .

3 2 3 3 3 3 2 2 2 . . . . . . . . . . . .

3 2 3 3 3 3 2 . 2 . 2 . . 1 . . . . . . .

3 3 3 3 3 3 1 2 2 . . . . . . . . . . . .

V V V V V V V V IV III III II II II II II II II I I I

2 2

3 1

2 2

3 1

2 2

3 1

3 1

3 1

V V

(continued)

4.3 Vegetation

65

Table 4.8 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Solanum lorentzii Vernonanthura squamulosa Cestrum parqui Pavonia sepium Verbesina macrophylla var. nelidae Clematis haenkeana Baccharis capitalensis Chamissoa altissima Mimosa polycarpa Dendrophorbium bomanii Senna pendula var. eriocarpaa Lantana trifoliaa Dolichandra ungis-cati Rubus imperialis Boehmeria caudata Phenax laevigatus Smilax campestris Verbesina suncho Acalypha plicata Herb layer Rivinia humilis Solanum aloysiifolium Elephantopus mollis Justicia goudotii Petiveria alliacea Tagetes terniflora Sida cabreriana Mikania micrantha Ruellia erythropus Galinsoga caracasana Bromelia serra Fleischmannia schickendantzii Bidens subalternans Orthopappus angustifolius Parthenium hysteriophorus Praxelis clematidea

103 100 66 53 1 2 1 1 . 1 . . 1 1 1 1 1 . . . 1 . 1 1

103 100 58 54 2 1 1 . + 1 . . . . 1 1 . . 1 1 1 1 1 1

103 100 36 56 3 1 1 1 . . . 1 . . 1 1 . + 1 1 . . . .

104 100 39 55 4 2 1 . + 1 1 . 1 1 . . . . . . . 1 . .

104 100 34 57 5 1 1 1 + 1 1 . . 1 . . . . . . . . . .

104 100 27 60 6 1 1 1 + . . . . . . . . . . . . . . .

104 100 33 59 7 1 1 1 + . . 1 . . . . . . . . . . . .

104 100 38 58 8 1 1 . . . 1 1 1 . . . 1 + . . . 1 . .

9 V V IV IV III II II II II II II II II II II II II II I

3 1 2 + + + 1 + + . 3 1 . + + +

4 2 2 1 + + 1 + + + 2 1 . . . .

3 3 2 . + + 1 . . . . . . . + .

4 1 2 + . . . . + + 2 . + + . .

3 2 2 . . . . . + . . . . . + +

3 2 2 . . . . + . . . . . . . .

3 2 . + + + . . . + . . + . . +

3 2 2 + + + 1 + . + . 1 + + . .

V V V IV IV IV III III III III II II II II II II

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

66 Table 4.8 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Chaptalia nutans Acalypha communis Duchesnea indica Verbena litoralis Tradescantia boliviana Salvia personata Anredera cordifolia Jungia pauciflora Pharus lappulaceus Tragia volubilis

103 100 66 53 1 . 1 + + + + + + + +

103 100 58 54 2 + 1 + + + . . . . .

103 100 36 56 3 . . . . . + . . . .

104 100 39 55 4 . . . . . . + + + +

104 100 34 57 5 + . . . . . . . . .

104 100 27 60 6 . . . . . . . . . .

104 100 33 59 7 . . . . . . . . . .

104 100 38 58 8 . . . . . . . . . .

9 II II II II II II II II II II

Other species: All species I in 9 (Syn.). Scrub layer: Cnidoscolus tubulosus + in 1; Carica glandulosa and Hebanthe occidentalis 1 in 2. Herb layer: Dicliptera squarrosa 1, Modiolastrum malvifolium, Leonurus japonicus, Cuphea racemosa, Cenchrus latifolius and Samolus valerandi + in 1; Adenostemma brasilianum, Desmodium affine, Mirabilis jalapa and Phytolacca bogotensis + in 2; Cantinoa mutabilis + in 3; Nicandra physalodes + in 8. Holotypus ass.: Relevé 5 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 9 is synthetic

4.3.6

Nomenclature of the Communities

The available information on the phytosociology of subtropical mountain woodlands in the province of Jujuy is still very scarce, and there are currently only two previous studies (Martin 2014; Haagen Entrocassi 2014); the same occurs in regard to the determination of the nomenclature and syntaxonomical position of the communities within these woodlands. The closest studies in the geographic literature are found in the work conducted in Bolivia by Navarro and Maldonado (2002). In this regard and given the complexity of this plant formation due to its species diversity, communities and layers, it is difficult to decide on a definitive nomenclature for the communities established in the present study. For this reason, and in order to contribute to future studies that extend the current floristic-phytosociological information and allow a syntaxonomical checklist to be compiled for these woodlands, the following associations are proposed:

4.3 Vegetation

67

Table 4.9 Schino bumeloidis-Allophyletum edulis ass. nova (Juglandi australis-Phoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Canopy layer Allophylus edulis Acacia aroma Vassobia breviflora Xylosma pubescens Condalia buxifoliab Schinus bumeloidesa Celtis iguanaea Sebastiania brasiliensis Chloroleucon tenuiflorum Manihot grahami Acacia caven Cnicothamnus lorentzii Sebastiania commersoniana Senna spectabilis Jacaranda mimosifolia Geoffroea decorticansb Eucalyptus sp. Celtis ehrenbergiana var. discolora Schinus gracilipesa Zanthoxylum petiolare Sapium haematospermum Seral species Enterolobium contortisiliquum Parapiptadenia excelsa Anadenanthera colubrina var. cebil Tipuana tipu Blepharocalyx salicifolius Erythrina falcata Scrub layer Urera baccifera Chamissoa altissima Vernonanthura squamulosa

109 100 56 61 1

109 100 62 62 2

109 100 43 65 3

109 100 47 63 4

109 100 53 64 5

109 100 35 66 6

109 100 39 67 7

110 100 44 68 8

110 100 46 69 9

110 100 51 70 10

11

4 3 3 3 2 3 1 1 1 1 1 1 . 1 . . . .

4 3 3 3 2 3 2 2 2 1 . 2 1 1 1 . 1 1

3 2 3 3 3 3 2 2 2 2 2 . . 1 . . . .

4 3 3 2 2 2 2 1 1 . 1 1 . . 1 1 . .

4 3 2 3 3 3 2 1 2 1 2 . 1 . . . . 1

4 2 3 3 3 3 2 2 1 1 1 1 . . . . 1 .

3 3 3 2 2 2 2 2 2 2 . . 1 1 . . . .

4 3 2 2 3 2 1 2 2 2 2 . 1 . . . . .

4 3 2 2 3 2 2 1 1 1 2 1 1 1 1 1 1 .

3 3 3 3 3 3 2 1 . 1 2 1 1 1 1 1 . .

V V V V V V V V V V IV III III III II II II I

1 . .

. 1 1

1 . .

. . .

. . .

. . .

. . .

. . .

. . .

. . .

I I I

1 1 .

1 . 1

2 . 1

. 2 1

1 2 .

. 1 .

2 1 .

2 1 .

1 . .

. 1 1

IV IV II

. . .

1 1 1

. 1 .

. . .

1 . .

1 . .

. 1 1

. . .

. . .

. . .

II II I

3 1 1

3 1 1

3 1 .

3 . 1

2 1 1

3 2 1

2 1 .

2 1 1

3 1 1

2 1 1

V V IV

(continued)

68

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.9 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Dendrophorbium bomanii Smilax campestris Barnadesia odorata Cestrum parqui Senecio rudbeckiifolius Capsicum chacoense Lantana canescensa Clematis haenkeana Senna pendula var. eriocarpaa Mimosa debilis Muehlenbeckia sagittifolia Rubus imperialis Sida rhombifolia Baccharis microdonta Baccharis capitalensis Croton saltensisa Acalypha amblyodonta Budleja iresinoides Solanum lorentzii Malvastrum coromandelianum Buddleja stachyoides Carica glandulosa Senna occidentalis Herb layer Elephantopus mollis Mirabilis jalapa Praxelis clematidea Samolus valerandi Modiolastrum malvifolium Parthenium hysteriophorus Tagetes terniflora Justicia goudotii Fleischmannia schickendantzii Jungia pauciflora Verbena litoralis Hypochaeris microcephala

109 100 56 61 1 1 + 1 . 1 + 1 1 1 + . + . . 1 1 . . . 1 1 1 .

109 100 62 62 2 1 . . 1 1 . . . 1 . + . + . . 1 1 . 1 . . . .

109 100 43 65 3 . . 2 2 1 . . . 1 . + . . . 1 . . . 1 . + . .

109 100 47 63 4 . + 1 . . + 1 1 . + + + . 1 1 . . 1 . 1 + . .

109 100 53 64 5 1 + . 1 1 + . 1 1 + . + + 1 . 1 1 1 . . . . .

109 100 35 66 6 1 + . . . . . . . . . . . . . 1 . . . . . . 1

109 100 39 67 7 1 . 1 1 1 + 1 . 1 . . + + 1 . . 1 . 1 . . 1 .

110 100 44 68 8 . + 1 1 . . 1 1 . + + . . . 1 . . 1 . . + . 1

110 100 46 69 9 1 + . . . + . 1 . . . + + . . . . . . . . . .

110 100 51 70 10 1 + 1 1 1 + 1 . . + + . + 1 . . . . . 1 . . .

11 IV IV III III III III III III III III III III III II II II II II II II II I I

3 2 1 2 + + + 1 . 1 + .

2 3 1 3 + + . 1 1 1 + 1

3 3 1 3 + + . . 1 1 . 1

3 3 1 . . . + 1 1 . . .

3 3 . 3 + + + . . 1 + 1

2 2 1 . + . . . . . . .

3 . 1 3 . . + . 1 . + .

3 2 1 1 + + + 1 1 1 . .

2 2 1 2 . + + 1 1 . + 1

2 3 1 2 + + + 1 . 1 + 1

V IV V IV IV IV IV III III III III III

(continued)

4.3 Vegetation

69

Table 4.9 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Tradescantia boliviana Salpichroa origanifolia Gamochaeta pensylvanica Viguiera tucumanensis var. oligodonta Rivinia humilis Bidens pilosa Bromus catharticus Cantinoa mutabilis Desmodium subsericeum Oenothera roseaa Galinsoga caracasana Leonurus japonicus Mimosa xanthocentra Cajophora hibiscifolia Plantago australis Zinnia peruviana Glandularia tweedieana Bidens subalternans Petiveria alliacea Axonopus compressus Pseudechinolaena polystachya Eleusine indica Ruellia ciliatiflora Digitaria insularis Solanum tenuispinuma Anredera cordifolia Sida cabreriana

109 100 56 61 1 1 + + .

109 100 62 62 2 1 + . .

109 100 43 65 3 1 . . .

109 100 47 63 4 . . + .

109 100 53 64 5 1 . . 1

109 100 35 66 6 1 . + .

109 100 39 67 7 . . . 1

110 100 44 68 8 . + . 1

110 100 46 69 9 . + + 1

110 100 51 70 10 . 1 + .

11 III III III II

+ + + + . . . . . . . . . . + . . . . + . . +

. + + . + + . . + + . + . + + . . . + . 1 . +

+ + + . . + . . . . + + . . . . . . . . . + .

. . . + . . . + . . + . . . + + + + . . . + .

+ + . . + . + . . + . . + . . . . . . + 1 . .

. . . + . + . . . . . . + . . + + + + . . . .

. . . . . . . . . . . . . + . . . . . . . . .

+ . . . + . . + . . . . + . . . . . . . . . .

. . . . + + + . + . + + . . . . . . . . . . .

. . + + . . + + + + . . . . . . . . . . . . .

II II II II II II II II II II II II II I II I I I I I I I I

Other species: All I in 11. Scrub layer: Baccharis coridifolia 1, Hebanthe occidentalis + in 1; Jungia polita 1 in 4. Herb layer: Cuphea racemosa and Salvia personata + in 1; Mikania micrantha + in 4; Solanum sisymbriifolium + in 5. Holotypus ass.: Relevé 2 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec.; relevé 11 is synthetic

70

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.10 Xylosmo pubescentis-Blepharocalycetum salicifolii ass. nova (Juglandi australisPhoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper canopy layer Blepharocalyx salicifolius Parapiptadenia excelsa Erythrina falcata Tipuana tipu Lower canopy layer Sebastiania brasiliensis Scutia buxifolia Xylosma pubescens Allophylus edulis Condalia buxifoliab Sebastiania commersoniana Celtis ehrenbergiana var. discolora Schinus bumeloidesa Celtis iguanaea Jacaranda mimosifolia Vassobia breviflora Acacia aroma Zanthoxylum petiolare Chloroleucon tenuiflorum Geoffroea decorticansb Sapium haematospermum Senna spectabilis Eucalyptus sp. Scrub layer Urera baccifera Capsicum chacoense Cestrum parqui Senna occidentalis Baccharis capitalensis Barnadesia odorata

111 100 38 71 1

111 100 52 72 2

111 100 28 80 3

111 100 51 73 4

111 100 40 74 5

112 100 27 75 6

112 100 31 79 7

112 100 30 76 8

112 100 32 77 9

113 100 33 78 10

11

4

4

4

3

3

4

4

4

4

4

V

2 . .

2 . 1

1 . .

1 . .

2 . .

. . .

1 1 .

2 1 .

2 . .

1 . .

V I I

3 3 2 2 1 .

3 3 2 2 1 1

3 3 2 2 1 2

4 3 2 2 1 .

4 3 2 1 . 1

4 3 3 2 1 1

4 3 2 2 1 2

4 2 3 2 1 .

4 2 2 2 1 1

4 3 2 2 . 1

V V V V IV IV

1

1

.

.

1

1

1

1

.

1

IV

1 . 1 . . . .

. . . 1 . 1 .

1 1 1 . . 1 .

1 1 . . 1 . 1

1 1 1 1 . . .

. . . . . . .

1 1 . . 1 . .

1 . . . 1 . .

1 1 . 1 . . .

. . . . . 1 1

IV III II II II II I

. .

. 1

. .

1 .

. .

. .

. .

1 .

. .

. 1

I I

. 1

1 .

. .

. .

. .

. .

. .

1 .

. .

. .

I I

3 + . 1 . .

3 + . . 1 1

2 + 1 . 1 1

1 . 1 1 1 .

3 + . 1 . 1

1 + 1 . 1 1

3 + 1 1 . 1

2 . . . . .

3 + 1 1 . .

3 . 1 1 1 .

V IV III III III III

(continued)

4.3 Vegetation

71

Table 4.10 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Dendrophorbium bomanii Senecio rudbeckiifolius Senna pendula var. eriocarpaa Rubus imperialis Baccharis microdonta Chamissoa altissima Heimia montanaa Lantana canescensa Solanum lorentzii Vernonanthura squamulosa Acalypha amblyodonta Carica glandulosa Weddelia saltensis Buddleja stachyoides Hebanthe occidentalis Budleja iresinoides Clematis haenkeana Pavonia sepium Heteropterys sylvatica Muehlenbeckia sagittifolia Ophryosporus piquerioides Smilax campestris Mimosa debilis Herb layer Mirabilis jalapa Elephantopus mollis Parthenium hysteriophorus Petiveria alliacea Cantinoa mutabilis Oenothera roseaa Tagetes terniflora Verbena litoralis

111 100 38 71 1 1

111 100 52 72 2 1

111 100 28 80 3 .

111 100 51 73 4 .

111 100 40 74 5 1

112 100 27 75 6 .

112 100 31 79 7 .

112 100 30 76 8 1

112 100 32 77 9 1

113 100 33 78 10 .

11 III

. 1

1 .

1 .

1 1

. .

. 1

. 1

1 .

. 1

1 .

III III

1 . . 1 1 1 .

. 1 1 . . . 1

. 1 . . . . .

1 . 1 1 1 1 .

. 1 . . 1 . 1

. . 1 1 . . .

+ . . . . . .

+ . . 1 . 1 1

. . 1 . 1 1 1

+ 1 . . . . .

III II II II II II II

. . 1 + . 1 . 1 + .

1 1 . . + 1 1 . . +

. . . . . . . . . .

. . 1 . . . . 1 . +

1 1 1 + + . 1 . + .

. . . . . . . . . .

. . . + + . . . . .

. . . . . . . . . .

. . . . . . . . . .

1 1 . . . . . . . .

II II II II II I I I I I

.

+

.

+

.

.

.

.

.

.

I

. .

. +

. .

+ .

. .

. .

. .

. .

+ .

. .

I I

1 1 +

. 1 +

1 . +

1 . +

1 1 .

. . +

1 1 +

1 1 .

1 . .

1 1 .

IV III III

+ + . + .

1 . + . .

. . . + .

1 . + . +

. + . + +

. . . . +

. + + + +

+ . . + .

. + + . +

+ + + . .

III III III III III

(continued)

72

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.10 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Justicia goudotii Praxelis clematidea Viguiera tucumanensis var. oligodonta Fleischmannia schickendantzii Solanum tenuispinuma Anredera cordifolia Bromus catharticus Galinsoga caracasana Mikania micrantha Salpichroa origanifolia Axonopus compressus Cajophora hibiscifolia Cuphea racemosa Desmodium subsericeum Gamochaeta pensylvanica Glandularia tweedieana Jungia pauciflora Rivinia humilis Ruellia ciliatiflora Dicliptera squarrosa Salvia personata Solanum aloysiifolium Bidens pilosa Eleusine indica Panicum trichanthum Plantago australis Sida cabreriana

111 100 38 71 1 . . .

111 100 52 72 2 1 . 1

111 100 28 80 3 . . 1

111 100 51 73 4 1 1 .

111 100 40 74 5 . 1 1

112 100 27 75 6 1 . .

112 100 31 79 7 . 1 .

112 100 30 76 8 . 1 1

112 100 32 77 9 1 . .

113 100 33 78 10 . . .

11 II II II

.

1

1

.

1

.

.

.

.

.

II

1 . + . + . . . . . .

. + . + + + + . + . .

. . + + . . . + . . .

. . + + . + + + . . .

1 . . . . . . . . + +

. + . . . + . + + + .

. + . . . . . . . . .

. . . . + . + . . . .

. + . . . . . . . + +

1 . + + + + . . + . +

II II II II II II II II II II II

. . + . . 1 1 . . . . .

+ + + . 1 . 1 . + + + .

. + . . . . . . . . . .

+ . + + . 1 . + + + + +

. . . . 1 . . . . . . .

+ . . + . . . . . . . .

. . . . . . . . . . . .

. . . . . . . + . . . +

. . . + . . . . . . . .

. + . . . . . . . . . .

II II II II I I I I I I I I

Other species: All species I in 11. Herb layer: Digitaria insularis + in 1; Modiolastrum malvifolium and Pseudechinolaena polystachya + in 4; Bidens subalternans + in 5; Leonurus japonicus + in 6; Mimosa xanthocentra + in 9. Holotypus ass.: Relevé 8 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec.; relevé 11 is synthetic

4.3 Vegetation

73

Table 4.11 Jacarando mimosifoliae-Vassobietum breviflorae ass. nova (Juglandi australisPhoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Dominant tree layer Vassobia breviflora Celtis iguanaea Acacia aroma Schinus fasciculatus Jacaranda mimosifolia Xylosma pubescens Allophylus edulis Chloroleucon tenuiflorum Sebastiania commersoniana Condalia buxifoliab Sebastiania brasiliensis Acacia caven Geoffroea decorticansb Schinus bumeloidesa Tecoma stans Prosopis albab Senna spectabilis Seral species Enterolobium contortisiliquum Erythrina falcata Tipuana tipu Parapiptadenia excelsa Anadenanthera colubrina var. cebil Eucalyptus sp. Scrub layer Barnadesia odorata Dendrophorbium bomanii Cestrum parqui Mimosa polycarpa Vernonanthura squamulosa Solanum lorentzii Baccharis capitalensis Baccharis microdonta Buddleja stachyoides Budleja diffusa Iresine diffusa Jungia polita

119 100 44 81 1

120 100 46 82 2

120 100 39 83 3

120 100 40 84 4

112 100 36 85 5

120 100 39 86 6

7

4 4 4 2 2 2 2 1 1 . 1 . . . . . .

4 4 3 3 2 1 2 2 1 . 1 2 . . 1 . .

4 4 4 2 2 2 2 1 1 1 1 1 . . 1 . .

4 4 4 3 2 2 1 2 1 . . . 1 1 . . 1

4 4 4 3 2 2 2 1 1 1 . . . . . . .

4 4 4 2 2 2 1 1 1 1 . . 1 1 . 1 .

V V V V V V V V V III III II II II II I I

1 1 . 1 1 .

2 1 . . . .

2 . 1 . . .

1 . . . . .

2 . . . . 1

1 . . . . .

V II I I I I

3 2 1 . 1 1 . 1 . 1 1 .

2 2 1 1 . 1 1 1 1 1 . 1

3 1 2 1 1 . 1 . 1 . 1 1

2 2 1 2 1 1 . 1 . 1 1 .

3 2 1 2 1 1 1 . 1 . . 1

3 2 1 1 1 . 1 1 . . . .

V V V V V IV IV IV III III III III (continued)

74

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.11 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Malvastrum coromandelianum Rubus imperialis Senna pendula var. eriocarpaa Sida rhombifolia Trixis grisebachii Smilax campestris Clematis haenkeana Acalypha amblyodonta Vernonanthura pinguis Herb layer Leonurus japonicus Modiolastrum malvifolium Parthenium hysteriophorus Mikania micrantha Jungia pauciflora Mimosa xanthocentra Bromus catharticus Cantinoa mutabilis Bidens pilosa Bidens subalternans Conyza tunariensis Cuphea racemosa Eleusine indica Glandularia tweedieana Nicandra physalodes Praxelis clematidea Solanum tenuispinuma Anagallis arvensis Galinsoga caracasana Gamochaeta pensylvanica Zinnia peruviana Verbena litoralis

119 100 44 81 1 1 1 1 . 1 + . . .

120 100 46 82 2 . . 1 1 . . + 1 1

120 100 39 83 3 1 1 . . . . . . 1

120 100 40 84 4 . . 1 . 1 . . 1 .

112 100 36 85 5 1 . . 1 . + + . .

120 100 39 86 6 . 1 . 1 . + + . .

7 III III III III II III III II II

+ 1 1 + 1 . + + . + + + . . . . 1 . + + . .

+ . 1 + 1 1 + . . . . . + + + + 1 . . . . .

+ 1 . . . . . + . + . + + + . + . . + + . +

+ 1 1 + 1 1 + + + + + + . . + . . + . . + .

+ 1 1 + . 1 . + + . . . + + . + . . . . . .

+ 1 1 + 1 1 + . + . + . . . + . . + . . + +

V V V V IV IV IV IV III III III III III III III III II II II II II II

Other species: All species I in 7. Scrub layer: Chamissoa altissima and Senna occidentalis 1 in 1 Solanum palinacanthuma 1 in 2; Capsicum chacoense + in 5; Collaea argentina 1 in 6. Herb layer: Desmodium affine and Justicia goudotii + in 1; Anredera cordifolia, Digitaria insularis, Mirabilis jalapa, Orthopappus angustifolius and Rivinia humilis + in 2; Solanum sisymbriifolium + in 3. Holotypus ass.: Relevé 4 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec.; relevé 7 is synthetic

4.3 Vegetation

75

Table 4.12 Erythrino falcatae-Tipuanetum tipi ass. nova (Juglandi australis-Phoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1 = 10 m asl) Area (1 = 10 m2) N. species Ref. N. Relevé N. Upper canopy layer Tipuana tipu Erythrina falcata Parapiptadenia excelsa Anadenanthera colubrina var. cebil Blepharocalyx salicifolius Juglans australis Enterolobium contortisiliquum Cinnamomum porphyrium Cedrela angustifolia Lower canopy layer Acacia aroma Sebastiania brasiliensis Allophylus edulis Celtis iguanaea Condalia buxifoliab Sapium haematospermum Xylosma pubescens Tecoma stans Acacia caven Sebastiania commersoniana Chloroleucon tenuiflorum Vassobia breviflora Schinus myrtifolius Senna spectabilis Celtis ehrenbergiana var. discolora Zanthoxylum coco Schinus gracilipesa

112 100 47 106 1

121 100 59 1 2

122 100 58 2 3

122 100 52 3 4

122 100 57 4 5

123 100 39 22 6

123 100 44 5 7

124 100 54 21 8

125 100 62 6 9

125 100 53 7 10

126 100 55 20 11

126 100 46 8 12

13

1 1 3

1 1 1

2 2 1

2 1 .

1 2 1

1 1 1

2 . .

2 . .

2 1 1

1 1 1

. 1 .

1 . 1

V IV IV

.

1

.

1

.

3

1

4

.

.

3

1

III

.

1

.

1

.

2

.

4

1

.

5

1

III

. .

1 1

. .

. .

1 .

2 .

. 1

. .

1 .

. .

. .

1 1

III II

.

1

.

1

.

.

.

.

.

1

1

.

II

.

.

.

.

.

.

.

.

1

.

.

.

I

2 2

2 4

2 4

1 5

2 5

2 .

2 5

1 3

1 3

2 4

1 .

1 4

V V

1 . . .

3 3 2 1

3 3 2 1

4 2 2 1

4 3 3 2

. . . 1

4 3 2 1

. 1 1 .

4 2 3 2

3 2 3 2

1 1 . 1

3 2 3 .

V V IV IV

. 4 . .

2 1 1 2

3 . . 1

2 2 1 1

2 2 1 .

. . 1 .

2 1 . 2

. 1 1 1

3 . 1 1

3 1 1 1

. . . 1

3 1 2 .

IV IV IV IV

.

1

1

2

1

.

.

1

2

2

.

.

III

. . . .

2 . 2 1

1 2 . .

. 2 2 .

2 1 . 1

. . . .

2 . 1 1

. . 1 1

. 2 1 2

1 2 . .

1 1 . .

1 . 2 2

III III III III

. .

1 2

. 1

1 .

. 1

. .

. 1

1 .

1 1

1 .

. .

1 .

III III

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

76 Table 4.12 (continued) Altitude (1 = 10 m asl) Area (1 = 10 m2) N. species Ref. N. Relevé N. Scutia buxifolia Escallonia millegrana Myrcianthes pseudomato Jacaranda mimosifolia Carica quercifolia Citrus sp. Geoffroea decorticansb Randia micrantha Myrsine laetevirens Solanum riparium Morus alba Cnicothamnus lorentzii Prosopis albab Scrub layer Urera baccifera Barnadesia odorata Vernonanthura squamulosa Jungia polita Vernonanthura pinguis Acalypha amblyodonta Senecio rudbeckiifolius Smilax campestris Sida rhombifolia Mimosa debilis Chamissoa altissima Buddleja stachyoides Cestrum parqui Dolichandra ungiscati Baccharis coridifolia

112 100 47 106 1 . . .

121 100 59 1 2 1 . .

122 100 58 2 3 . . .

122 100 52 3 4 . . .

122 100 57 4 5 1 . .

123 100 39 22 6 . 1 .

123 100 44 5 7 . . .

124 100 54 21 8 . 1 .

125 100 62 6 9 . . 1

125 100 53 7 10 1 1 .

126 100 55 20 11 . . .

126 100 46 8 12 . . 2

13 III II I

.

.

1

.

.

.

.

.

1

.

.

.

I

. . .

. . .

1 1 .

. . 1

. 1 .

. . .

. . .

. . .

. . .

. . 1

. . .

1 . .

I I I

. . . . .

. . . . .

. 1 . . .

. . . . .

1 . . 1 .

. . . . 1

. . . . .

. . . . .

. 1 1 . .

1 . . 1 .

. . 1 . .

. . . . .

I I I I I

.

.

1

.

.

.

.

.

.

.

.

.

I

3 . 1

4 1 2

4 2 2

4 1 1

4 1 1

. 3 1

5 1 .

1 2 1

5 1 1

3 2 .

1 2 1

3 . 1

V V V

1 2

+ 3

+ 2

. 3

2 2

2 .

1 4

2 .

+ 4

2 2

. .

+ 4

V IV

.

.

1

1

.

1

2

2

2

.

1

1

IV

1

.

1

.

1

1

1

1

.

1

1

.

IV

. + . 1 1 . 1

2 . + 1 . . .

1 1 + . . 1 1

. . . . 1 . .

2 1 + . 1 . .

. 1 . . 1 . .

2 . . 1 . 1 1

2 1 . . 1 1 1

1 . + 1 . . 1

1 1 + . . 1 .

. 1 + 2 1 1 1

. . + 1 . 1 .

III III III III III III III

.

.

.

.

+

.

+

1

1

.

1

1

III

(continued)

4.3 Vegetation

77

Table 4.12 (continued) Altitude (1 = 10 m asl) Area (1 = 10 m2) N. species Ref. N. Relevé N. Clematis haenkeana Dendrophorbium bomanii Solanum palinacanthuma Senna pendula var. eriocarpaa Baccharis capitalensis Carica glandulosa Budleja diffusa Baccharis microdonta Croton saltensisa Malvastrum coromandelianum Heteropterys sylvatica Budleja iresinoides Lantana canescensa Iresine diffusa Solanum lorentzii Abutilon grandifolium Boehmeria caudata Rubus imperialis Ophryosporus piquerioides Baccharis trimera Achyrocline flaccida Agalinis genistifolia Aldama mollis Clinopodium bolivianum Chromolaena laevigata Mimosa polycarpa Pavonia sepium

112 100 47 106 1 1 1

121 100 59 1 2 + 1

122 100 58 2 3 + .

122 100 52 3 4 . .

122 100 57 4 5 + 2

123 100 39 22 6 . .

123 100 44 5 7 . .

124 100 54 21 8 . .

125 100 62 6 9 . .

125 100 53 7 10 + 2

126 100 55 20 11 + 1

126 100 46 8 12 . .

13 III III

.

.

+

1

+

.

.

+

+

.

.

.

III

1

1

.

1

.

.

1

.

.

.

1

.

III

.

.

.

.

.

1

1

.

1

.

1

1

III

1 1 .

1 . .

. . 1

1 . .

. 1 .

. 1 .

. 1 1

. . 1

1 . .

. . 1

. . .

. . .

II II II

1 .

1 1

. .

. .

1 1

. .

. .

. 1

. .

. .

1 1

. .

II II

1

.

+

+

.

.

.

.

1

.

.

.

II

. . . 1 .

. . 1 . .

1 . . . .

. . . . .

1 . 1 . .

. . . . .

. . 1 . .

1 1 . . 1

. . . . .

. 1 . . .

. 1 . 1 .

. . . . 1

II II II I I

. 1 .

. . .

. . .

. . +

. . .

. . .

. + .

1 . .

. . .

. . .

1 . .

. . +

I I I

. . . 1 .

. . . . .

. . . . .

. . . . .

. . . . .

1 . 1 . 1

. . . . .

. . . . .

. . . . .

. 1 . . .

. . . . .

. . . . .

I I I I I

.

.

.

.

.

.

.

.

.

.

1

.

I

1 1

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

I I

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

78 Table 4.12 (continued) Altitude (1 = 10 m asl) Area (1 = 10 m2) N. species Ref. N. Relevé N. Tournefortia paniculata Verbesina macrophylla var. nelidae Verbesina suncho Herb layer Elephantopus mollis Cantinoa mutabilis Modiolastrum malvifolium Bidens pilosa Salvia personata Justicia goudotii Jungia pauciflora Tagetes terniflora Tagetes filifolia Bidens subalternans Bromus catharticus Hypochaeris microcephala Leonurus japonicus Cuphea racemosa Viguiera tucumanensis var. oligodonta Praxelis clematidea Anredera cordifolia Desmodium subsericeum Galinsoga caracasana Samolus valerandi Zinnia peruviana Stevia yaconensis var. subeglandulosaa Petiveria alliacea Mimosa xanthocentra

112 100 47 106 1 .

121 100 59 1 2 .

122 100 58 2 3 .

122 100 52 3 4 .

122 100 57 4 5 .

123 100 39 22 6 .

123 100 44 5 7 .

124 100 54 21 8 .

125 100 62 6 9 .

125 100 53 7 10 .

126 100 55 20 11 1

126 100 46 8 12 .

13 I

1

.

.

.

.

.

.

.

.

.

.

.

I

1

.

.

.

.

.

.

.

.

.

.

.

I

. . .

2 1 1

2 1 1

1 2 +

2 . 1

. + +

2 1 .

1 + 1

2 1 .

2 . 1

. + 1

1 1 +

IV IV IV

. . + 1 + + . + +

+ 1 . . 1 1 + 1 1

. 2 1 + 1 1 + . .

+ 1 . . 1 . + + 1

. 1 1 + . 1 . + .

+ . . 3 . . + . .

+ . 1 . . + . . .

+ . . 2 + + + + .

+ 2 1 . 1 . . + 1

+ 2 . 1 . . . . 1

1 . 1 1 1 1 1 + 1

. 1 1 . . . + . .

IV III III III III III III III III

. + .

. + 1

1 . .

. + .

+ . .

. . 1

. . .

+ . 1

+ + 1

1 + 1

+ + .

. . .

III III III

. 1 .

. . +

+ . +

+ + .

1 . .

. + +

. . .

. + .

1 . +

1 . .

. . +

. + .

III III III

+

.

.

+

.

+

.

.

+

.

+

.

III

+ + .

. + .

+ . .

+ + 1

. . .

. . .

. . 1

. . .

+ . 1

+ + .

. . .

. + 1

III III II

2 1

. .

+ .

. .

. .

. .

. .

. +

. +

+ .

. .

+ +

II II

(continued)

4.3 Vegetation

79

Table 4.12 (continued) Altitude (1 = 10 m asl) Area (1 = 10 m2) N. species Ref. N. Relevé N. Anagallis arvensis Parthenium hysteriophorus Nicandra physalodes Conyza tunariensis Glandularia tweedieana Solanum sisymbriifolium Turnera sidoides Urtica chamaedryoides Plantago australis Verbascum virgatum Salpichroa origanifolia Rivinia humilis Fleischmannia schickendantzii Gorgonidium vermicidum Eleusine indica Galium richardianum Polygonum punctatum Tradescantia boliviana Veronica arvensis Gamochaeta pensylvanica

112 100 47 106 1 . .

121 100 59 1 2 . +

122 100 58 2 3 + +

122 100 52 3 4 . .

122 100 57 4 5 + .

123 100 39 22 6 . +

123 100 44 5 7 . .

124 100 54 21 8 . +

125 100 62 6 9 + .

125 100 53 7 10 . .

126 100 55 20 11 + .

126 100 46 8 12 . .

13 II II

. . .

1 . .

. . .

. + .

. . +

. . .

1 . .

. + .

. + +

. . .

. . +

+ . .

II II II

.

.

+

.

+

.

.

.

.

.

+

.

II

. .

. +

+ .

. .

. +

. .

+ .

. .

. .

. +

+ .

. .

II II

. + +

+ . .

. . .

+ . +

. + .

. . .

. . +

. + .

+ . .

. . .

. . .

. . .

II II II

3 .

. .

. .

. .

. .

. 1

. 1

1 .

. .

. .

. .

. .

II I

.

.

1

.

.

+

.

.

.

.

.

.

I

. .

. .

+ .

. .

+ .

. +

. .

. .

. .

. +

. .

. .

I I

.

.

.

.

+

.

.

.

.

.

.

+

I

.

+

.

.

.

.

.

.

+

.

.

.

I

. .

. .

. +

+ +

. .

. .

+ .

. .

. .

. .

. .

. .

I I

Other species: All species I in 13. Herbs: Ruellia erythropus + in 1; Bromelia serra 2, Cortaderia hieronymi 1, Festuca hieronymi + in 6; Oenothera roseaa 1, Conyza sumatrensis and Begonia boliviensis var. boliviensis + in 8; Cortaderia selloana 1 in 10; Begonia micranthera var. micranthera and Cajophora hibiscifolia + in 11; Veronica persica + in 12. Holotypus ass.: Relevé 5 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec.; relevé 13 is synthetic

80

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.13 Schinetum myrtifolio-gracilipedis ass. nova (Juglandi australis-Phoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper canopy layer Schinus gracilipesa Allophylus edulis Sebastiania brasiliensis Schinus myrtifolius Sebastiania commersoniana Vassobia breviflora Condalia buxifoliab Chloroleucon tenuiflorum Scutia buxifolia Xylosma pubescens Acacia aroma Emerging Anadenanthera colubrina var. cebil Scrub layer Jungia polita Rubus imperialis Vernonanthura squamulosa Malvastrum coromandelianum Baccharis microdonta Buddleja stachyoides Clematis haenkeana Croton saltensisa Dendrophorbium bomanii Senna pendula var. eriocarpaa Urera baccifera Baccharis coridifolia Barnadesia odorata Chamissoa altissima Senecio rudbeckiifolius Smilax campestris Solanum palinacanthuma Herb layer Jungia pauciflora Zinnia peruviana Eleusine indica Mirabilis jalapa Praxelis clematidea Fleischmannia schickendantzii

124 50 16 42 1

124 100 21 39 2

124 50 19 40 3

124 100 28 36 4

124 100 28 37 5

6

4 4 3 2 3 3 2 . . . .

5 4 4 4 3 3 3 . . 1 .

5 4 3 3 3 2 2 . 1 . .

4 4 4 3 3 2 2 1 1 1 1

5 3 3 4 2 3 2 1 . . .

5 5 5 5 5 5 5 2 2 2 1

.

1

.

1

.

2

1 1 1 1 1 . . . . . . . . . . . .

1 1 1 . . 1 1 . . 1 + . . . . . .

. 1 1 1 . . . 1 1 . . . . . . . .

1 . 1 1 . . . . . 1 . 1 . . 1 . .

1 1 . . 1 1 1 1 1 . + . 1 1 . + +

4 4 4 3 2 2 2 2 2 2 2 1 1 1 1 1 1

1 . + . + .

. 1 + + . 1

1 . . + + .

. 1 + + . 1

1 1 . . + .

3 3 3 3 3 2 (continued)

4.3 Vegetation

81

Table 4.13 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Hypochaeris microcephala Bidens subalternans Justicia goudotii Parthenium hysteriophorus Tagetes terniflora

124 50 16 42 1 1 . . . .

124 100 21 39 2 . . . + .

124 50 19 40 3 . + + . +

124 100 28 36 4 . + . + +

124 100 28 37 5 1 . + . .

6 2 2 2 2 2

Other species: All species I in 6. Herb layer: Digitaria insularis, Mimosa xanthocentra and Tagetes filifolia + in 4; Bidens pilosa, Modiolastrum malvifolium and Nicandra physalodes + in 5. Holotypus ass. : Relevé 2 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 6 is synthetic

4.3.7

List of Associations Proposed for the Study Area

• Enterolobio contortisilici-Anadenantheretum cebilis ass. nova (Association 1; Cluster 1A; Table 4.8). • Schino bumeloidis-Allophyletum edulis ass. nova (Association 2; Cluster 2B; Table 4.9). • Xylosmo pubescentis-Blepharocalycetum salicifolii ass. nova (Association 3; Cluster 2C; Table 4.10). • Jacarando mimosifoliae-Vassobietum breviflorae ass. nova (Association 4; Cluster 2A; Table 4.11). • Erythrino falcatae-Tipuanetum tipi ass. nova (Association 5; Cluster 1C; Table 4.12). • Schinetum myrtifolio-gracilipedis ass. nova (Association 6; Cluster 3A; Table 4.13). • Juglandi australis-Blepharocalycetum salicifolii ass. nova (Association 7; Cluster 3B; Table 4.14). • Zanthoxylo cocoi-Blepharocalycetum salicifolii ass. nova (Association 8; Cluster 1B; Table 4.15). • Tecomo stantis-Anadenantheretum cebilis ass. nova (Association 9; Cluster 5C; Table 4.16). • Myrciantho pseudomatoi-Blepharocalycetum salicifolii ass. nova (Association 10; Cluster 5A; Table 4.17). • Cinnamomo porphyrium-Blepharocalycetum salicifolii ass. nova (Association 11; Cluster 5B-5D; Table 4.18). • Pruno tucumanensis-Podocarpetum parlatorei Navarro and Maldonado (2002) cedreletosum angustifoliae subass. nova (Association 12; Cluster 4; Table 4.19). • Salici humboldtianae-Acacietum aromae ass. nova (Association 13; Cluster 6B; Table 4.20).

82

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.14 Juglandi australis-Blepharocalycetum salicifolii ass. nov (Juglandi australisPhoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper canopy layer Juglans australis Blepharocalyx salicifolius Anadenanthera colubrina var. cebil Erythrina falcata Parapiptadenia excelsa Tipuana tipu Lower canopy layer Sebastiania brasiliensis Allophylus edulis Vassobia breviflora Scutia buxifolia Condalia buxifoliab Schinus gracilipesa Schinus myrtifolius Celtis iguanaea Sebastiania commersoniana Xylosma pubescens Celtis ehrenbergiana var. discolora Senna spectabilis Acacia aroma Morus alba Chloroleucon tenuiflorum Sapium haematospermum Scrub layer Jungia polita Senna pendula var. eriocarpaa Vernonanthura squamulosa Chamissoa altissima Dendrophorbium bomanii Heteropterys sylvatica

124 100 24 41 1

124 100 30 32 2

124 100 23 33 3

124 100 24 34 4

124 100 22 38 5

124 100 27 31 6

124 100 25 35 7

125 100 26 30 8

125 100 40 28 9

125 100 35 29 10

125 100 39 27 11

12

. . .

3 2 1

3 2 .

3 1 .

. . .

3 1 .

4 2 .

4 3 1

3 3 .

4 1 .

4 2 .

V V I

1 . .

. 1 .

. . .

. . .

. . .

. . .

. . .

. . .

1 . .

. . 1

. . .

I I I

3 4 2 1 2 3 3 . 3 . .

3 2 2 2 1 2 2 2 . 2 .

3 2 2 2 1 1 2 2 2 2 .

3 2 1 2 1 1 2 2 . . .

3 4 3 1 2 5 4 . 2 1 1

3 2 1 2 1 2 2 1 2 . 1

3 2 2 2 1 1 2 2 2 2 1

2 2 2 1 1 2 . 1 2 2 .

3 3 2 2 1 2 1 2 . . 1

3 2 2 1 1 . 2 2 3 2 .

2 3 2 2 1 1 . 2 3 2 .

V V V V V V V V IV IV II

. . . . .

1 1 . . .

. . . . .

. . . . .

. . . 1 .

. . . . .

. . . . .

. . . . .

. . . . 1

1 . . . .

1 1 1 . .

II I I I I

1 1

2 2

3 2

4 .

1 1

2 2

2 2

. 1

3 1

3 .

4 2

V V

1 1 . .

. 1 . .

1 . 1 .

1 1 . 1

1 . . 1

1 2 1 1

. . 1 .

. . . .

1 . 1 1

1 1 1 .

. 1 . 1

IV III III III

(continued)

4.3 Vegetation

83

Table 4.14 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Malvastrum coromandelianum Rubus imperialis Acalypha amblyodonta Barnadesia odorata Mimosa debilis Smilax campestris Urera baccifera Baccharis microdonta Clematis haenkeana Baccharis coridifolia Buddleja stachyoides Croton saltensisa Senecio rudbeckiifolius Sida rhombifolia Dolichandra ungis-cati Solanum palinacanthuma Baccharis dracunculifolia Senecio hieronymi Herb layer Jungia pauciflora Zinnia peruviana Cantinoa mutabilis Justicia goudotii Mikania micrantha Praxelis clematidea Fleischmannia schickendantzii Gorgonidium vermicidum Hypochaeris microcephala Galinsoga caracasana Mirabilis jalapa Parthenium hysteriophorus Tagetes terniflora Bidens pilosa Digitaria insularis Eleusine indica

124 100 24 41 1 .

124 100 30 32 2 .

124 100 23 33 3 .

124 100 24 34 4 1

124 100 22 38 5 .

124 100 27 31 6 .

124 100 25 35 7 .

125 100 26 30 8 1

125 100 40 28 9 1

125 100 35 29 10 1

125 100 39 27 11 1

12 III

. 1 1 . + + . . . . . . + . . . .

1 . . + + + 1 1 . . . . . . . . .

. . 1 . . . . . . . . . . . . . .

1 . . . . . . . . . 1 . . . . . 1

. 1 . + . . 1 . . . . . . . . . .

. 1 . . . . . . . 1 . . . . . . .

1 . 1 . . + . . . . . . . . + . .

. . . . . . . . 1 . . 1 . + . . .

. 1 1 + + + 1 1 1 . . 1 1 . . 1 .

1 . . . + . . 1 . . 1 . . + . . .

1 . . + . . . . . 1 . . . . + . .

III II II II II II II II I I I I I I I I I

1 1 + + . + .

2 . . + + + .

2 1 . + . . 1

2 1 + . + . .

1 1 + . . + .

1 1 + . + . 1

2 1 . . . + .

2 1 . + . . 1

2 1 . + + . .

2 . + . + + .

1 1 + + + + 1

V V III III III III II

. . . . . . . . .

1 . . . . . . . +

. . . + . . . . .

. 1 . . . . . . .

. . . . . . . . .

. . + . + . . . .

. . . . . + . . .

1 1 . + . . + . .

1 . . + . + . + .

. 1 + . + + . . +

. . + . + . + + .

II II II II II II I I I

(continued)

84

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.14 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Nicandra physalodes Oenothera roseaa Panicum trichanthum

124 100 24 41 1 . . .

124 100 30 32 2 . . .

124 100 23 33 3 + . .

124 100 24 34 4 . . +

124 100 22 38 5 . . .

124 100 27 31 6 . . .

124 100 25 35 7 . . +

125 100 26 30 8 . + .

125 100 40 28 9 + . .

125 100 35 29 10 . + .

125 100 39 27 11 . . .

12 I I I

Other species: All species I in 12. Mimosa xanthocentra + in 1; Conyza tunariensis + in 9; Leonurus japonicus and Solanum sisymbriifolium + in 10; Axonopus compressus, Bromus catharticus, Cajophora hibiscifolia, Modiolastrum malvifolium and Tagetes filifolia + in 11. Holotypus ass.: relevé 9 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec.; relevé 12 is synthetic

The names of each of the associations established in the present study were assigned based on the species that were considered typical of the plant community. Thus the first name is that of a species in the representative cluster of the community (Fig. 4.6) with higher or significant phytosociological values and a high presence; that is, with high fidelity values or a marked preference for this community (Appendix C). Most of these species showed a clear association with the environmental variables, and only two showed a less significant relation, probably due to the fact that their abundance and distribution was favoured by other environmental factors; this is the case of Tecoma stans, a species which in the study area is characteristic of woodlands on high gorges, gently to moderately sloping hillsides, and river terraces (Associations 9 and 13); and Salix humboldtiana, an edaphohygrophilous species characteristic of the riparian community (Association 13). Finally, the second name corresponds to that of a dominant species that confers the physiognomy on the community. The relation between the composition and distribution of the plant associations and environmental variables is finally represented in the ordination diagram in Fig. 4.6: five groups are mainly distinguished along the first two axes of the triplot: the first is formed by associations representing the woodlands in the lower Mesotropical-lower and upper Subhumid belt located in the lower, warmer and less humid stratum of the mountain forest (“basal forest”) (Associations 1, 2 and 3); the second cluster is formed by associations representing the woodlands in the lower Mesotropical-upper Subhumid belt, also located in the “basal forest” but at a higher altitude. These are less warm and more humid than the previous ones (Associations 4, 5, 6, 7 and 8). The third cluster is formed by the associations growing in the upper Mesotropical-lower Humid belt, and comprise the forests in the high, temperate and humid areas of the mountain forest (“high forest”) (Associations 9, 10 and 11). The fourth cluster is formed by the association that represents

Altitude (1=10 m asl) Area (1=10 m2) N. Species Ref. N. Relevé N. Upper canopy layer Blepharocalyx salicifolius Anadenanthera colubrina var. cebil Tipuana tipu Juglans australis Cinnamomum porphyrium Parapiptadenia excelsa Erythrina falcata Cedrela angustifolia Enterolobium contortisiliquum Lower canopy layer Celtis iguanaea Sebastiania brasiliensis Vassobia breviflora Allophylus edulis Acacia aroma Zanthoxylum coco Scutia buxifolia Sebastiania commersoniana Schinus myrtifolius Celtis ehrenbergiana var. discolora

124 100 41 24 2 4 3 1 . 2 . . . . 2 3 1 2 1 . . . . 2

123 100 36 23 1

2 3 1 2 . . . . .

. 3 . 1 2 . . . . .

2 . 2 1 2 1 . . . .

4 3 2 . 2 1 1 1 .

125 100 62 25 3

2 3 2 2 2 1 . 2 . 1

3 4 2 2 2 . . 1 .

125 100 77 26 4

1 3 2 1 1 1 1 2 1 2

5 2 2 3 . 1 1 . .

126 100 65 19 5

2 3 2 1 . . 1 2 . 1

4 3 1 3 1 . 1 . .

126 100 59 18 6

3 5 1 3 2 1 . 1 2 1

1 1 1 . . 1 1 . 1

126 100 53 9 7

2 1 1 1 1 3 2 1 1 .

3 3 3 2 2 1 . 1 .

126 100 73 16 8

3 1 2 2 1 3 1 1 1 2

3 3 1 3 3 2 1 . .

126 100 73 17 9

2 1 1 2 2 3 2 1 1 2

2 3 3 3 3 1 1 . .

126 100 73 15 10

3 1 2 1 1 3 2 2 1 .

3 1 1 3 2 4 1 1 .

127 100 92 10 11

2 1 2 1 2 3 2 1 1 2

3 2 2 3 3 3 1 1 .

127 100 78 14 12

2 1 1 . 1 2 1 1 1 .

3 1 2 3 3 4 . . .

127 100 73 13 13

2 1 2 1 . 2 1 1 2 2

3 1 2 2 3 4 1 . 1

127 100 84 12 14

V V V V V IV IV IV IV IV

V V V IV IV IV IV II I

16

(continued)

3 1 2 1 2 3 2 2 2 1

3 2 2 3 3 4 1 . .

128 100 99 11 15

Table 4.15 Zanthoxylo cocoi-Blepharocalycetum salicifolii ass. nova (Juglandi australis-Phoebion porphyrae, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi)

4.3 Vegetation 85

2 1 2 1

1 1 1 .

1 1 2 2

1 1 1 2

125 100 77 26 4 1 1 . . 1 . 1 . 1 . 1 . 1 . . . . 2 2 2 2

126 100 65 19 5 1 1 1 1 . 2 1 1 . . . . . . . 1 . 3 1 2 2

126 100 59 18 6 1 2 1 1 . 1 1 1 . . . . . 1 . . . 2 1 . 4

126 100 53 9 7 1 . 1 2 . 1 . 1 . 1 . . . 1 2 . . 2 1 2 1

126 100 73 16 8 1 1 . . . . 1 . . . 1 . . . . 1 2 1 2 3 1

126 100 73 17 9 1 1 . . 1 2 1 1 1 . . . 2 . . . . 2 1 2 2

126 100 73 15 10 . . 1 1 1 1 . 1 1 1 1 1 . . 1 . . 3 2 1 2

127 100 92 10 11 1 2 1 1 1 2 . 1 . 1 . 1 1 1 . . . 4 1 2 2

127 100 78 14 12 1 1 . . 1 1 1 . . 1 . 1 . . . . . 2 2 2 2

127 100 73 13 13 . 1 1 1 2 . . . 1 . . . . . + . . 3 1 1 1

127 100 84 12 14 . . 1 . 1 1 . 1 . 2 1 1 . . . . . 3 2 2 2

128 100 99 11 15 1 1 . 1 1 2 1 . 1 2 . 1 2 1 + 1 .

V V V V

16 IV IV III III III III III III III II II II II II II I I

Altitude (1=10 m asl) Area (1=10 m2) N. Species Ref. N. Relevé N. Acacia caven Tecoma stans Chloroleucon tenuiflorum Condalia buxifoliab Sapium haematospermum Senna spectabilis Myrsine laetevirens Solanum riparium Cnicothamnus lorentzii Myrcianthes pseudomato Kaunia lasiophthalma Schinus gracilipesa Randia micrantha Escallonia millegrana Xylosma pubescens Duranta serratifolia Salix humboldtianac Scrub layer Barnadesia odorata Vernonanthura squamulosa Acalypha amblyodonta Urera baccifera

125 100 62 25 3 1 1 1 1 1 . . . 1 . . . . . . . 2

Table 4.15 (continued) 124 100 41 24 2 1 . . . . . . . . . . . . . . . 1

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

123 100 36 23 1 . . 1 1 . . . . 1 . 1 . . . . . .

86

Boehmeria caudata Clinopodium bolivianum Jungia polita Chamissoa altissima Heimia montanaa Abutilon grandifolium Croton salternsisa Cestrum parqui Buddleja stachyoides Carica glandulosa Dolichandra ungis-cati Lantana canescensa Senecio rudbeckiifolius Senna pendula var. eriocarpaa Smilax campestris Verbesina macrophylla var. nelidae Chromolaena laevigata Hebanthe occidentalis Manetia jorgenseniia Heteropterys sylvatica Lantana trifoliaa Mimosa polycarpa Rubus imperialis Vernonanthura pinguis Achyrocline flaccida Weddelia saltensis Acalypha plicata Senecio hieronymi

1 1 1 . . 1 . + 1 . 1 1 . . . . . . . . . . . . . . . .

. . . . . 1 . + . 1 . . . . . . . . 1 . . . . . . . . .

1 1 2 1 1 1 1 . . . . 1 1 1 1 . . . . . 1 1 . . . . . 1

2 1 2 2 1 . . 1 . . 1 1 1 . 1 . 1 1 . 1 . . . 1 . . . 1

. . 2 1 . 1 1 . . . . 1 1 . 1 1 . . . 1 . . . . . . . .

2 1 . 1 . 1 . 1 1 . 1 . . 1 2 1 . 1 1 . 1 1 . . . . . .

1 . . 1 1 . 1 1 1 1 1 . 1 1 . . . . . . 1 . + 3 1 . 1 .

2 2 1 1 1 1 1 1 . 1 1 . . . 1 1 . 1 1 . . . 1 . . 1 1 .

3 2 1 1 2 . 1 1 + . . 1 1 1 . 2 1 . . + . 1 1 . 1 . . .

2 2 2 . 2 . 1 . 1 1 1 . 1 1 . 1 1 . . 1 . 1 . . 1 1 . .

3 2 2 2 2 1 1 1 . 1 . 1 . 1 1 1 . 1 . . . . 1 2 . 1 . 1

. 2 2 1 2 1 + . 1 2 . 1 1 1 1 . 1 . 1 1 . . . 2 1 . 1 1

1 2 2 1 2 1 . 1 . 2 1 1 . . 1 1 . 1 1 . . 1 1 2 . . 1 .

1 3 1 2 1 1 1 . . 1 . 1 1 1 . 2 1 . . . 1 . 1 . 1 1 . .

IV IV IV IV IV IV IV IV III III III III III III III III II II II II II II II II II II II II (continued)

2 2 2 2 2 . 1 1 1 2 1 . 1 . . . 1 1 1 1 1 1 . 2 . 1 1 1

4.3 Vegetation 87

125 100 62 25 3 . . . . . 1 . . . . . . . . 1 . . . . . . .

125 100 77 26 4 1 1 . . . 1 1 . . 1 1 . . . 1 . . . . . . .

126 100 65 19 5 . . 1 1 . . . 1 . . . 1 . . . . . . . . . .

126 100 59 18 6 1 . . . . . . . 1 . . . . . . . . + . . . .

126 100 53 9 7 . . . 1 . . . . 1 . . . . . . . . . . . . .

126 100 73 16 8 . . . . 2 1 . . 1 1 . 1 1 1 . 1 . . 1 . . .

126 100 73 17 9 . . 1 . . . . . . 1 1 . . . . . . . . 1 . .

126 100 73 15 10 1 1 1 1 . . 1 . . . . . . . . . 1 . . . . .

127 100 92 10 11 1 . . . . . 1 1 . . 1 . 1 1 . . 1 . . . . .

127 100 78 14 12 . 1 . . 2 . . . . . . . . . . . . . . . . 1

127 100 73 13 13 . . . . . . . 1 . . . 1 1 1 . . . + . 1 . 1

127 100 84 12 14 1 1 1 1 1 . . . 1 1 1 1 . . . . 1 . . . . .

128 100 99 11 15 . . 1 . 2 . 1 . . . . . 1 . . . . + 1 . . .

16 II II II II II II II II II II II II II I I I I I I I I I

Altitude (1=10 m asl) Area (1=10 m2) N. Species Ref. N. Relevé N. Solanum lorentzii Baccharis capitalensis Baccharis dracunculifolia Baccharis microdonta Dendrophorbium bomanii Baccharis salicifoliusc Bidens squarrosa Budleja diffusa Budleja iresinoides Collaea argentina Koanophyllon solidaginoides Malvastrum coromandelianum Thalictrum venturii Baccharis coridifolia Baccharis trimera Tessaria dodoneifoliac Trixis grisebachii Clematis haenkeana Acalypha lycioides Iresine diffusa Lippia suffruticosa Sida rhombifolia

124 100 41 24 2 . 1 . 1 . 1 . 1 . . . . . . . 1 . . . . 1 .

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.15 (continued)

123 100 36 23 1 . . . . . . . . . . . . . . 1 1 . . . . 1 .

88

Solanum betaceum Muehlenbeckia sagittifolia Mimosa debilis Herbs Jungia pauciflora Tagetes terniflora Elephantopus mollis Cortaderia selloana Justicia goudotii Salvia personata Urtica chamaedryoides Petiveria alliacea Tibouchina paratropica Fleischmannia schickendantzii Adenostemma brasilianum Rivinia humilis Mikania micrantha Anredera cordifolia Cortaderia hieronymi Axonopus compressus Phytolacca bogotensis Dicliptera squarrosa Bidens pilosa Begonia boliviensis var. boliviensis Cajophora hibiscifolia Conyza sumatrensis Duchesnea indica Digitaria insularis

. . . 3 + . 1 . . . . . 1 + . . + 1 . . . . + . + . .

. . +

. + . . 1 . . . . . + . . . 1 + . . . . . . . .

3 + 1 1 1 . . . . 1 . . + + 1 . . 1 1 . + . + +

. . . 3 + 1 1 1 1 + . 1 . 1 1 . + 1 . . 1 . + . + + .

. . . 3 1 1 . . 1 + 1 . . 1 1 + . . + . . 1 . . . . .

. + + 3 + . 1 1 . + 1 1 1 . 1 . . . + . . 1 . . . . +

. . . 1 . 2 1 . 2 + . . 1 . . . + . . . 1 . . . . . .

. . . 2 . 1 2 1 2 + 1 1 1 . . . . 1 + . . + . + . . .

. . . + 1 1 2 1 2 + 1 1 . 1 + . + . . 1 . . + . . . +

. . . 2 1 + . 1 1 . . . . . . + . . + . 1 1 + . . . .

1 . . 2 1 1 1 1 2 . 2 . 1 + 1 + + 1 . 1 1 . + + + + .

. + . 1 + 1 2 1 2 + 1 1 . 1 . + . . . 1 . + . . + + +

. . . . + + . . . + . 1 1 + . . + 1 . 1 . . . + . + .

. . . 1 1 1 2 1 . . 1 1 . . 1 + . . + 1 1 . . + + . +

. . . V V IV IV IV III III III III III III III III III III III II II II II II II II II

I I I

(continued)

1 1 2 . 1 2 + 2 1 1 . 1 + + . + 1 . . + . . . .

1 . .

4.3 Vegetation 89

Altitude (1=10 m asl) Area (1=10 m2) N. Species Ref. N. Relevé N. Leonurus japonicus Mirabilis jalapa Solanum tenuispinuma Acalypha communis Oenothera roseaa Viguiera tucumanensis var. oligodonta Cantinoa mutabilis Bromelia serra Bomarea edulis Chaptalia nutans Cuphea racemosa Gorgonidium vermicidum Panicum trichanthum Sida cabreriana Tagetes filifolia Deyeuxia polygamaa Conyza tunariensis Bidens subalternans Alternanthera philoxeroidesc Galium hypocarpium Pseudechinolaena polystachya Scoparia ericaceaa Modiolastrum malvifolium

Table 4.15 (continued)

123 100 36 23 1 + . . . . . . 1 . . . . . . . . . . + . . . .

124 100 41 24 2 . . . . . . . . . . . . . . . + . . . + . . .

125 100 62 25 3 . 1 . . . . . + + . . . . . . + . . . . . . .

125 100 77 26 4 . 1 1 1 . . . 1 . + . . . . . + . . . . . . .

126 100 65 19 5 . 2 1 . 1 . . . + . + + . . . . + + . . . . .

126 100 59 18 6 . 1 . . . . . . . . . . + + . . . . . . + . +

126 100 53 9 7 . . . . . . . . . . . . . . . . . . . . . . .

126 100 73 16 8 . . . . . . . . . + . . . . + . . . . . . . .

126 100 73 17 9 . . . . . + . . . . . . . + . . + + . . . + .

126 100 73 15 10 + . . . 1 . 1 . + . + . . . . . . + . . + . .

127 100 92 10 11 . . 1 1 . . . . . + + + + + . . . . . + . . .

127 100 78 14 12 . . . . 1 1 + . . . . + + . . . . . + . . + +

127 100 73 13 13 + . . 1 . 1 1 . . . . . . . + . + . . . + . .

127 100 84 12 14 + . . 1 . . . 1 . . . + + + . . . . . + . . +

128 100 99 11 15 + . 1 . 1 1 1 . + + + . . . 1 . . . + . . + . 16 II II II II II II II II II II II II II II I I I I I I I I I

90 4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

. . . 1 . . . . . . . . . . . . . . . .

. + . . . . . . . . . . . . . . + . . .

+ . . 1 . 1 . . . . . . . . . . + . . .

. . + . . . . . . + . + . + . . . . . .

. . . . . . . . . . + . . . . . . . . .

. . . . . . . . . . . . . . . . . . . .

. . . . . . . . + . . . . . . . . . . .

. + . . . + . + . . . + . . . . . . . .

+ . . . . . . . . . . . . . . . . + . .

. . . . . . . . . . . . + . . . . . . .

+ . + . 1 . . . . . . . . . + . . . . .

. . . . . . + . . . . . . . . . . . . .

. . . . . . . + . . . . + . . + . + + +

. . . . . . 1 . . + + . . . + . . . . .

. + + . 1 . . . + . . . . + . + . . + +

I I I I I I I I I I I I I I I I I I I I

Other species: All species I in 16. Lower canopy layer: Tessaria integrifoliac 1 in 2; Zanthoxylum petiolare 1 in 5; Geoffroea decorticansb 1 in 7; Morus alba 1 in 12. Scrub layer: Cnidoscolus tubulosus + in 3; Agalinis genistifolia 1 in 4; Ludwigia peruviana 1 in 8; Aldama mollis 1 in 9; Chiropetalum boliviense 1 in 10; Chamissoa maximiliani 1 in 14. Herbs: Eleusine indica + in 1; Asclepias curassavica + in 2; Calceolaria chelidonioides, Seemannia gymnostoma and Veronica persica + in 4; Hypochaeris microcephala 1 in 5; Borreria spinosa + in 8; Anagallis arvensis + in 10; Primula malacoides 1 and Tradescantia boliviana + in 11; Gamochaeta pensylvanica + in 12; Plantago australis and Samolus valerandi + in 14; Desmodium subsericeum and Setaria parviflora + in 15. Holotypus ass.: Relevé 11 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec. b Relevant biogeographic spec. c Stenoic species; relevé 16 is synthetic

Pilea jujuyensisa Polygonum punctatumc Galium lilloia Festuca hieronymi Solanum aloysiifolium Hydrocotyle bonariensis Praxelis clematidea Stevia jujuyensisa Bromus catharticus Begonia micranthera var. micranthera Galinsoga caracasana Festuca superbaa Mimosa xanthocentra Oplismenus hirtellus Muhlenbergia schreberi Parthenium hysteriophorus Paspalum distichum Zinnia peruviana Leptochloa virgata Calceolaria elatior

4.3 Vegetation 91

92

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.16 Tecomo stantis-Anadenantheretum cebilis ass. nova (Myrciantho callicomaePodocarpion parlatorei, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper canopy layer Anadenanthera colubrina var. cebil Tipuana tipu Erythrina falcata Parapiptadenia excelsa Cedrela angustifolia Enterolobium contortisiliquum Lower canopy layer Tecoma stans Sebastiania brasiliensis Acacia aroma Allophylus edulis Trema micrantha Stillingia tenella Aralia soratensis Schinus gracilipesa Duranta serratifolia Zanthoxylum petiolare Celtis iguanaea Scrub layer Verbesina suncho Vernonanthura pinguis Verbesina macrophylla var. nelidae Acalypha plicata Justicia kuntzei Lantana canescensa Vernonanthura squamulosa Dendrophorbium bomanii Lantana trifoliaa Solanum lorentzii Clematis haenkeana Chamissoa altissima Dolichandra ungis-cati Heteropterys sylvatica Budleja diffusa Chromolaena laevigata Hebanthe occidentalis Jungia polita

118 100 42 107 1

122 100 53 108 2

125 100 51 109 3

126 100 62 110 4

128 100 61 111 5

130 100 57 113 6

131 100 58 114 7

8

1 . . 3 . 1

. 1 . 3 . .

4 1 1 . . .

4 . . . . .

4 1 1 . . .

3 2 . . . .

3 2 1 . 1 .

V IV II II I I

4 2 2 . . . . . . . 1

3 2 2 1 1 . 1 . . . .

2 2 1 . . . . . 1 1 1

2 2 2 1 1 . . 1 . . .

1 2 2 1 . 2 1 . 1 1 .

2 2 1 1 . 2 . . . . .

1 1 1 . 1 . . 1 . . .

V V V III III II II II II II II

1 2 . 1 1 1 2 1 1 1 . . . 1 1 . . .

2 1 1 1 1 1 2 . 1 . . 1 1 1 . . . .

2 2 2 2 . 1 . 1 . 1 2 . 1 . 1 . . 1

2 1 2 1 1 . 1 1 1 1 3 1 . . 1 1 1 .

1 2 2 1 1 1 1 1 1 1 3 1 1 1 . 1 . 1

1 1 2 2 1 1 1 1 1 . . . . 1 . . 1 1

2 2 2 . 1 1 . . . 1 3 1 1 . . 1 1 .

V V V V V V IV IV IV IV III III III III III III III III

(continued)

4.3 Vegetation

93

Table 4.16 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Manetia jorgenseniia Mimosa debilis Pavonia sepium Phenax laevigatus Rubus imperialis Senecio rudbeckiifolius Weddelia saltensis Urera baccifera Abutilon grandifolium Acalypha lycioides Aldama mollis Baccharis latifolia Baccharis microdonta Boehmeria caudata Clinopodium bolivianum Cnidoscolus tubulosus Croton saltensis Heimia montanaa Justicia mandoniia Senna pendula var. eriocarpaa Solanum abutiloides Solanum confusuma Tournefortia paniculata Budleja iresinoides Herbs Rivinia humilis Petiveria alliacea Pharus lappulaceus Oplismenus hirtellus Adenostemma brasilianum Salvia personata Jungia pauciflora Acalypha communis Anredera cordifolia Solanum aloysiifolium Urtica chamaedryoides Digitaria insularis Fleischmannia schickendantzii Desmodium affine Acalypha boliviensis

118 100 42 107 1 . . . . . 1 . 4 . . . . . . . . . . . . . . . .

122 100 53 108 2 . . . . . . . 4 . . 1 . 1 . . . . . . 1 . . . .

125 100 51 109 3 . . . 1 . . 1 . . . . 1 . . . . 1 . . . 1 . 1 +

126 100 62 110 4 1 1 1 1 1 1 . . . 1 . . . 1 . 1 . 1 . 1 . 1 . .

128 100 61 111 5 . . . . . . 1 . 1 1 1 . 1 . . . . . 1 . . . 1 .

130 100 57 113 6 1 1 1 . 1 1 . . 1 . . 1 . . 1 1 1 . 1 . . 1 . +

131 100 58 114 7 1 1 1 1 1 . 1 . . . . . . 1 1 . . 1 . . 1 . . .

8 III III III III III III III II II II II II II II II II II II II II II II II II

3 2 1 1 + + 1 1 . 1 . + . 1 +

3 1 1 1 + + . 1 + 1 + + 1 + +

+ 1 2 2 + . 3 2 2 . + 1 . . .

2 2 2 1 + + 2 1 2 1 1 . 1 . +

2 2 2 2 + + . . . . 1 1 1 + .

1 2 2 2 + + 2 2 1 1 . 1 . + .

2 2 2 2 + + 3 . 1 1 1 . 1 . +

V V V V V V IV IV IV IV IV IV III III III

(continued)

94

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.16 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Duchesnea indica Pseudechinolaena polystachya Salpichroa origanifolia Sida cabreriana Dicliptera squarrosa Justicia goudotii Phytolacca bogotensis Primula malacoides Chaptalia nutans Cuphea racemosa Ruellia erythropus Scoparia ericaceaa Tragia volubilis Cortaderia hieronymi Mimosa xanthocentra Solanum tenuispinuma Parthenium hysteriophorus Tibouchina paratropica Tradescantia boliviana Axonopus compressus Cantinoa mutabilis Conyza sumatrensis Desmodium subsericeum Galium hypocarpium Plantago australis Samolus valerandi Setaria parviflora Tagetes filifolia Tagetes terniflora Valeriana effusa Veronica persica

118 100 42 107 1 + + + + 1 . . . + . + . . . . . . . . + + . . . . . . . . . .

122 100 53 108 2 + + . + 1 . . 1 + + . . + . 1 1 1 . . . . . . . . . . + + . .

125 100 51 109 3 . . + . . . 1 . . + . . . 1 . . 1 . 1 . . . . . + + + + . . .

126 100 62 110 4 . . . + 1 1 1 . + + . + + . . . . . . 1 . + + . . . . . + + .

128 100 61 111 5 + + + . . 1 . 1 . . + + + . 1 . . 1 1 . + . . + + . + . . . +

130 100 57 113 6 . . . + . 1 1 . . . + . . 1 . 1 . 1 . . . + . . . . . . . + .

131 100 58 114 7 + + + . . . . 1 . . . + . . . . . . . . . . + + . + . . . . +

8 III III III III III III III III III III III III III II II II II II II II II II II II II II II II II II II

Other species: All I in 8. Lower canopy layer: Cordia saccelia 1 in 2; Coutarea hexandra and Kaunia lasiophthalma 1 in 6; Chrysophyllum marginatum and Pisonia zapallo 1 in 7. Scrub layer: Carica glandulosa 1 and Smilax campestris + in 2; Acalypha amblyodonta 1 and Sida rhombifolia + in 2; Senecio hieronymi 1 in 5; Chiropetalum boliviense and Koanophyllon solidaginoides 1 in 6; Bidens squarrosa and Buddleja stachyoides 1, Achyrocline flaccida + in 7. Herbs: Praxelis clematidea + in 1; Veronica arvensis and Zinnia peruviana + in 3; Galinsoga caracasana, Hypochaeris microcephala and Ruellia ciliatiflora + in 4; Cajophora hibiscifolia and Galium lilloia + in 7. Holotypus ass.: Relevé 7 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 8 is synthetic

4.3 Vegetation

95

Table 4.17 Myrciantho pseudomatoi-Blepharocalycetum salicifolii ass. nova (Myrciantho callicomae-Podocarpion parlatorei, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatoreiTipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Upper canopy layer Blepharocalyx salicifolius Cinnamomum porphyrium Parapiptadenia excelsa Cedrela angustifolia Anadenanthera colubrina var. cebil Erythrina falcata Juglans australis Tipuana tipu Lower canopy layer Myrcianthes pseudomato Myrsine laetevirens Allophylus edulis Sebastiania brasiliensis Prunus tucumanensis Cnicothamnus lorentzii Kaunia lasiophthalma Myrcianthes pungens Solanum riparium Xylosma pubescens Pisonia zapallo Schinus gracilipesa Scrub layer Smilax campestris Justicia mandoniia Acalypha plicata Justicia kuntzei Rubus imperialis Verbesina suncho Dolichandra ungis-cati Vernonanthura pinguis Chamissoa altissima Lantana canescensa Verbesina macrophylla var. nelidae

132 100 26 95 1

132 100 29 94 2

133 100 22 93 3

133 100 20 92 4

134 500 18 91 5

134 100 28 89 6

136 100 34 88 7

136 100 37 87 8

9

4 4 2 2 2 1 . .

4 4 2 . 2 . . .

4 4 2 . . . . .

4 4 2 . . . . 1

4 4 2 . . . . .

5 3 2 1 . . . .

5 4 3 2 . . 1 .

5 3 2 1 . . . .

V V V III II I I I

3 1 . 2 . . . . . . . .

4 1 1 2 . 1 . 1 . 1 . .

4 1 . 2 1 . . . 1 . . .

4 . 1 3 . 1 . . . . . .

4 1 1 2 . . . . . . . .

3 . 1 . . . . . 1 1 . .

3 2 1 . 2 . 1 . . . . 1

3 2 1 . 3 . 1 1 . . 1 .

V IV IV IV II II II II II II I I

2 . 1 1 1 2 1 1 . . .

2 1 . . 2 2 1 1 1 1 .

1 2 1 1 1 2 . . 1 . 1

2 2 1 . 1 . . . . 1 1

2 2 1 1 1 . . . . . .

2 1 1 2 . . . . 1 . .

1 2 1 1 1 1 1 1 . . 1

2 2 2 1 . 1 1 1 . 1 .

V V V IV IV IV III III II II II

(continued)

96

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.17 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Acalypha amblyodonta Acalypha lycioides Campovassouria cruciata Clinopodium bolivianum Dendrophorbium bomanii Ludwigia peruviana Phenax laevigatus Senecio rudbeckiifolius Solanum lorentzii Vernonanthura squamulosa Herb layer Justicia goudotii Bromelia serra Petiveria alliacea Acalypha communis Pharus lappulaceus Rivinia humilis Cortaderia hieronymi Jungia pauciflora Mimosa xanthocentra Deyeuxia polígamaa Panicum trichanthum

132 100 26 95 1 . . 1 . . 1 . 1 . .

132 100 29 94 2 1 . 1 . 1 . . 1 . 1

133 100 22 93 3 . 1 . . . . . . . .

133 100 20 92 4 . . . . 1 . . . . .

134 500 18 91 5 . . . . . 1 . . . .

134 100 28 89 6 . . . . . . 1 . . .

136 100 34 88 7 1 . . 1 . . . . 1 .

136 100 37 87 8 . 1 . 1 . . 1 . 1 1

9 II II II II II II II II II II

1 2 . . . . 1 1 . . .

1 2 . . . . 1 . 1 . .

1 1 . . . . . 1 . . .

1 . 1 . . . . . 1 . .

2 1 . . . . 1 . . + .

1 1 2 1 2 1 . . . + .

1 2 2 1 2 1 . . . . +

2 1 1 2 1 1 . . . . +

V V III II II II II II II II II

Other species: All I in 9. Scrub layer: Muehlenbeckia sagittifolia and Senecio hieronymi 1 in 1; Baccharis microdonta 1 in 2; Mimosa polycarpa 1 in 3; Senna pendula var. eriocarpaa 1 in 4; Hebanthe occidentalis and Heteropterys sylvatica 1, Ophryosporus piquerioides + in 6; Boehmeria caudata 1 in 7; Chiropetalum boliviense and Weddelia saltensis 1 in 8. Herbs: Seemannia gymnostoma + in 1; Veronica persica 1 in 3; Salpichroa origanifolia + in 4; Plantago australis + in 5; Acalypha boliviensis and Fleischmannia schickendantzii 1, Pseudechinolaena polystachya and Ruellia ciliatiflora + in 6; Urtica chamaedryoides 1, Cenchrus latifolius and Galium hypocarpium + in 7; Adenostemma brasilianum 1, Cantinoa mutabilis, Muhlenbergia schreberi and Sida cabreriana + in 8. Holotypus ass.: Relevé 7 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 9 is synthetic

4.3 Vegetation

97

Table 4.18 Cinnamomo porphyrium-Blepharocalycetum salicifolii ass. nova (Myrciantho callicomae-Podocarpion parlatorei, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatoreiTipuanetea tipi) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Upper canopy layer Cinnamomum porphyrium Blepharocalyx salicifolius Parapiptadenia excelsa Anadenanthera colubrina var. cebil Tipuana tipu Cedrela angustifolia Juglans australis Enterolobium contortisiliquum Cedrela saltensis Erythrina falcata Lower canopy layer Allophylus edulis Myrcianthes pseudomato Kaunia lasiophthalma Sebastiania brasiliensis Solanum riparium Tecoma stans Vassobia breviflora Myrcianthes pungens Aralia soratensis Bougainvillea stipitata Stillingia tenella Sapium haematospermum Celtis iguanaea

130 100 55 43 1

134 100 75 44 2

134 100 73 45 3

131 100 52 47 4

136 100 68 48 5

130 100 62 49 6

130 100 57 50 7

137 100 56 51 8

137 100 78 115 9

133 100 66 117 10

140 100 50 118 11

143 100 81 120 12

13

4

4

4

3

3

4

4

3

4

4

3

4

V

3

3

4

3

3

4

4

4

3

3

3

3

V

2

2

2

1

1

2

2

2

2

2

.

2

V

2

2

2

2

1

2

.

2

2

1

2

2

V

2 . . 1

2 1 . .

2 . . .

1 1 . .

2 . . 1

2 1 . 1

2 1 . .

1 . . .

. 2 2 .

1 3 2 .

. 3 2 .

. 3 2 .

IV IV II II

. .

. .

. .

. 1

. .

. .

1 .

. .

1 .

. .

. .

. .

I I

2 .

2 .

2 .

1 2

. 2

1 2

1 1

2 1

2 2

2 2

2 2

2 2

V IV

1

1

.

1

1

.

1

.

1

1

1

1

IV

1

1

1

1

1

1

1

1

.

.

.

.

IV

2 1 1 . 1 .

2 2 1 . . .

1 1 1 . 1 1

. 1 1 . . .

1 1 1 . . 1

1 . . 1 . 1

. 1 1 . . .

1 1 1 1 1 1

1 . . 1 1 .

. . . 1 1 1

. . . 1 . .

. . . 1 1 .

III III III III III III

. .

1 .

1 .

. .

2 .

. .

2 .

. .

. 2

. 1

. 1

. 1

II II

1

.

1

.

1

.

1

.

.

.

.

.

II

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

98 Table 4.18 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Schinus gracilipesa Duranta serratifolia Prunus tucumanensis Alnus acuminata Pisonia zapallo Myrsine laetevirens Scrub layer Acalypha plicata Verbesina macrophylla var. nelidae Boehmeria caudata Baccharis latifolia Justicia mandoniia Phenax laevigatus Vernonanthura pinguis Vernonanthura squamulosa Cestrum parqui Justicia kuntzei Weddelia saltensis Verbesina suncho Dendrophorbium bomanii Rubus imperialis Urera baccifera Aphelandra hieronymi Solanum lorentzii Heimia montanaa Clinopodium bolivianum Dolichandra ungiscati Chromolaena laevigata

130 100 55 43 1 . . . . . .

134 100 75 44 2 . . . . . .

134 100 73 45 3 . . . . 1 .

131 100 52 47 4 . . . . . .

136 100 68 48 5 1 . . . . .

130 100 62 49 6 . . . . . .

130 100 57 50 7 . 1 . . . .

137 100 56 51 8 . . . . . .

137 100 78 115 9 . 1 . . . .

133 100 66 117 10 1 . . . . 1

140 100 50 118 11 . . 2 1 1 .

143 100 81 120 12 1 1 2 1 . 1

13 II II I I I I

2 2

1 1

2 2

2 2

2 2

2 1

1 1

1 .

2 2

2 2

2 1

2 1

V V

. 1 1 1 2

1 . 1 2 2

1 1 2 . 2

1 1 1 2 2

1 . . . 1

2 1 1 2 1

. 1 1 . 1

2 1 1 2 2

1 1 . 2 .

2 2 1 2 .

1 1 + 1 .

1 1 1 2 .

V V V IV IV

1

1

.

1

1

.

1

1

1

1

.

1

IV

2 . 1 2 .

2 2 . 2 1

2 1 1 1 .

2 2 . 2 1

2 2 1 2 .

3 . 1 2 1

1 1 . 2 .

2 . 1 . .

. 1 1 . 1

. . . . 1

. 1 1 . 1

. 1 1 . 1

IV IV IV III III

. 1 .

. 1 1

1 1 .

1 1 .

1 1 1

. . 2

. . 1

1 . .

1 . 1

. 1 1

1 . .

1 1 .

III III III

1 . .

1 . 1

. 1 .

1 . .

. . 1

1 1 .

1 . .

. . 1

1 1 1

. 1 1

. 1 .

1 1 1

III III III

1

.

1

.

1

.

.

.

1

.

1

1

III

.

.

1

1

.

.

1

1

1

.

.

.

III

(continued)

4.3 Vegetation

99

Table 4.18 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Koanophyllon solidaginoides Piper hieronymi Abutilon grandifolium Croton saltensisa Ludwigia peruviana Cnidoscolus tubulosus Manetia jorgenseniia Hebanthe occidentalis Mimosa polycarpa Senecio rudbeckiifolius Muehlenbeckia sagittifolia Solanum abutiloides Solanum betaceum Solanum confusuma Ophryosporus piquerioides Acalypha lycioides Baccharis microdonta Bidens squarrosa Chamissoa altissima Heteropterys sylvatica Pavonia sepium Senna pendula var. eriocarpaa Solanum aligeruma Austroeupatorium inulifolium Carica glandulosa

130 100 55 43 1 1

134 100 75 44 2 1

134 100 73 45 3 .

131 100 52 47 4 .

136 100 68 48 5 .

130 100 62 49 6 .

130 100 57 50 7 1

137 100 56 51 8 .

137 100 78 115 9 1

133 100 66 117 10 1

140 100 50 118 11 .

143 100 81 120 12 .

13 III

. .

1 .

. 1

. .

1 1

. .

1 .

. 1

1 .

. .

1 .

. 1

III II

1 . .

. . .

. . 1

. 1 .

. 1 .

1 . .

. 1 .

1 . 1

. 1 .

. . 1

. . .

1 . 1

II II II

. .

. .

1 1

. .

1 1

. .

. .

. 1

1 .

1 .

. 1

. .

II II

1 .

. 1

. .

. .

. 1

1 .

. .

. .

. 1

1 .

. .

1 1

II II

.

.

1

.

.

.

.

.

.

1

1

1

II

. . 1 .

1 . . +

. . . .

. . 1 +

. . . .

. 1 1 .

1 . . .

. . . +

1 . . .

. 1 . .

1 1 . .

. 1 1 +

II II II II

. 1

. .

1 1

. .

. .

1 .

. .

. .

1 1

. .

. .

. .

II II

. . 1

1 1 1

. . .

. . .

1 . .

. 1 1

1 . .

. . .

. 1 .

. . .

. . .

1 . .

II II II

. .

. .

. .

. .

. .

1 1

. .

. .

. 1

1 .

. 1

1 .

II II

. .

1 .

. .

. .

. .

. .

. .

. .

. 1

1 .

. 1

1 .

II I

.

.

.

.

.

.

.

.

1

.

1

.

I

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

100 Table 4.18 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Ophryosporus lorentziia Tournefortia paniculata Thalictrum venturii Herbs Elephantopus mollis Acalypha communis Pharus lappulaceus Petiveria alliacea Rivinia humilis Phytolacca bogotensis Justicia goudotii Tibouchina paratropica Desmodium subsericeum Adenostemma brasilianum Salvia personata Cuphea racemosa Dicliptera squarrosa Solanum aloysiifolium Duchesnea indica Onoseris alata Anredera cordifolia Urtica chamaedryoides Pseudechinolaena polystachya Panicum trichanthum Conyza sumatrensis Primula malacoides Desmodium affine

130 100 55 43 1 .

134 100 75 44 2 .

134 100 73 45 3 .

131 100 52 47 4 .

136 100 68 48 5 .

130 100 62 49 6 .

130 100 57 50 7 .

137 100 56 51 8 .

137 100 78 115 9 1

133 100 66 117 10 .

140 100 50 118 11 .

143 100 81 120 12 1

13 I

1

.

.

.

1

.

.

.

.

.

.

.

I

.

.

.

.

.

.

.

.

.

+

.

+

I

3 2 1 2 2 1

3 2 2 1 2 1

3 1 1 1 2 1

2 2 2 1 1 .

2 1 2 2 2 1

3 2 1 2 1 1

3 2 1 2 2 1

3 2 1 2 2 1

2 1 2 2 1 2

1 2 2 1 1 3

2 2 2 1 . 2

2 2 2 1 1 2

V V V V V V

2 1

1 .

1 1

1 1

1 .

. 1

1 .

1 1

1 1

1 1

. 1

1 1

V IV

.

+

1

+

.

1

.

1

1

1

1

1

IV

.

+

+

.

+

.

+

+

1

1

.

1

IV

. + 1 .

+ + 1 1

+ + . 1

. + . .

+ . 1 1

+ + 1 1

. . . .

+ + . 1

+ + 1 1

+ + . 1

. . 1 .

+ . 1 .

IV IV III III

. . 1 .

. + . 1

+ . 1 .

+ . . .

+ + + 1

. . . 1

. + . .

+ + + +

+ + 1 1

+ + . .

. . 1 .

+ + . +

III III III III

+

.

+

.

+

+

.

.

+

.

1

.

III

. + . .

+ + 1 .

+ . . +

. + . .

. . 1 1

+ . . .

+ . 1 .

. + . +

+ + + +

+ . . .

. . . .

. + 1 +

III III III III

(continued)

4.3 Vegetation

101

Table 4.18 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Chaptalia nutans Galinsoga caracasana Muhlenbergia schreberi Ruellia ciliatiflora Tragia volubilis Digitaria insularis Solanum tenuispinum Samolus valerandi Plantago australis Mikania micrantha Begonia boliviensis var. boliviensis Galium hypocarpium Galium lilloi Mirabilis jalapa Bomarea edulis Acalypha boliviensis Axonopus compressus Cajophora hibiscifolia Calceolaria elatior Petunia occidentalis Salpichroa origanifolia Sibthorpia conspicua Sida cabreriana Sinningia warmingii Veronica arvensis Seemannia gymnostoma Ruellia erythropus Valeriana effusa Scoparia ericacea

130 100 55 43 1 . .

134 100 75 44 2 + +

134 100 73 45 3 . .

131 100 52 47 4 . +

136 100 68 48 5 . .

130 100 62 49 6 . +

130 100 57 50 7 + .

137 100 56 51 8 . .

137 100 78 115 9 + +

133 100 66 117 10 + +

140 100 50 118 11 . .

143 100 81 120 12 + .

13 III III

.

.

.

+

.

.

.

+

+

.

+

+

III

. + . . . . . .

+ . 2 . . . . +

+ + . 1 . + + .

. . . 1 + . . .

+ . 1 1 . . . .

. + . . . + + .

+ . 1 . + . . +

. . . 1 . . . .

. . 1 . . + + +

+ + . . + . + +

. + . . . . . .

. . . . + + . .

III III II II II II II II

. . . 1 . .

. + . . + +

+ . . . . .

. . . . + +

+ + . . . .

. . . . . .

+ . . 1 + +

+ . . . . .

. . 2 . . .

. . 1 . . .

. + . . . .

. + 2 1 . .

II II II II II II

.

.

+

.

.

.

+

.

.

+

.

.

II

+ . +

. + .

+ . .

. . .

. + .

. . +

. . .

. . .

. . .

+ . .

. . +

. + .

II II II

. . . + .

+ . . . .

. . . . .

+ . . . .

. . . . .

. + . + .

. . . . .

. . . . .

. + + . .

. . + . 1

. + . . .

+ . + + 1

II II II II I

. . .

. . .

+ . .

. . .

. . .

. + .

. . +

. . .

. . +

. + .

. . .

+ . .

I I I

(continued)

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

102 Table 4.18 (continued) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé. N. Fleischmannia schickendantzii Cenchrus latifolius Modiolastrum malvifolium Stevia jujuyensis Cortaderia hieronymi Jungia pauciflora Tagetes terniflora Leptochloa virgata

130 100 55 43 1 .

134 100 75 44 2 .

134 100 73 45 3 .

131 100 52 47 4 1

136 100 68 48 5 .

130 100 62 49 6 .

130 100 57 50 7 .

137 100 56 51 8 1

137 100 78 115 9 .

133 100 66 117 10 .

140 100 50 118 11 .

143 100 81 120 12 .

13 I

. .

+ +

. .

. .

. .

. .

. .

+ +

. .

. .

. .

. .

I I

. 1 . . +

. . . + .

+ . 1 . +

. . . . .

. 1 1 + .

+ . . . .

. . . . .

. . . . .

. . . . .

. . . . .

. . . . .

. . . . .

I I I I I

Other species: All I in 13. Canopy layer (E2): Coutarea hexandra 1 in 1; Acacia aroma and Citrus sp. 1 in 2; Trema micrantha 1 in 3; Chrysophyllum marginatum 1 in 5; Ilex argentina 1 in 11. Scrub layer: Aldama mollis, Chiropetalum boliviense and Malvastrum coromandelianum 1 in 2; Buddleja stachyoides and Mimosa debilis 1, Achyrocline flaccida + in 3; Senecio hieronymi 1 in 4; Lantana canescens 1 in 6; Cnidoscolus vitifolius 1 in 7. Herbs: Mimosa xanthocentra and Tradescantia boliviana 1 in 2; Alternanthera philoxeroides and Orthopappus angustifolius + in 3; Cantinoa mutabilis and Oenothera rosea + in 5; Borreria spinosa + in 6; Setaria parviflora + in 7; Gorgonidium vermicidum + in 11; Stevia yaconensis var. subeglandulosa 1, Parthenium hysteriophorus and Polygonum punctatum + in 12. Holotypus ass.: Relevé 6 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 13 is synthetic

the mountain woodland at the upper extreme of the altitudinal gradient, within the same bioclimatic belt but under the most temperate and humid conditions in the whole study area (Association 12). Finally, the last cluster is formed by the association that represents the subhumid and humid riparian woodlands in the mountain forest (“base forest”) within the lower Mesotropical belt (Association 13). The list of the characteristic or indicator species in the proposed associations are shown in Table 4.21.

4.3 Vegetation

103

Table 4.19 Pruno tucumanensis-Podocarpetum parlatorei Navarro and Maldonado 2002 cedreletosum angustifoliae subass. nova (Myrciantho callicomae-Podocarpion parlatorei, Tipuano-Podocarpetalia parlatorei, Podocarpo parlatorei-Tipuanetea tipi) Altitude (1=10 m asl) Area (m2) N. species Ref. N. Relevé N. Upper canopy layer Cedrela angustifolia Juglans australis Podocarpus parlatorei Blepharocalyx salicifolius Cinnamomum porphyrium Parapiptadenia excelsa Erythrina falcata Lower canopy layer Prunus tucumanensis Alnus acuminata Sambucus nigra subsp. peruviana Allophylus edulis Schinus gracilipesa Cnicothamnus lorentzii Anadenanthera colubrina var. cebil Myrcianthes pseudomato Myrsine laetevirens Kaunia lasiophthalma Stillingia tenella Duranta serratifolia Escallonia millegrana Ilex argentina Scutia buxifolia Myrcianthes pungens Bougainvillea stipitata Vassobia breviflora Zanthoxylum coco Scrub layer Rubus imperialis Austroeupatorium inulifolium Baccharis latifolia Solanum aligeruma Clinopodium bolivianum Campovassouria cruciata Boehmeria caudata Lepechinia vesiculosa

149 100 62 96 1

152 100 60 97 2

153 100 40 98 3

155 100 42 99 4

158 100 54 101 5

159 100 38 102 6

160 100 31 103 7

162 100 32 104 8

9

4 2 . 3 4 1 1

4 2 . 3 4 1 .

3 2 1 3 4 . .

3 2 1 3 3 . .

2 2 5 1 . . .

1 2 5 . . . .

1 3 5 . . . .

1 2 5 . . . .

V V IV IV III II I

1 2 1 2 . . 4 2 2 1 . 1 . . . . . 1 .

2 2 2 2 1 1 3 1 1 1 . . . . . 1 1 . .

2 2 1 1 1 . 4 2 1 1 1 1 1 . 2 . . . .

3 2 2 1 . 1 3 2 . 1 1 . 1 1 2 1 . . .

4 3 2 1 1 1 . . 1 . 1 1 1 1 . . . . .

4 4 2 2 1 . . . . . 1 . . . . . . . 1

5 4 2 1 1 1 . . . . . . . 1 . . . . .

4 4 2 1 . 1 . . . . . . . . . . . . .

V V V V IV IV III III III III III II II II II II I I I

3 1 1 1 3 . 1 .

3 . 1 1 3 . 1 1

2 1 1 . . . . .

2 2 . 1 2 1 1 1

1 2 . . . 2 1 1

. 2 1 1 1 2 1 .

2 2 1 1 . 2 . 1

2 2 1 1 1 2 . 1

V V IV IV IV IV IV IV

(continued)

104

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.19 (continued) Altitude (1=10 m asl) Area (m2) N. species Ref. N. Relevé N. Muehlenbeckia sagittifolia Ophryosporus lorentziia Phenax laevigatus Solanum confusuma Pavonia sepium Senecio hieronymi Senecio rudbeckiifolius Baccharis dracunculifolia Barnadesia odorata Dolichandra ungis-cati Bidens squarrosa Koanophyllon solidaginoides Solanum lorentzii Achyrocline flaccida Thalictrum venturii Acalypha plicata Aldama mollis Baccharis microdonta Berberis jobii Buddleja stachyoides Heimia montanaa Lantana canescensa Manetia jorgenseniia Solanum betaceum Vernonanthura squamulosa Ophryosporus piquerioides Trixis grisebachii Abutilon grandifolium Aphelandra hieronymi Cestrum parqui Cnidoscolus tubulosus Croton saltensisa Dendrophorbium bomanii Justicia kuntzei Justicia mandoniia Piper hieronymi Urera baccifera

149 100 62 96 1 1 . 1 1 . . . . 1 . 1 1 . + . . . . . . 1 . 1 . 1 + . 1 . . 1 . 1 1 . 1 .

152 100 60 97 2 . 1 1 . 1 . 1 1 . . 1 1 1 . + 1 . . . . . . 1 1 1 + . . 1 1 . 1 . . 1 . .

153 100 40 98 3 1 1 . 1 . . . 1 1 1 . . . . . 1 . . . 1 1 1 . 1 . . . . . . . . . . . . .

155 100 42 99 4 . . . . . 1 1 . . . 1 1 . . . . . 1 . . . . . . . . + . . . . . . . . . .

158 100 54 101 5 1 1 1 1 1 1 . . 1 1 . . 1 + . . 1 1 . 1 . 1 . . . . + . . . . . . . . . 1

159 100 38 102 6 . . 1 . 1 . 1 1 . 1 . . 1 + + . 1 . . . . . . . . . . . . . . . . . . . .

160 100 31 103 7 1 1 1 1 . 1 1 . . . . . . . + . . . 1 . . . . . . . . . . . . . . . . . .

162 100 32 104 8 1 1 . 1 1 1 . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . .

9 IV IV IV IV III III III II II II II II II II II II II II II II II II II II II II II I I I I I I I I I I

(continued)

4.3 Vegetation

105

Table 4.19 (continued) Altitude (1=10 m asl) Area (m2) N. species Ref. N. Relevé N. Herb layer Duchesnea indica Tibouchina paratropica Stevia yaconensis var. subeglandulosaa Sibthorpia conspicuaa Elephantopus mollis Urtica chamaedryoides Axonopus compressus Galium hypocarpium Conyza sumatrensis Pilea jujuyensisa Tradescantia boliviana Cenchrus latifolius Cajophora hibiscifolia Desmodium affine Deyeuxia poligamaa Mikania micrantha Phytolacca bogotensis Jungia pauciflora Festuca hieronymi Dicliptera squarrosa Acalypha communis Anredera cordifolia Calceolaria teucrioides Muhlenbergia schreberi Begonia micranthera var. micranthera Parthenium hysteriophorus Adenostemma brasilianum Chaptalia nutans Galinsoga caracasana

149 100 62 96 1

152 100 60 97 2

153 100 40 98 3

155 100 42 99 4

158 100 54 101 5

159 100 38 102 6

160 100 31 103 7

162 100 32 104 8

9

1 1 1

+ . 1

1 1 .

+ 1 +

1 1 +

1 1 1

1 1 1

1 1 1

V V V

+ + + + + + + + . . + . + . 1 . 1 1 1 . . .

+ + + + . . . + . + . . + + . . 1 1 . . . +

. . . + . + + . . . + . . . 1 . . . . . . .

+ . . . + + . . + + . . . . . 1 . . . . . .

. + + + + . . + + + + + + . . 1 . . . . + .

+ + + . + + . . + . . . . + . . . . 1 . + +

+ . . . . . + . . . . + . . . . . . . + . .

+ + + . . . + + . . . + . + . . . . . + . .

IV IV IV III III III III III II II II II II II II II II II II II II II

. . + +

+ + + +

. + . .

+ . . .

. . . .

. . . .

. . . .

. . . .

II II II II

Other species: All I in 8. Solanum tenuispinuma 1, Desmodium subsericeum, Digitaria insularis, Justicia goudotii and Tragia volubilis + in 1; Bomarea edulis 1, Galium lilloia + in 2; Onoseris alataa + in 4; Cortaderia hieronymi and Mimosa xanthocentra 1, Veronica arvensis + in 5. Holotypus subass.: Relevé 4 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) a Endemic spec.; relevé 9 is synthetic

106

4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Table 4.20 Salici humboldtianae-Acacietum aromae ass. nova (Prosopion albae, Salici humboldtiani-Prosopietalia albae, Salici humboldtiani-Prosopietea albae) Altitude (1=10 m asl) Area (1=10 m2) N. species Ref. N. Relevé N. Dominant canopy layer Acacia aroma Salix humboldtiana Parapiptadenia excelsa Tessaria integrifolia Sebastiania brasiliensis Celtis iguanaea Trema micrantha Vassobia breviflora Acacia caven Allophylus edulis Tecoma stans Emerging Anadenanthera colubrina var. cebil Enterolobium contortisiliquum Scrub layer Urera baccifera Baccharis salicifolius Tessaria dodoneifolia Clematis haenkeana Acalypha amblyodonta Carica glandulosa Herb layer Dicliptera squarrosa Polygonum punctatum Paspalum distichum

102 50 19 52 1

103 100 40 105 2

3

3 3 2 2 2 2 3 2 1 . .

3 2 3 3 2 1 . . . 1 4

2 2 2 2 2 2 1 1 1 1 1

. .

1 1

1 1

2 2 2 . . .

4 2 2 2 1 1

2 2 2 1 1 1

1 1 1

1 1 1

2 2 2

Other species: All 1 in 3. Alternanthera philoxeroides, Asclepias curassavica, Galinsoga caracasana and Nicandra physalodes + in 1; Sida rhombifolia, Smilax campestris, Duchesnea indica, Samolus valerandi, Bromus catharticus, Cuphea racemosa, Justicia goudotii, Praxelis clematidea, Tagetes filifolia, Tagetes terniflora, Veronica persica and Zinnia peruviana +, Anredera cordifolia, Hydrocotyle bonariensis, Jungia pauciflora, Mimosa xanthocentra, Parthenium hysteriophorus, Rivinia humilis and Tradescantia boliviana 1 in 2. Holotypus ass.: Relevé 1 Localities: All relevés from Serranías de Zapla Multiple Use Ecology Reserve (Jujuy province, Argentina) Relevé 3 is synthetics

Name Enterolobio contortotisilici-Anadenantheretum cebilis (Table 4.8)

Schino bumeloidis-Allophyletum edulis (Table 4.9)

Xylosmo pubescentis-Blepharocalycetum salicifolii (Table 4.10)

Jacarando mimosifoliae-Vassobietum breviflorae (Table 4.11)

Erythrino falcatae-Tipuanetum tipi (Table 4.12)

Schinetum myrtifolio-gracilipedis (Table 4.13)

Juglandi australis-Blepharocalycetum salicifolii (Table 4.14)

Assoc. N. 1

2

3

4

5

6

7

Characteristic or Indicator species Parapiptadenia excelsa, Anadenanthera colubrina var. cebil, Enterolobium contortisiliquum, Sebastiania brasiliensis, Acacia aroma, Sebastiania commersoniana, Vassobia breviflora, Celtis iguanaea, Zanthoxylum petiolare, Carica quercifolia, Urera baccifera, Rivinia humilis, Solanum aloysiifolium, Elephantopus mollis, Myroxylon peruiferum, Terminalia triflora; (16 species) Allophylus edulis, Acacia aroma, Vassobia breviflora, Xylosma pubescens, Schinus bumeloides, Manihot grahami, Urera baccifera, Capsicum chacoense, Elephantopus mollis, Mirabilisjalapa, Samolus valerandi; (11 species) Blepharocalyx salicifolius, Sebastiania brasiliensis, Scutia buxifolia, Xylosma pubescens, Urera baccifera, Capsicum chacoense, Senna occidentalis; (7 species) Vassobia breviflora, Celtis iguanaea, Acacia aroma, Schinus fasciculatus, Jacaranda mimosifolia, Chloroleucon tenuiflorum, Geoffroea decorticans, Prosopis alba, Barnadesia odorata, Dendrophorbium bomanii, Modiolastrum malvifolium; (11species) Tipuana tipu, Erythrina falcata, Sebastiania brasiliensis, Allophylus edulis, Urera baccifera, Acalypha amblyodonta, Baccharis coridifolia, Solanum palinacanthum, Cantinoa mutabilis, Modiolastrum malvifolium, Prosopis alba, Turnera sidoides; (12 species) Schinus gracilipes, Allophylus edulis, Sebastiania brasiliensis, Schinus myrtifolius; (4 species) Juglans australis, Blepharocalyx salicifolius, Sebastiania brasiliensis, Allophylus edulis, Schinus myrtifolius, Jungia polita, Senna pendula var. eriocarpa, Jungia pauciflora; (8 species)

Table 4.21 List of species characteristics or indicators of the phytosociological associations identified in the study area

(continued)

74

46

140

79

92

99

Total rel. 93

4.3 Vegetation 107

Tecomo stantis-Anadenantheretum cebilis (Table 4.16)

Myrciantho pseudomatoi-Blepharocalycetum salicifolii (Table 4.17) Cinnamomo porphyrium-Blepharocalycetum salicifolii (Table 4.18)

Pruno tucumanensis-Podocarpetum parlatorei (Table 4.19)

Salici humboldtianae-Acacietum aromae (Table 4.20)

9

10

12

13

11

Name Zanthoxylo cocoi-Blepharocalycetum salicifolii (Table 4.15)

Assoc. N. 8

Table 4.21 (continued) Characteristic or Indicator species Blepharocalyx salicifolius, Anadenanthera colubrina var. cebil, Zanthoxylum coco, Barnadesia odorata, Urera baccifera, Thalictrum venturii, Jungia pauciflora; (7 species) Anadenanthera colubrina var. cebil, Tecoma stans, Trema micrantha, Verbesina suncho, Vernonanthura pinguis, Verbesina macrophylla var. nelidae, Petiveria alliacea, Rivinia humilis, Pharus lappulaceus, Oplismenus hirtellus; (10 species) Blepharocalyx salicifolius, Cinnamomum porphyrium, Myrcianthes pseudomato, Justicia mandonii; (4 species) Blepharocalyx salicifolius, Cinnamomum porphyrium, Cedrela angustifolia, Cedrela saltensis, Kaunia lasiophthalma, Myrcianthes pungens, Aralia soratensis, Bougainvillea stipitata, Stillingia tenella, Duranta serratifolia, Acalypha plicata, Baccharis latifolia, Justicia mandonii, Phenax laevigatus, Aphelandra hieronymi, Piper hieronymi, Solanum betaceum, Elephantopus mollis, Acalypha communis, Phytolacca bogotensis, Onoseris alata, Petunia occidentalis; (22 species) Podocarpus parlatorei, Prunus tucumanensis, Alnus acuminata, Cedrela angustifolia, Ilex argentina, Austroeupatorium inulifolium, Solanum aligerum, Clinopodium bolivianum, Campovassouria cruciata, Lepechinia vesiculosa, Ophryosporus lorentzii, Solanum confusum, Tibouchina paratropica, Stevia yaconensis var. subeglandulosa, Sibthorpia conspicua, Sambucus nigra subsp. peruviana, Berberis jobii, Calceolaria teucrioides; (18 species) Acacia aroma, Salix humboldtiana, Tessaria integrifolia, Tecoma stans, Urera baccifera, Baccharis salicifolius, Tessaria dodoneifolia, Polygonum punctatum, Paspalum distichum, Alternanthera philoxeroides, Asclepias curassavica; (11 species) 45

111

165

77

127

Total rel. 183

108 4 Geobotany of Serranías de Zapla Multiple Use Ecology Reserve:. . .

Chapter 5

Biodiversity Analysis: A Geobotanic Interpretation

5.1

Floristic Composition in Detail

The Serranías de Zapla Multiple Use Ecology Reserve contains a representation of 257 species belonging to 194 genera and 66 botanical families. Of the total species recorded, 65 correspond to the tree layer, 85 to the shrub layer and 107 to the herbaceous layer (Appendix B). The flora of the province of Jujuy is one of the most diverse in the Republic of Argentina. It comprises a total of 170 families, 972 genera and 2831 species (Zuloaga et al. 1999) distributed in the various ecological belts and plant formations throughout the region. According to these figures, the study area contains a representation of 39% the families, 20% of the genera and 9.1% of the species in the province. Specifically, for the subtropical mountain woodlands or Yungas in northwest Argentina, and especially in the province of Jujuy, a complete floristic catalogue has yet to be compiled. Many of the studies have focused on the tree layer and very few have undertaken the combined analysis of the three main vegetation layers; added to this is the fact that practically none has employed a theoretical approach and the phytosociological methodology. In this regard, and specifically for the province of Jujuy, only two phytosociological studies have recently been done in these forests, one of which analyses the vegetation at the level of the three layers (Martín 2014), while the other focuses solely on arboreal vegetation (Haagen Entrocassi 2014). The floristic composition showed variations throughout the altitudinal gradient existing in the study area: one cluster of species is distributed along the whole of the altitudinal gradient, while another appears exclusively within a particular interval of the gradient (Appendix C). It was observed that 181 (70.4%) of the first cluster of species are distributed along the whole of the altitudinal gradient, most in a discontinuous manner, but all are present in the two bioclimatic belts in the Reserve (lower and upper Mesotropical) (Appendix C). These species generally have a wide range of ecological tolerance that allows them to occupy a range of environments (widehabitat or eurioic species). However, most have their optimum distribution and © Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_5

109

110

5 Biodiversity Analysis: A Geobotanic Interpretation

frequency within certain altitudinal intervals, It and Io and only some are more indifferent to these variables. These include some species that behave as dominant and contribute significantly to the physiognomy of the plant communities of which they form part. This can be seen from the positions they occupy in the CCA ordination diagram (Fig. 4.7), where they are located close to the central coordinates or above zero on axis 1. Examples are tree species such as Parapiptadenia excelsa (Para), Anadenanthera colubrina var. cebil (Anad), Allophylus edulis (Allo) and Schinus gracilipes (Schg). Most of these species (108) occur with frequencies
20%; 62 have a frequency of between 21–50%, and only 11 species have a high frequency (over 50%). These include Allophylus edulis (84%), Blepharocalyx salicifolius (58%), Parapiptadenia excelsa (61%), Acacia aroma (56%), Anadenanthera colubrina var. cebil (56%), Elephantopus mollis (52%), Jungia pauciflora (54%), Justicia goudotii (54%), Sebastiania brasiliensis (79%), Urera baccifera (59%), Vassobia breviflora (61%) and Vernonanthura squamulosa (66%) (Appendices B and C). The first three of these species have also been reported as the most frequent throughout the different altitudinal and latitudinal gradients analysed in the tree layer in Las Yungas in Argentina, and have their greatest ecological importance at altitudes between 1300–1500 m asl (Malizia et al. 2012). Elsewhere, 76 species (29.6%) were recognised that are not present along the whole altitudinal gradient but are distributed in one or another of the bioclimatic belts. Within the study area these species show a narrower ecological distribution range and function as ecological and bioclimatic indicators: 41 species (53.9%) are distributed exclusively in the lower Mesotropical belt, and 35 (46.1%) in the upper Mesotropical belt (Appendix C). This is consistent with the commonly observed fact that species have an interval of tolerance to particular environmental factors and are therefore distributed within a limited range that is specific to that interval, where they tend to be more abundant and more frequent, and to decrease or disappear towards both extremes of the environmental gradient (Matteucci and Colma 1982). The following are exclusive to the lower Mesotropical belt in the study area: Acacia caven, Anagallis arvensis, Baccharis capitalensis, Baccharis coridifolia, Bidens pilosa, Bidens subalternans, Bromus catharticus, Capsicum chacoense, Carica quercifolia, Celtis ehrenbergiana var. discolor, Chamissoa maximiliani, Chloroleucon tenuiflorum, Condalia buxifolia, Conyza tunariensis, Eleusine indica, Galium richardianum, Gamochaeta pensylvanica, Geoffroea decorticans, Glandularia tweedieana, Iresine diffusa, Jacaranda mimosifolia, Leonurus japonicus, Manihot grahami, Morus sp., Myroxylon peruiferum, Nicandra physalodes, Prosopis alba, Randia micrantha, Schinus bumeloides, Schinus fasciculatus, Schinus myrtifolius, Sebastiania commersoniana, Senna occidentalis, Senna spectabilis, Solanum palinacanthum, Solanum sisymbriifolium, Terminalia triflora, Turnera sidoides, Verbascum virgatum, Verbena litoralis and Viguiera tucumanensis var. oligodonta. This assessment does not include the species typical of riparian environments or Eucalyptus sp., as this is an introduced species. All these species require the warmer and less humid environments in the study area, as indicated in the ordination chart of the Canonical Correspondence Analysis

5.1 Floristic Composition in Detail

111

(Fig. 4.7: Sector I and II), where they show a positive response to high values of It (399–429) and low values of Io (4.7–5.5) and altitude (1015–1275 m asl) (Appendix A: Clusters 1A, 2B, 2C, 2A, 1C, 3A, 3B, 1B, and 6B). Some even have more “xeric” preferences as will be indicated below. Many of them (17 species; 44.7%) have a diagnostic value and behave as characteristic of the communities occupying the lower level of the mountain forest in the Reserve (“basal forest”, between 1015–1275 m asl) (Appendix C and Fig. 4.7: Sector I and II). However, it is worth noting that these species are also frequent in the lower altitudinal belt of the subtropical mountain woodlands, in what is known as the foothill woodland (Cabrera 1994). This vegetation layer grows in the foothills and low mountain ranges (between 350–550 m asl) and is therefore not represented in the Reserve; furthermore, some of these species descend to the more xeric areas of the Chacoan woodland where they have the lower limit of their distribution (Cabido et al. 1991; Giorgis et al. 2011). In particular, the species that ascend from the foothill woodland exploit the warm and subhumid conditions prevailing in the “basal forest” in the Reserve, although they are neither abundant or frequent within the vegetation; with the exception of Sebastiania commersoniana and Condalia buxifolia, which are better represented (Appendix C). These last species, along with Geoffroea decorticans and Prosopis alba, are characteristic of the Chacoan biogeographical province and therefore come from more xeric areas. The presence of these thermophilous species is fairly common in areas of the foothill woodland ecotone with Chacoan woodland, in what are known as the “transitional Yungas”, where they form structurally simpler and less diverse woodlands (Brown et al. 2002). However, under certain environmental conditions these three species ascend throughout the altitudinal gradient and become established in some subhumid areas within Las Yungas sensu stricto, such as those in the study area, where they form part of the pluviseasonal woodland. This would explain their presence in the lower Mesotropical belt and their absence from the higher areas of the Reserve, namely in the upper Mesotropical belt with its increased humidity and lower temperatures. These results indicate that these species find the upper limit of their distribution interval in the study area, and are limited to the lowest level of the altitudinal gradient in the mountain forest (“basal forest”) (Fig. 4.7: Sector I and II). These species disappear at high elevations outside the intervals of this gradient. They therefore behave as indicator species of the lower Mesotropical-lower Subhumid and upper Subhumid belt that characterises the lower, warm and subhumid areas of the subtropical mountain woodlands in the Reserve. The following species are exclusive to the upper Mesotropical belt in the study area: Acalypha boliviensis, Alnus acuminata, Aphelandra hieronymi, Aralia soratensis, Austroeupatorium inulifolium, Baccharis latifolia, Berberis jobii, Calceolaria teucrioides, Campovassouria cruciata, Cedrela saltensis, Chrysophyllum marginatum, Cnidoscolus vitifolius, Cordia saccelia, Coutarea hexandra, Ilex argentina, Justicia kuntzei, Justicia mandonii, Lepechinia vesiculosa, Myrcianthes pungens, Onoseris alata, Ophryosporus lorentzii, Petunia occidentalis, Piper hieronymi, Pisonia zapallo, Podocarpus parlatorei, Prunus tucumanensis, Sambucus nigra ssp. peruviana, Sibthorpia conspicua, Sibthorpia repens, Sinningia

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5 Biodiversity Analysis: A Geobotanic Interpretation

warmingii, Solanum abutiloides, Solanum aligerum, Solanum confusum, Stillingia tenella and Valeriana effusa. These species are less thermophilous and have greater humidity requirements. They have their optimum distribution in higher, temperate humid environments in the study area; this can be seen from the ordination diagram in the Canonical Correspondence Analysis (Fig. 4.7: Sector III and IV) where they show a positive response to high values of altitude (1260–1620 m asl) and Io (7.7–8.7) and low values of It (352–395) (Appendix A: Clusters 5C, 5A, 5B–5D and 4). Most of them (25 species, 71.4%) have a diagnostic value and behave as characteristics of the communities occupying the upper Mesotropical belt. Taking into account their distribution within this bioclimatic belt, one cluster of these species is found only in the higher stratum of the mountain forest (“high-mountain forest”) (between 1260–1433 m asl; It ¼ 374–395; Io ¼ 7.7–8.2) (Appendix A: Clusters 5C, 5A and 5B–5D); it includes Acalypha boliviensis, Aralia soratensis, Cedrela saltensis, Chrysophyllum marginatum, Cnidoscolus vitifolius, Cordia saccelia, Coutarea hexandra, Onoseris alata, Petunia occidentalis, Pisonia zapallo, Sibthorpia repens, Sinningia warmingii, Solanum abutiloides and Valeriana effusa (Appendix C: Clusters 5C, 5A and 5B– 5D). Another cluster of species is practically exclusive to the highest vegetation belts corresponding to the so-called mountain woodland (Cabrera 1994), which in the study area extends mainly along the edges and borders of mountain ranges above 1488 m asl (It ¼ 352–367; Io ¼ 8.3–8.7) (Appendix A: Cluster 4; Fig. 4.7: Sector IV). Examples include Alnus acuminata, Austroeupatorium inulifolium, Berberis jobii, Calceolaria teucrioides, Campovassouria cruciata, Ilex argentina, Lepechinia vesiculosa, Ophryosporus lorentzii, Podocarpus parlatorei, Prunus tucumanensis, Sambucus nigra ssp. peruviana, Sibthorpia conspicua and Solanum aligerum (Appendix C: Cluster 4). Finally, another cluster of species is distributed with varying degrees of presence throughout the mountain forest and mountain woodland in the whole of the upper Mesotropical belt, such as for example Aphelandra hieronymi, Baccharis latifolia, Justicia kuntzei, Justicia mandonii, Myrcianthes pungens, Piper hieronymi and Stillingia tenella (Appendix C; Fig. 4.7). These species therefore behave as indicators of the upper Mesotropical-lower Humid belt and characterise the higher, temperate and humid areas in the subtropical mountain woodlands in the Reserve. These results also show that within the upper Mesotropical belt there is also a sectorisation in the distribution of some species, where species occupying sites at higher elevations (mountain woodland) can be differentiated from those that are distributed at a lower altitude (high-mountain forest) within the same bioclimatic belt. Of the 25 endemic species recorded in the study area, six are distributed exclusively in the upper Mesotropical belt, three in the lower Mesotropical, and the rest in both belts. The occurrence of a greater number of endemic species in the upper Mesotropical could indicate greater singularity in terms of floristic composition in this belt.

5.2 Species Richness

5.2

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Species Richness

With regard to the total species richness of the subtropical mountain woodlands in Argentina, there are only specific data on the number of tree species, due to the value of many of them for biology and forestry, making them the object of greater interest both to science and in terms of their conservation. Until now, approximately 204 tree species have been recorded in this woodland formation (Brown and Malizia 2007), so the 65 tree species identified in the study area would represent 32% of the total existing in Las Yungas. These characteristics highlight the floristic diversity of the tree layer in the Serranías de Zapla Multiple Use Ecology Reserve, particularly when considering the small size of the territory covered by this Reserve (37,139 hectares) within the subtropical mountain woodland formation, which in Argentina covers an approximate area of 5.2 million hectares; it extends from the Bolivian border (22 S) to the north of Catamarca Province (29 S) and includes the provinces of Salta, Jujuy and Tucumán (Brown et al. 2002; Brown and Malizia 2007). These results differ from those reported in another study on the tree layer in one sector of Serranías de Zapla (Cuyckens 2005) which recorded 46 tree species; the differences found may be linked to the smaller size of the total area sampled by Cuyckens (2005) (30,000 m2), leading to an incomplete representation of the tree species in the location. However, 54 tree species were identified in a similar area (27,500 m2) in the El Caulario site in the lower layer of the subtropical mountain woodlands (between 935–1150 m asl), revealing the high diversity existing in this altitudinal vegetation belt (mountain forest) (Haagen Entrocassi 2014). The results obtained for the families with the greatest species richness (Asteraceae, Fabaceae, Poaceae, Solanaceae and Euphorbiaceae) (Appendix B; Fig. 4.1) coincide with the findings of Zuloaga et al. (1999), who included these families within the ten most represented in the province of Jujuy; on this point it can be confirmed that 11% of the Asteraceae and Fabaceae species, 6% of the Poaceae species, 12% of the Solanaceae species and 26% of the Euphorbiaceae species in the whole province were found in the study area. Particularly in regard to the tree layer, of the 31 families recorded, the best represented were Fabaceae (12 species), Euphorbiaceae (5 species), Anacardiaceae and Myrtaceae (4 species), and to a lesser degree Asteraceae, Celtidaceae and Rutaceae (3 species) (Fig. 4.2). Similarly, in the aforementioned study on the arboreal communities in the El Caulario site (Haagen Entrocassi 2014) the best represented family was Fabaceae (11 species) from a total of 28 families identified. These results once again differ from those found in the study by Cuyckens (2005) in a sector of the Serranías de Zapla, where four Fabaceae species, two Euphorbiaceae, one Anacardiaceae and one Rutaceae are reported, in addition to the occurrence of some families that were not relevéd in the present study (Polygonaceae and Proteaceae) or that were included here in the shrub layer (Urticaceae and Piperaceae). Missing from her study are species belonging to the Celtidaceae, Bignoniaceae, Moraceae, Rubiaceae, Salicaceae, Sapotaceae and Verbenaceae families, whereas the number of Asteraceae and Mirtaceae species was similar. It was observed that the greatest species richness is found in the shrub (25 families with 85 species) and herbaceous layers (37 families with 107 species) (Appendix B);

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in the shrub layer the most represented families were Asteraceae (28 species), Solanaceae (8), Euphorbiaceae (7), Fabaceae (5) and Malvaceae (4) (Fig. 4.3), and Asteraceae (23) and Poaceae (18) in the herbaceous layer (Fig. 4.4). All these families are also among the ten best represented in the province of Jujuy, according to the list of Zuloaga et al. (1999). These results differ in part from those reported by Martín (2014) in her study on the study of the arboreal, shrub and herbaceous vegetation of the subtropical mountain woodlands in the middle basin of the Chijra river (San Salvador de Jujuy). 172 species, 120 genera and 67 families were recorded, and the number of species belonging to the three layers was lower: 37 tree species belonging to 23 families, with Fabaceae (5), Asteraceae (4) and Euphorbiaceae (3) also coinciding as the best represented families; 47 shrub species corresponding to 16 families, and 88 herbaceous species distributed in 22 families, with Asteraceae and Solanaceae the best represented families in both layers, in agreement with the findings of this present study. Possibly the differences observed in species richness are also due to the smaller size of the total areas sampled by Martín (2014) (41,500 m2 approximately). The species richness of the relevés (Fig. 4.5) showed a correspondence with the size and location of the sampling plots (transects). So the relevés located in both the foothills and on gentle to moderate slopes on the edges of mountain ranges generally had a greater species richness, which may be mainly due to the presence of more stable environments with generally more developed soils. Most of the relevés (91) with an area of 1000 m2 contained between 20 and 60 species; another set of 24 relevés with the same area had the highest richness values with over 60 species, and there were even exceptional values in a few sites (over 90 species). This is probably due to their location in sectors that are generally difficult to access and therefore better conserved, whereas the relevés with smaller surface areas (500 m2) had a low species richness (up to 20 species), very probably owing to their location in more heterogeneous and unstable sites whose specific geomorphological characteristics restrict the establishment of many species (such as ravines and steep gorges, steeply sloping foothills and riparian environments) (Appendix B: relevés 40, 42, 52, 90 and 91). The lower species richness observed in the riparian relevés is probably determined by the typical features of these environments (immature and poorly drained soils, or soils subject to seasonal flooding) that reveal a degree of environmental instability and condition the development of a specific flora and vegetation (Sirombra and Mesa 2010). This may therefore influence the species richness, as not all species can adapt to live in these environments. Similar results were also observed in the studies of subtropical mountain woodland vegetation in the Chijra (Martín 2014) and Caulario river basins (Haagen Entrocassi 2014). Some studies conducted in subtropical mountain woodlands (Grau et al. 1995; Morales et al. 1995; Blundo et al. 2012) have reported a decrease in species richness along clearly defined altitudinal gradients. In the present study, although the species richness of the relevés varied along the altitudinal gradient, no general declining pattern was detected with an increase in altitude (Table 4.21). This could be due to the fact that the altitudinal interval studied (600 m) is not sufficiently pronounced to reflect a decrease in species richness in relation to the increase in altitude. These findings

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coincide with the results obtained in the studies on subtropical mountain woodland vegetation in the Chijra river basin for an altitudinal gradient of 500 m (Martín 2014) and in the El Caulario site, for a gradient of 250 m (Haagen Entrocassi 2014).

5.3

Species Frequency

The analysis of species frequency revealed the existence of a few species with a high frequency and many species with low frequency (Appendix B). It was observed that six species appear in over 60% of the relevés and therefore have a high presence index (IV and V): Allophylus edulis (84%), Celtis ehrenbergiana var. discolor (60%), Parapiptadenia excelsa (61%), Sebastiania brasiliensis (79%), Vassobia breviflora (61%) and Vernonanthura squamulosa (66%). With the exception of this last species, all the others are trees. Many of these species have a broad distribution in the subtropical mountain woodlands in Jujuy, and their high capacity to adapt to different environmental conditions allows them to grow in more sites, thus increasing their frequency. Most of them behave as dominant species in some of the communities in the study area and contribute to their physiognomy. In the work by Martín (2014) and Haagen Entrocassi (2014) in the subtropical mountain woodlands of Jujuy, some of these species occur with a high frequency, e.g. Allophylus edulis (70–81%), Sebastiania brasiliensis (100%) and Vassobia breviflora (62%). A large number of species (173) were observed with a low frequency and were present in fewer than 20% of the relevés and with low presence indices (I) (Appendix B); these include many species that appear along the whole of the altitudinal gradient, but in a discontinuous or scattered manner. They do not have a preference for any particular community and behave as companion species; others of these fairly infrequent species generally have a more restricted distribution and a narrower optimum tolerance interval (stenoic); they are limited to certain environmental conditions and have a value as diagnostic species in their community; finally, others are occasional or very infrequent species. A very low frequency of exotic tree species was also observed (Citrus sp. 3%, Eucalyptus sp. 4%, and Morus alba 5%), and their isolated presence in the woodland is linked to the proximity of rural settlements. Similarly, there was a low frequency of shrub and herbaceous exotics, except for Duchesnea indica (25%), Leonurus japonicus (19%) and Mirabilis jalapa (28%), adventitious species that have become naturalised in some places in Las Yungas, and whose frequency indicates a certain degree of anthropic intervention in some sectors of the study area. It was generally observed in the field that high-frequency species tended to be more abundant. However, this observation was subjective, as the abundance of each species (total number of individuals) was not determined owing to the complexity of the woodlands studied, mainly in terms of their diversity and structure, and for operational reasons given the total surface area sampled (117,500 m2). The frequency of a species provides a measure of its abundance (Raunkiaer 1918). It has been empirically observed in most communities that many species are represented by a few individuals

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(less frequent) and a few very numerous species (very frequent); according to this, it could also be assumed that the most frequent species recorded in the study area would therefore be the most abundant, and the opposite would occur with the least frequent. Specifically, this positive correlation between species abundance and frequency has been corroborated in other studies of nearby subtropical woodlands with a lesser extension (Cuyckens 2005; Martín 2014; Haagen Entrocassi 2014).

5.4

Biological Spectrum

The analysis of the biological spectrum based on the Raunkiaer classification (Raunkiaer 1934; Ellenberg and Mueller-Dombois 1967) effectively shows the type of woodland formation growing in the study area, where phanerophytes constitute the dominant biotype (57%). The lifeforms recorded reveal the stratification of these woodlands: the tree layer is distributed between microphanerophytes and mesophanerophytes; in the lower level there is a shrub layer with a variable height and cover, as it depends on the height and density of the tree canopy. This layer is mainly represented by nanophanerophytes and a very low proportion of chamaephytes, then there is a herbaceous layer whose cover is associated mainly to the degree of light in the woodland and where there is a predominance of perennial biotypes such as hemicryptophytes, followed by therophytes and geophytes. Raunkiaer’s proposal is based on the correspondence between lifeforms and climate: the predominance of certain lifeforms as an indicator of the climate conditions in a particular area. The biological spectrum obtained resembles Raunkiaer’s proposal for warm and humid tropical regions with an oceanic character, characterised by a “phanerophytic” phytoclimate where the unfavourable season corresponds to the cold dry season, and where phanerophytes represent over 50%, followed by hemicryptophytes, chamaephytes, therophytes and finally geophytes. This spectrum also shows a correspondence with Dansereau’s climate diagram (1951), which associates the dominant lifeforms with temperature and precipitation. According to this author, the dominant biotypes in woodlands and forests in warm humid climates are phanerophytes and hemicryptophytes, frequently accompanied by chamaephytes.

5.5

Analysis, Interpretation and Characterisation of the Plant Communities in the Study Area

The plant communities established in the present study belong to the subtropical mountain woodland formation or Yungas, equivalent to the Bolivian-Tucuman sub-Andean pluviseasonal mountain vegetation in the Bolivian-Tucuman biogeographical province (Navarro and Maldonado 2002). According to the classic

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phytogeographical typology of Cabrera and Willink (1980) and Cabrera (1994) for Argentina, these woodlands belong to the mountain forest and mountain woodland, which are the altitudinal vegetation layers characteristic of the phytogeographical province of Las Yungas. Two types of woodlands develop in the study area based on their environmental location: terrestrial woodlands (non-riparian) which occupy a greater area and grow on foothills, hillsides, gorges, ravines and the edges of mountain ranges; and riparian woodlands occupying low-lying floodable terraces and beaches of rivers and streams. Structurally they conform microwoodlands (between 4 and 8 metres high) and mesowoodlands (over 8 metres high) according to the classification of Cabrera and Willink (1980). According to the percentage of tree canopy, we can distinguish woodlands that are open or disperse (25–50%), semi-open or interrupted (50–75%), and closed or continuous (over 75%) (Matteucci and Colma 1982). Two types of woodland can be distinguished based on the duration of the foliage: deciduous (over 60% of individuals in the tree canopy and/or understorey are deciduous due to drought), semi-deciduous (part of the canopy and/or understorey have up to 60% of deciduous individuals due to drought), and seasonally evergreen (with green foliage all year round; however there is a reduction in the foliage in the season due to partial defoliation) (Navarro and Maldonado 2002). These are subtropical pluviseasonal woodlands as they are located to the south of the Tropic of Capricorn and are subject to the seasonality of the precipitation. They therefore depend closely on the alternation between the dry season (the end of autumn, winter and early spring) and the wet season (end of spring and summer and early autumn). From the bioclimatic point of view, the plant communities in the study area are distributed along an altitudinal gradient of 600 m (between 1015 and 1620 m asl) which comprises two bioclimatic belts: lower Mesotropical-lower Subhumid and upper Subhumid, distributed between 1015 and 1275 m asl (It ¼ 399–429; Io ¼ 4.7–5.5); and upper Mesotropical-lower Humid between 1278 and 1620 m asl (It ¼ 352–392; Io ¼ 7.8–8.7). It also includes the transitional band between both layers, located at approximately 1260 m asl (It ¼ 395; Io ¼ 7.7). The mountain forest extends throughout both bioclimatic belts, up to the level between 1433–1488 m asl approximately, whereas the mountain woodland is only distributed in the higher section of the upper Mesotropical belt (between 1488–1620 m asl). This altitudinal gradient is the main factor responsible for the variations in temperature and humidity, and thus for the changes in the floristic composition and spatial distribution of the communities. Thirteen (13) plant communities were recognised within these bioclimatic belts and based on the altitudinal gradient, eight (8) of which belong to the woodlands that extend throughout warm and subhumid areas at lower elevations in the lower Mesotropical-lower and upper Subhumid belt (Associations 1, 2, 3, 4, 5, 6, 7 and 8), although one reappears in some sectors with a lower Humid ombrotype (Association 5); four (4) communities correspond to woodlands growing at higher altitudes in more temperate and humid areas within the upper Mesotropical-lower Humid belt (Associations 9, 10, 11 and 12) and in the transitional band between both bioclimatic belts; and finally one (1) community belongs to the subhumid-

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WESTERN AREA

CENTRAL AREA

SOUTHERN AREA

Fig. 5.1 Distribution of the plant communities in the Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina). Solid yellow line: boundary of the Reserve. Dotted line (green): sampled areas. Associations: 1–13

humid riparian woodlands that occupy the lower Mesotropical belt (Association 13) (Fig. 4.9; Fig. 5.1). The floristic composition of the plant communities varies along the altitudinal gradient (Appendix C). A variety of species find their ecological optimum for their development in the communities in the lower Mesotropical belt under the bioclimatic conditions characteristic of this thermotype. Many are exclusive species that disappear at lower altitudes; this is also the case of a cluster of species typical of the upper Mesotropical belt that are absent at lower altitudes (Appendix C; Fig. 4.9). As mentioned earlier, these results show that the species are more abundant and more frequent within an optimum interval of ecological conditions along the altitudinal gradient, and that outside this optimum they decrease or disappear, thus affecting both the floristic composition and the species richness of the communities. The appearance and disappearance of species along environmental gradients, and particularly along altitudinal gradients, has also been reported in other studies on subtropical mountain woodlands in the region (Malizia et al. 2006; Brown et al. 2007; Blundo et al. 2012). This phenomenon implies that species appear and disappear within the communities as we ascend the altitudinal gradient in the Reserve, and a gradual exchange of species can be seen from one end of the gradient to the other and between bioclimatic belts. However, in the transition from the lower Mesotropical to the upper Mesotropical belt (at approximately 1260 m asl) there is a greater exchange of species in the tree layer, indicating a significant shift in

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environmental conditions and producing an important replacement of the tree cover within the mountain forest; of the 65 tree species recorded in the study area, 29 (44.6%) undergo an exchange as they go from one bioclimatic belt to another: 16 tree species present in the lower Mesotropical disappear (Carica quercifolia, Celtis ehrenbergiana var. discolor, Chloroleucon tenuiflorum, Condalia buxifolia, Geoffroea decorticans, Jacaranda mimosifolia, Manihot grahami, Myroxylon peruiferum, Prosopis alba, Randia micrantha, Schinus bumeloides, Schinus fasciculatus, Schinus myrtifolius, Sebastiania commersoniana, Senna spectabilis and Terminalia triflora) and 13 new species appear in the upper Mesotropical, of which six appear only in the mountain forest and disappear at a higher altitude (Aralia soratensis, Cedrela saltensis, Chrysophyllum marginatum, Cordia saccelia, Coutarea hexandra and Pisonia zapallo); five species are practically exclusive to the mountain woodland (Alnus acuminata, Ilex argentina, Podocarpus parlatorei, Prunus tucumanensis and Sambucus nigra ssp. peruviana); and two species can be found in both vegetation layers (Myrcianthes pungens and Stillingia tenella) (Appendix C). This also reveals that even in the upper Mesotropical belt there is a significant replacement of tree species. This exchange begins at the limit between the mountain forest and the mountain woodland (after 1488 m asl approximately) and points to a more pronounced variation in the environmental conditions above this altitude, mainly in regard to the decrease in temperature and the increase in humidity. It should be noted that some studies on subtropical mountain woodlands in Argentina report a greater exchange of tree species above 1500 m asl, in the transition from mountain forest to mountain woodland; that is, at a higher altitude than observed in this work (Cuyckens 2005; Malizia et al. 2006; Malizia et al. 2012; Martín 2014). Finally, the remaining tree species (36) are distributed throughout the whole of the altitudinal range in the study area. Most (29) have a low to moderate frequency (below 50%) and only a few species (7) occur with a high frequency (over 50%), and generally behave as dominant species in their respective communities (Appendices B and C). In contrast, the species exchange is less in the shrub layer: of the 85 shrub species recorded, 20 (23.5%) undergo replacement. In this regard, six species disappear when moving from the communities in the lower Mesotropical to the upper Mesotropical belt (Baccharis capitalensis, Baccharis coridifolia, Capsicum chacoense, Iresine diffusa, Senna occidentalis and Solanum palinacanthum—this last is endemic—); and 14 new shrub species appear (Aphelandra hieronymi, Austroeupatorium inulifolium, Baccharis latifolia, Campovassouria cruciata, Cnidoscolus vitifolius, Justicia kuntzei, Lepechinia vesiculosa, Piper hieronymi, Solanum abutiloides, Berberis jobii, Justicia mandonii, Ophryosporus lorentzii, Solanum aligerum and Solanum confusum—the last five are endemic–). The remaining shrub species (65) are distributed along the whole altitudinal gradient with differing frequencies, most with less than 20% and only two with more than 50% (Urera baccifera, 59% and Vernonanthura squamulosa, 66%) (Appendices B and C). The exchange is also lower in the herbaceous layer: of the 107 species recorded, 19 (17.7%) undergo replacement. In this regard, when moving from the communities in the lower Mesotropical belt to those of the upper Mesotropical belt, 11 species

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disappear (Bidens subalternans, Eleusine indica, Galium richardianum, Gamochaeta pensylvanica, Glandularia tweedieana, Leonurus japonicus, Solanum sisymbriifolium, Turnera sidoides, Verbascum virgatum, Verbena litoralis and Viguiera tucumanensis var. oligodonta) and aparecen 8 nuevas species (Acalypha boliviensis, Calceolaria teucrioides, Onoseris alata, Sibthorpia conspicua, Sibthorpia repens, Sinningia warmingii, Valeriana effusa and Petunia occidentalis—this last is endemic—). The remaining herbaceous species (89) are distributed along the whole altitudinal gradient with different frequencies. Most have a frequency of less than 20%, another fraction has between 20–40%, and only three species have frequencies of over 50%: Elephantopus mollis (52%), Jungia pauciflora (54%) and Justicia goudotii (54%), which behave as dominant in some communities, except the last (Appendices B and C). The species richness of the plant communities also reveals variations along the altitudinal gradient; however, they do not follow the general pattern observed for the altitudinal gradients in the Neotropical Kingdom (Gentry 2001), and particularly for the subtropical mountain woodlands in northwest Argentina, which show a decrease in species richness in relation to the increase in altitude (Morales et al. 1995; Brown et al. 2001; Malizia et al. 2006). As mentioned earlier, this is probably due to the fact that the altitudinal interval studied is not sufficiently broad to show a marked pattern of decrease in richness along the altitudinal gradient. Other factors that could play a role in the species richness of the communities must also be considered, such as their location, the number and size of the representative relevés, and their state of conservation, among others. It can be seen that the species richness of the communities in the lower Mesotropical belt varies from 93 species in the lowest community (Association 1: Enterolobio contortisilici-Anadenantheretum cebilis) to 183 species in the highest community in this layer (Association 8: Zanthoxylo cocoi-Blepharocalycetum salicifolii) (Table 4.21). The remaining communities distributed within this bioclimatic belt had between 45 and 138 species. Specifically, the communities with the lowest richness are represented by fewer relevés, some of which are smaller in size (500 m2), and located in more unstable and heterogeneous areas that probably restrict the confluence of a large number of species. This is the case of riparian communities (Association 13: Salici humboldtianae-Acacietum aromae, with 45 species) and of those located in foothills with sleep slopes and subject to rockfalls and landslides (known locally as “volcanoes”) during the period of greatest precipitation (Association 6: Schinetum myrtifolio-gracilipedis, with 46 species) (Table 4.21). Subsequently, when moving from the lower to the upper Mesotropical belt (between 1180–1300 m asl), the richness declines from 183 species to 127 (Association 9: Tecomo stantis-Anadenantheretum cebilis). Continuing along up the gradient, the richness once again increases to 165 species (Association 11: Cinnamomo porphyrium-Blepharocalycetum salicifolii) and finally decreases to 111 species above 1488 m asl (Association 12: Pruno tucumanensis-Podocarpetum parlatorei) (Table 4.21).

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These results reveal that there are two peaks of high species richness: one in the lower Mesotropical (with 183 species), and the other in the upper Mesotropical (with 165 species). Both peaks are present in mountain forest communities, whereas species richness tends to decrease in the transition to the mountain woodland, which is consistent with the findings of other studies that point to the mountain forest as the most diverse vegetation belt within the subtropical mountain woodlands. They also establish that the peaks of maximum species richness occur between 1000 and 1500 m asl (Morales et al. 1995; Brown et al. 2001; Cuyckens 2005; Malizia et al. 2006; Martín 2014). The analysis of the species richness of the vegetation layers in each bioclimatic belt revealed the presence of 48 tree, 70 shrub and 100 herbaceous species in the communities in the lower Mesotropical belt, whereas the communities in the upper Mesotropical have a total of 41 tree, 74 shrub and 89 herbaceous species. Although these values are not substantially different, they indicate certain characteristics of the communities growing in the two bioclimatic belts, mainly in terms of their state of conservation. The differences in richness in the herbaceous layer reflect the fact that the plant communities in the lower Mesotropical belt are more open and luminous, which favours the growth of a more extensive herbaceous cover containing more thermophilous species. This may indicate the poorer state of conservation of the vegetation. In contrast, the richness values for the communities in the upper Mesotropical belt are a reflection of the presence of closed, more humid woodland communities with deeper soils and less light, which determines the development of a less dense herbaceous cover with umbrophilous species and abundant pteridophytes. This denotes that these are well conserved communities, most probably because they are located in areas that are difficult for humans to access. The state of conservation of the communities in the study was also revealed by the high number of exotic and cosmopolitan species present in each bioclimatic belt; so the communities occupying the lower Mesotropical belt contain all the species recorded with this status (18), whereas only half are present in the upper Mesotropical belt, which is once again an indicator of the more favourable state of conservation of these last communities. The abundance-dominance and frequency of the species in the communities also varies along the altitudinal gradient (Appendix B). These results show again that the species are more abundant and more frequent within an optimum interval of environmental conditions, and that outside this interval they decrease or disappear. This therefore affects the floristic composition of the communities and the abundance-dominance values of the species that conform them (Matteucci and Colma 1982). The variations observed in the abundance-dominance of the species in the communities in the study area are determined by the prevailing environmental gradient in the area, and primarily the altitudinal gradient, which is in turn responsible for the variations in temperature and precipitation. Certain environments (e.g. steeply sloping sites, deep gorges and the beaches of rivers and streams) are also influenced by other factors such as geomorphology, soil type, and the orientation of the slopes. As a result, some species behave as dominant in one or more communities, while acting as companion species in others. Many of them have a

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broad interval of tolerance to environmental factors which allows them to be present in more sites along the altitudinal gradient and to be dominant in some communities. This is the case of 13 tree species (e.g. Acacia aroma, Allophylus edulis, Anadenanthera colubrina var. cebil, Blepharocalyx salicifolius, Celtis iguanaea, Cinnamomum porphyrium, Parapiptadenia excelsa, Schinus gracilipes, Scutia buxifolia, Sebastiania brasiliensis, Sebastiania commersoniana, Tipuana tipu and Vassobia breviflora); six shrub species (e.g. Barnadesia odorata, Dendrophorbium bomanii, Jungia polita, Senna pendula var. eriocarpa and Urera baccifera) and six herbaceous species (e.g. Elephantopus mollis, Jungia pauciflora, Mirabilisjalapa, Petiveria alliacea, Rivinia humilis and Samolus valerandi). Only a few species that behave as dominant have a more restricted distribution (e.g. Podocarpus parlatorei, Solanum aloysiifolium and Pharus lappulaceus) (Appendix C). Other species behave as exclusive, selective or preferential, or differential, and tend to be restricted to a single community where they find their ecological optimum, although they may well appear in other communities but with fairly insignificant or irrelevant abundance-dominance values from the phytosociological point of view. They include 32 tree species (e.g. Alnus acuminata, Aralia soratensis, Bougainvillea stipitata, Carica quercifolia, Cedrela angustifolia, Cedrela saltensis, Chloroleucon tenuiflorum, Duranta serratifolia, Enterolobium contortisiliquum, Erythrina falcata, Geoffroea decorticans, Ilex argentina, Jacaranda mimosifolia, Juglans australis, Kaunia lasiophthalma, Manihot grahami, Myrcianthes pseudomato, Myrcianthes pungens, Myroxylon peruiferum, Prosopis alba, Prunus tucumanensis, Sambucus nigra ssp. peruviana, Schinus bumeloides, Schinus fasciculatus, Schinus myrtifolius, Stillingia tenella, Tecoma stans, Terminalia triflora, Trema micrantha, Xylosma pubescens, Zanthoxylum coco and Zanthoxylum petiolare); 25 shrub species (e.g. Acalypha amblyodonta, Acalypha plicata, Aphelandra hieronymi, Austroeupatorium inulifolium, Baccharis coridifolia, Baccharis latifolia, Berberis jobii, Campovassouria cruciata, Capsicum chacoense, Clinopodium bolivianum, Justicia kuntzei, Justicia mandonii, Lepechinia vesiculosa, Ophryosporus lorentzii, Phenax laevigatus, Piper hieronymi, Senna occidentalis, Solanum aligerum, Solanum betaceum, Solanum confusum, Solanum palinacanthum, Thalictrum venturii, Verbesina macrophylla var. nelidae, Verbesina suncho and Vernonanthura pinguis; and 12 herbaceous species (e.g. Acalypha communis, Calceolaria teucrioides, Cantinoa mutabilis, Modiolastrum malvifolium, Onoseris alata, Oplismenus hirtellus, Petunia occidentalis, Phytolacca bogotensis, Sibthorpia conspicua, Stevia yaconensis var. subeglandulosa, Tibouchina paratropica and Turnera sidoides. This cluster also contains a few species that behave as stenoic and are generally indicators of the specific influence of some environmental factor. This is the case of riparian species such as Alternanthera philoxeroides, Asclepias curassavica, Baccharis salicifolius, Paspalum distichum Polygonum punctatum, Salix humboldtiana, Tessaria dodoneifolia and Tessaria integrifolia (Appendix C). Of the 257 species recorded in the whole of the study area, 103 (40%) are diagnostic (characteristic or indicator): 29 are dominant, 67 are distributed between exclusive, selective, preferential and differential and seven are stenoic. In addition, of these diagnostic species, 48 are trees (46.6%), 32 are shrubs (31.1%) and 23 are

5.5 Analysis, Interpretation and Characterisation of the Plant. . .

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herbaceous (22.3%) (Tables 3.3 and 3.4). It can be deduced from this that the tree layer has the highest percentage of diagnostic species, which would explain the diversity of woodland communities identified. The mountain forest communities have a higher percentage of diagnostic species (82.5%: 85 species) than the mountain woodland community (17.5%: 18 species). These differences could be related to the greater species richness found in the mountain forest, although the fact that most of the communities identified belong to this vegetation layer may also have an influence. The remaining species identified in the study area (152) behave as companions. Most appear in different communities along the whole of the altitudinal gradient. They are generally distributed in a discontinuous or scattered pattern and proved to have no diagnostic value (Appendix C). In terms of the current knowledge of the communities identified in the present study, it should be noted that they all are described for the first time for the province of Jujuy; twelve (12) are new to science and one (1) has been described for Bolivia. However, some of these communities show certain analogies with those described by other authors (Pinazo et al. 2003; Perea et al. 2008; Grau et al. 2010; Martín 2014) and they are probably also found in other localities and in similar bioclimatic conditions. The field data must therefore be extended in the future in order to refine the information obtained. In 1976 Cabrera highlighted the need to conduct phytosociological studies in Las Yungas to further the knowledge of the current diversity of communities and ecological variants, due to the large number of dominant species and to the different combinations occurring between them. The works of Martín (2014), Haagen Entrocassi (2014) and Martín et al. (2016) established the first precedent for phytosociological studies on the subtropical mountain woodlands in the province of Jujuy; the only previous similar work was conducted specifically in the mountain woodland of the southernmost Yungas in the province of Tucumán (Aceñaloza 1996). Specifically, the work of Martín (2014) covered the mountain forest and woodland at an altitudinal interval of 1320–1802 m asl. The communities relevéd include the description of Blepharocalyx salicifolius-Juglans australis (1407–1453 m asl), whose floristic and phytosociological similarities point to a possible correspondence with the association Juglandi australis-Blepharocalycetum salicifolii identified in the present study (Association 7; Cluster 3B; Table 3B). The work of Haagen Entrocassi (2014) studied the lower layer of the mountain forest in the Caulario river basin (935–1149 m asl), and none of the communities described there can be assimilated with the ones identified in the present study, as although they have some species in common, certain floristic elements typical of the foothill woodland give the vegetation its own singular character. It should be noted that the study area is geographically located between the areas studied by the abovementioned authors, so the three territories conform a band of woodlands running in a southwest-northeast direction starting in the Chijra river basin, crossing the Serranías de Zapla and concluding in the Caulario river basin; it covers not only the altitudinal but also the latitudinal gradient (Fig. 5.1). The present study and doctoral thesis provides a record of the vegetation growing in this belt, and

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thus makes an important contribution to the knowledge of a sector of subtropical mountain woodlands in the province of Jujuy.

5.6

Influence of the Environmental Gradient on the Composition and Distribution of the Plant Communities in the Study Area

The composition and distribution of plant communities is primarily determined by the altitudinal gradient prevailing in the study area, which in turn is responsible for the variations in temperature and precipitation and thus for the gradient of the thermicity (It) and ombrothermic (Io) indices. Taken as a whole, these gradients determine the variations in the floristic composition and distribution of the communities. This is defined by axis 1 in the ordination diagram obtained with Canonical Correspondence Analysis (CCA) (Fig. 5.1), which clusters on one side the communities representing the more open micro- and mesowoodlands growing in the low-lying warm subhumid areas of the Reserve in the lower Mesotropical-lower and upper Subhumid belt; these communities belong to the lowest altitudinal belt in the mountain forest (“basal forest”) and are characterised by thermophilous species that respond positively to high values of It (399–429) and low values of Io (4.7–5.5) and altitude (1015–1275 m asl) (Fig. 5.1, negative side of axis 1, Sectors I and II). Clustered on the other side are communities that represent the more closed woodlands or mesowoodlands, which grow at higher altitudes under more temperate and humid conditions in the upper Mesotropical-lower Humid belt; these communities belong to the highest layer of the mountain forest (“high-mountain forest”) and to the mountain woodland, and are generally characterised by less thermophilous and sciophilous species that respond positively to high values of altitude (1260–1620 m asl) and Io (7.7–8.7) and low values of It (352–395) (Fig. 5.1, positive side of axis 1, Sectors III and IV). In some situations, the influence of the altitudinal gradient is accompanied by the effect of the topographic exposure of the slopes, so both factors ultimately intervene in the changes in temperature and precipitation. Hence in subtropical latitudes where the sun is inclined towards the north, the west- and northwest-facing slopes are somewhat warmer and drier, as they receive more solar radiation and often tend be affected by what is known as the “rain shadow”, while those facing east and southeast tend to be cooler and more humid, as they have less radiation and receive the moist air masses from the Atlantic Ocean. However, the orientation of the slopes did not constitute a determining factor in the general distribution pattern of the communities in the Reserve: its influence is localised and can be noted particularly in certain communities due to the appearance of species that have their optimum in the drier areas of the foothill and Chacoan woodlands (e.g. Condalia buxifolia, Geoffroea decorticans and Prosopis alba). In some sectors with specific environmental characteristics, the effects of geomorphological and soil factors can also be seen to overlap with the influence of

5.6 Influence of the Environmental Gradient on the Composition and. . .

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1620 m asl 12

11

a 10

11

UPPER MESOTROPICAL / LOWER HUMID

9

8 7

f 8

LOWER MESOTROPICAL / UPPER SUBHUMID

6

e

LOWER MESOTROPICAL / UPPER SUBHUMID LOWER HUMID

8 9

5

d

4 2

b

3

Los Blancos stream

13

1

LOWER MESOTROPICAL / LOWER SUBHUMID

c 13 Zapla river 1015 m asl Pacará stream WESTERN AREA

CENTRAL AREA

SOUTHERN AREA

Fig. 5.2 Altitudinal zonation of the vegetation in the Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina). Meteorological station localities: (a) Mina 9 de Octubre; (b) Socavón; (c) Arroyo Pacará; (d) Las Capillas; (e) Algarrobal; (f) El Cucho. Phytosociological associations: (1) Enterolobio contortotisilici-Anadenantheretum cebilis; (2) Schino bumeloidis-Allophyletum edulis; (3) Xylosmo pubescentis-Blepharocalycetum salicifolii; (4) Jacarando mimosifoliaeVassobietum breviflorae; (5) Erythrino falcatae-Tipuanetum tipi; (6) Schinetum myrtifoliogracilipedis; (7) Juglandi australis-Blepharocalycetum salicifolii; (8) Zanthoxylo cocoiBlepharocalycetum salicifolii; (9) Tecomo stantis- Anadenantheretum cebilis; (10) Myrciantho pseudomatoi-Blepharocalycetum salicifolii; (11) Cinnamomo porphyrium-Blepharocalycetum salicifolii; (12) Pruno tucumanensis-Podocarpetum parlatorei; (13) Salici humboldtianaeAcacietum aromae

altitude. As a result of the influence of the environmental gradient—mainly the altitudinal gradient—, the communities are organised spatially according to the existing environmental variations and distributed on foothills, hillsides, gorges, ravines and edges of mountain ranges (terrestrial associations 1–12) and on river terraces and beaches (riparian association 13) (Fig. 5.1). In general terms, the following altitudinal zonation of the vegetation can be recognised in the study area (Fig. 5.2): the subhumid mesowoodlands of Enterolobio contortisilici-Anadenantheretum cebilis (Association 1) are found at the lower end of the altitudinal gradient, in the lower Mesotropical-lower Subhumid bioclimatic belt, and are distributed in the southern part of the study area between 1032–1037 m asl (It ¼ 426–427; Io ¼ 4.7) (Appendix A: Cluster 1A). Ascending along the altitudinal gradient the ombrotype changes to upper Subhumid, which is verified at approximately 1085 m asl. From this point on and until 1200 m asl, there is a presence of subhumid woodlands of Schino bumeloidis-Allophyletum edulis (Association 2), Xylosmo pubescentis-Blepharocalycetum salicifolii (Association 3) and Jacarando mimosifoliae-Vassobietum breviflorae (Association 4), growing on gently sloping hillsides and foothills, generally west-, northwest- and southwestfacing in the western part of the study area (It ¼ 408–422; Io ¼ 4.8–4.9) (Appendix

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A: Clusters 2B, 2C and 2A). Subsequently, in the same zone and at a higher altitude, these woodlands are replaced on high foothills and gently to moderately sloping hillsides, generally east-, northeast- and southeast-facing, by subhumid micro- and mesowoodlands of Erythrino falcatae-Tipuanetum tipi (Association 5), Schinetum myrtifolio-gracilipedis (Association 6) and Juglandi australis-Blepharocalycetum salicifolii (Association 7), which are distributed approximately between 1200–1258 m asl (It ¼ 401–407; Io ¼ 4.9–5.5) (Appendix A: Clusters 1C, 3A and 3B); also on hillsides with a humid ombrotype in the central part of the study area there is a further presence of the woodlands of Erythrino falcatae-Tipuanetum tipi (Association 5) but at a lower altitude (1115 m asl; It ¼ 412; Io ¼ 7.4), which extend approximately through to the transition with the upper Mesotropical belt at 1260 m asl. Finally, on the last section of the lower Mesotropical belt and in different topographical exposures in both the central and western part of the Reserve, there are subhumid mesowoodlands of Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8) growing on more steeply sloping hillsides at 1233–1275 m asl (It ¼ 399–403; Io ¼ 5–5.5) (Appendix A: Cluster 1B). On low-lying river terraces and beaches in the central-southern and southern zone of the Reserve (between 1015–1035 m asl), there are riparian microwoodlands of Salici humboldtianae-Acacietum aromae (Association 13) that extend within the lower Mesotropical belt under subhumid (It ¼ 429; Io ¼ 4.7) to humid ombrotypes (It ¼ 422; Io ¼ 7.2) (Appendix A: Cluster 6B). The presence of this association in environments with different ombrotypes indicates that its distribution responds mainly to the geomorphological and soil conditions typical of riparian environments, where the water table is shallow and accessible to the roots, and does not depend essentially on the moisture supplied by rainfall. The influence of the geomorphological and soil variables prevailing in the riparian environments in the study area can be seen in the location of this association in the ordination diagram obtained in the Canonical Correspondence Analysis; its characteristic species are associated to axis 2, revealing a relatively insignificant relation with the environmental variables (altitude, It and Io) (Fig. 4.9: Sector I). The central part, which contains the highest elevations in the study area, marks the beginning of the transition to the upper Mesotropical-lower Humid bioclimatic belt. This can be seen at 1260 m asl according to the values shown by the bioclimatic indices (It ¼ 395; Io ¼ 7.7). In this part of the Reserve the upper humid ombrotype is widely distributed on the east-, northeast- and southeast-facing slopes in both bioclimatic belts, whereas on slopes exposed to the northwest, west and southwest, the presence of this ombrotype is restricted only to the upper Mesotropical belt. From the aforementioned bioclimatic transition and up to 1310 m asl, there are humid mesowoodlands of Tecomo stantis-Anadenantheretum cebilis (Association 9) (It ¼ 389–395; Io ¼ 7.7–7.9) (Appendix A: Cluster 5C), which replaces those of Erythrino falcatae-Tipuanetum tipi (Association 5) located at lower elevations and with lower moisture requirements; it also descends as edaphohygrophilous through humid gorges until the lower Mesotropical belt (1180 m asl; It ¼ 396–404; Io ¼ 7.5–7.7). Ascending the gradient between 1320–1433 m asl, there is a presence of humid mesowoodlands of Cinnamomo porphyrium-Blepharocalycetum

5.7 General Diagnosis of the Plant Communities in the Study Area

127

salicifolii (Association 11) which occupy both the east-, northeast- and southeastfacing slopes and some northwest-facing slopes where the humid ombrotype persists given that it receives the last moisture-laden air masses coming over the lower summits of the mountain ranges, where they find temperate and humid conditions to prosper (It ¼ 374–387; Io ¼ 7.9–8.2) (Appendix A: Cluster 5B–5D); on the slopes, this woodland descends to the limit of the lower Mesotropical belt and contacts with the mesowoodlands of Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8) forming a broad ecotone. The woodlands of Cinnamomo porphyrium-Blepharocalycetum salicifolii coexist altitudinally with the humid mesowoodlands of Myrciantho pseudomatoi-Blepharocalycetum salicifolii (Association 10), which tend to occupy more steeply-sloping and usually shadier sites between 1319–1360 m asl (It ¼ 382–388; Io ¼ 7.9–8) (Appendix A: Cluster 5A). All these associations belong to the highest stratum of the mountain forest (“highmountain forest”). Finally, at the upper extreme of the altitudinal gradient, there is a presence of the mesowoodlands of Pruno tucumanensis-Podocarpetum parlatorei (Association 12), distributed mainly along the edges and boundaries of the mountain ranges between 1488–1620 m asl, and belonging to the highest layer corresponding to the mountain woodland and constituting the last band of tree vegetation (It ¼ 352–367; Io ¼ 8.3–8.7) (Appendix A: Cluster 4). The altitudinal zonation described shows the spatial distribution of the vegetation as a result of the general influence of the environmental variables considered in the present study (altitude, It and Io), to which can be added in certain sectors the local effects caused by geomorphological and soil factors which have not been analysed quantitatively here. It can therefore be concluded that the composition and distribution of the plant communities in the study area are determined by the environmental gradient prevailing in it. Within this complex of gradients it is the variations in altitude and their influence on the It and Io gradients that fundamentally determine the variability of the floristic patterns and spatial organisation of the plant communities.

5.7

General Diagnosis of the Plant Communities in the Study Area

The study of the vegetation in the Serranías de Zapla Multiple Use Ecology Reserve allowed us to identify and eliminate the most representative plant communities and establish their correspondence with the environmental gradient, principally determined by the variations in altitude and by the bioclimatic indices (It and Io). These communities were also typified from the phytosociological point of view as new associations with a provisional character, with the exception of one that has been described for Bolivia (Navarro and Maldonado 202). The diagnosis of each of the plant communities identified here is presented below, with a description of their main floristic, structural, distributional, bioclimatic and

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ecological characteristics, supported by the corresponding phytosociological tables. There is first a description of the terrestrial communities in the lower Mesotropical bioclimatic belt, followed by the communities that occupy the upper Mesotropical belt, ordered according to ascending intervals of altitude. The riparian community is described at the end of the section.

5.8

Terrestrial Vegetation in the Lower Mesotropical Bioclimatic Belt

1. Enterolobio contortisilici-Anadenantheretum cebilis ass. nova hoc loco (Table 4.8, holotypus rel. 5; Cluster 1A; Appendix C). It is distributed in the southern limit of the study area, in the lower-lying and warmer areas of the lower Mesotropical-lower subhumid belt (1032–1037 m asl; It ¼ 426–427; Io ¼ 4.7) (Appendix A: Cluster 1A; Fig. 4.9). It occupies foothills and gently-sloping low-lying hillsides with well-developed soils and without a marked preference for orientation (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open and semi-deciduous mesowoodland with an average height of 15 m. Four structural layers can be distinguished: the upper tree layer (canopy) is dominated by tall specimens of Anadenanthera colubrina var. cebil (cebil colorado) and Parapiptadenia excelsa (horco cebil), accompanied mainly by Tipuana tipu (tipa blanca) and Enterolobium contortisiliquum (pacará), and less frequently by Myroxylon peruiferum (quina), Terminalia triflora (lanza amarilla), Erythrina falcata (ceibo jujeño), Cedrela angustifolia (cedro coya) and Cinnamomum porphyrium (laurel del cerro). The arboreal understorey (lower tree layer) grows to a height of approximately 8 m and has a high species richness. It is dominated by Sebastiania brasiliensis (leche-leche), Sebastiania commersoniana (blanquillo), Vassobia breviflora (pucancho), Celtis iguanaea (tala pispa) and Acacia aroma (tusca). The four first species are abundant in woodland areas with indirect solar radiation, whereas Acacia aroma is common in the sunny margins of the woodland; these species are accompanied mainly by Xylosma pubescens (coronillo), Schinus bumeloides, Sapium haematospermum (lecherón) and Zanthoxylum petiolare (naranjillo); there is a less frequent presence of Acacia caven (espinillo), Carica quercifolia (sacha higuera), Allophylus edulis (chal-chal), Myrsine laetevirens (palo San Antonio), Schinus fasciculatus, Scutia buxifolia (nocán), Solanum riparium (fumo bravo), Tecoma stans (guaranguay), Bougainvillea stipitata (alfilerillo), Randia micrantha, Trema micrantha and Morus alba; this last is an exotic species that has propagated and become naturalised in Las Yungas due to the edible nature of its fruits, which are consumed and dispersed by wild fauna (mainly birds and herbivores) and also used by the inhabitants of the rural communities in the surrounding area.

5.8 Terrestrial Vegetation in the Lower Mesotropical Bioclimatic Belt

129

The shrubby understorey does not exceed a height of 2 m, although some individuals from species known as tall herbs grow to a height of 3 m. It generally has an open and disperse cover and is dominated by Urera baccifera (ortiguilla) accompanied mainly by Barnadesia odorata (clavillo), Solanum lorentzii and Vernonanthura squamulosa, and less frequently by Cestrum parqui (revienta caballos), Pavonia sepium, Verbesina macrophylla var. nelidae, Senna pendula var. eriocarpa, Clematis haenkeana, Baccharis capitalensis, Boehmeria caudata, Dendrophorbium bomanii and Smilax campestris (sacha rosa), among others. The herbaceous understorey has a denser cover and is richer in species. It is dominated by Rivinia humilis, Solanum aloysiifolium and Elephantopus mollis. Companion species are frequently Justicia goudotii, Petiveria alliacea, Tagetes terniflora, Sida cabreriana, Mikania micrantha, Ruellia erythropus, Galinsoga caracasana, Acalypha communis and Fleischmannia schickendantzii, among others; on rockfall sites with large rocks there is a frequent presence of colonies of Bromelia serra (chagua). The total species richness of this community is 93 species (Table 4.21), of which 30 are trees, 24 shrubs and 39 belong to the herbaceous layer. Of the 16 species selected as characteristic or indicator in this community, most (12) belong to the canopy and arboreal understorey: Seven species behave as dominant (Anadenanthera colubrina var. cebil, Parapiptadenia excelsa; Sebastiania brasiliensis, Sebastiania commersoniana, Vassobia breviflora, Celtis iguanaea and Acacia aroma); three are selective or preferential (Enterolobium contortisiliquum, Zanthoxylum petiolare and Carica quercifolia) and two are differential species (Myroxylon peruiferum and Terminalia triflora); the four remaining species come from the shrub (Urera baccifera) and herbaceous layer (Rivinia humilis, Solanum aloysiifolium and Elephantopus mollis). This forest community represents the vegetation in the warmer and less humid environments in the study area (Fig. 4.9); this can be seen by the presence of some thermophilous floristic elements that have their optimum distribution in the lower and warmer layer of the foothill woodland, located at a lower altitude outside the study area. Examples include Myroxylon peruiferum and Terminalia triflora, species that ascend to the mountain forest favoured by the warmer and subhumid conditions of the lower Mesotropical belt in the Reserve, where they finally find the altitudinal limit of their distribution. The significant presence of these two differential species, along with Enterolobium contortisiliquum, which also has a broad distribution in the foothill woodland (Cabrera 1994), and the absence of floristic elements characteristic of atmospheres with greater humidity (e.g. Juglans australis and Blepharocalyx salicifolius) would explain the distribution of this community in the ordination diagram in the Canonical Correspondence Analysis (Fig. 4.9) in response to the environmental conditions in the area it occupies, characterised by high thermicity indices and low altitudes and ombrothermic indices (Fig. 4.9: Sector I). In the dendrogram in the Hierarchical Classification Analysis (Fig. 4.6), this community shows greater floristic-phytosociological similarities with the mesowoodlands of Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8) and Erythrino falcatae-Tipuanetum tipi (Association 5). This may be due to the fact that they

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share some species, particularly in the upper tree layer, which generally occur with high frequencies (e.g. Parapiptadenia excelsa, Anadenanthera colubrina var. cebil and Tipuana tipu). Based on its structural characteristics, this woodland constitutes the mature stage of the forest community, essentially due to the existence of four well-defined vegetation layers and to its high richness in tree species. Although this woodland is generally in a good state of conservation, it contains some cleared sectors that indicate a certain degree of anthropic intervention before it became protected; this can be seen particularly from the ingression towards the interior of more heliophilous species that usually have their optimum in the sunny margins of the woodland (e.g. Acacia aroma and Acacia caven). This community marks the southern boundary of the Reserve. From this point on, with the descent in altitude and the presence of rural settlements, it coexists with a degraded vegetation mosaic and fields used for the cultivation of tobacco that have encroached on its distribution area, thereby reducing its representation until it is completely replaced below approximately 900 m asl. In the study of the vegetation in the Caulario river basin (Haagen Entrocassi 2014), located towards the northeast of the Reserve and separated from it by the Serranías de Zapla, a community is described that is also dominated by Anadenanthera colubrina var. cebil (1000–1050 m asl). However, this community reveals a significantly different floristic composition, as there is a more frequent presence of the characteristic species of the foothill woodland and even of the Chacoan biogeographic areas. The following are notable for their biological and/or forestry value: Cordia trichotoma (afata), Trichilia claussenii, Schinopsis marginata (horco quebracho), Ruprechtia apetala (manzano del campo), Prockia crucis, Handroanthus impetiginosus (lapacho rosado), Gleditsia amorphoides (espina corona), Cybistax antisyphilitica (lapacho verde), Croton piluliferus (tinajero), Cordia americana (lanza blanca), Coccoloba tiliacea, Ceiba chodatii (palo borracho), Astronium urundeuva (urundel), Achatocarpus praecox var. praecox (palo tinta), Acacia tucumanensis (garabato), Myroxylon peruiferum (quina) and Terminalia triflora (lanza amarilla). This community is distributed within a similar altitudinal interval to the one occupied by the community analysed here, so we might expect to find more floristic and phytosociological similarities between them, as both belong to the mountain forest. It is highly probable that the watershed imposed by the Serranías de Zapla acts as a barrier or filter to the arrival of foothill woodland species from the river basin, thus determining the absence of many of them from the Reserve and the fact that others reach their upper distribution limit and disappear towards the west. However, of all the communities identified in the study area, this is the one that absorbs most floristic elements from the foothill woodland due to its geographic location. This is a discrete and clearly defined community and does not therefore represent a vegetation type with a transitional character. 2. Schino bumeloidis-Allophyletum edulis ass. nova hoc loco (Table 4.9, holotypus rel. 2; cluster 2B; Appendix C). It is distributed throughout the lower-lying areas in the western part of the Reserve within the lower Mesotropical-upper subhumid belt (1085–1102 m asl;

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It ¼ 420–422; Io ¼ 4.8) (Appendix A: Cluster 2B; Fig. 4.9). It occupies broad gently-sloping foothills towards the south, generally with developed soils (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open deciduous microwoodland that grows to a height of 5–6 m. Three different structural layers can be distinguished: the tree layer is rich in species and dominated by Allophylus edulis (chal-chal), Vassobia breviflora (pucancho) and Acacia aroma (tusca). This last is abundant until the margins of the woodland, and accompanied mainly by Xylosma pubescens (coronillo), Condalia buxifolia (piquillín), Schinus bumeloides, Celtis iguanaea (tala), Sebastiania brasiliensis (leche-leche), Chloroleucon tenuiflorum (tatané), Manihot grahami (falsa mandioca) and Acacia caven (espinillo); and less frequently by Cnicothamnus lorentzii (azafrán), Senna spectabilis (carnaval), Jacaranda mimosifolia (jacarandá) and Geoffroea decorticans (chañar), among others. Taller trees with a dispersed distribution emerge above the level of the crowns in this microwoodland, but do not form a clearly defined layer (e.g. Enterolobium contortisiliquum, Anadenanthera colubrina var. cebil, Parapiptadenia excelsa, Tipuana tipu, Blepharocalyx salicifolius and Erythrina falcata). The shrubby understorey is more open and grows to a height of 1–1.5 m. It is dominated by Urera baccifera (ortiguilla) accompanied mainly by Chamissoa altissima, Vernonanthura squamulosa, Dendrophorbium bomanii and Smilax campestris, with a less frequent presence of Barnadesia odorata, Cestrum parqui, Senecio rudbeckiifolius, Capsicum chacoense, Lantana canescens, Clematis haenkeana and Senna pendula var. eriocarpa, among others. The herbaceous understorey has a denser cover and a high species richness. It is dominated by Elephantopus mollis, Mirabilis jalapa and Samolus valerandi, accompanied mainly by Praxelis clematidea, Modiolastrum malvifolium, Parthenium hysteriophorus, Tagetes terniflora, Justicia goudotii, Fleischmannia schickendantzii, Jungia pauciflora, Verbena litoralis, Hypochaeris microcephala, Tradescantia boliviana, among others; and less frequently by Viguiera tucumanensis var. oligodonta, Rivinia humilis, Bidens pilosa, Bromus catharticus, Cantinoa mutabilis, Desmodium subsericeum, Oenothera rosea, etc. The total species richness of this community is 99 species (Table 4.21), of which 27 are trees, 29 shrubs and 43 belong to the herbaceous layer. Of the 11 characteristic or indicator species in this community, six belong to the tree layer: three behave as dominant (Allophylus edulis, Acacia aroma and Vassobia breviflora), two are preferential (Xylosma pubescens and Schinus bumeloides) and one behaves as a differential species (Manihot grahami). The rest of the characteristic species are distributed between the shrub (Urera baccifera and Capsicum chacoense) and herbaceous layer (Elephantopus mollis, Mirabilis jalapa and Samolus valerandi). Although this community exists under a more humid ombrotype than the previous one, species can be found here that have their optimum in drier Chacoan environments, and which succeed in becoming established in the sunnier enclaves favoured by the orientation of the slopes. Some of these species appear as isolated individuals (Geoffroea decorticans) while others are more frequent (Condalia buxifolia).

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Although this community is rich in tree species, its structural characteristics indicate that it probably represents a secondary woodland (preclimax) resulting from the perturbation of an old mature woodland. This can be deduced above all from the development of a single tree layer and the existence of long-lived tall trees that emerge from the canopy in a disperse manner, some of which belong to species that appear with greater cover in the adjoining mesowoodland, where they dominate and contribute to its physiognomy (e.g. Blepharocalyx salicifolius). This community may derive from the alteration and reduction of the distribution area of the adjoining mesowoodland which develops higher in the altitudinal gradient (Xylosmo pubescentis-Blepharocalycetum salicifolii; assoc. 3, see Tables 4.21 and 4.12), with which it shares greater floristic-phytosociological similarities, as seen in the dendrogram in the Hierarchical Classification Analysis (Fig. 4.6). The subsistence of these tall trees in the community probably represents the vestiges or relics of a mesowoodland modified by past cutting campaigns which have led to the consequent alteration in its original composition, structure and distribution and very probably to the establishment of the current microwoodland as a result of the secondary succession. 3. Xylosmo pubescentis-Blepharocalycetum salicifolii ass. nova hoc loco (Table 4.10, holotypus rel. 8; cluster 2C; Appendix C). It is distributed at a greater elevation in the western part of the study area within the lower Mesotropical-upper Subhumid belt (1105–1127 m asl; It ¼ 417–420; Io ¼ 4.8). It limits with the previous community and replaces it on higher and steeper generally west-, northwest- and southwest-facing slopes with well-developed soils (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open and semi-deciduous seasonally evergreen mesowoodland with a height of approximately 12 m. It is differentiated into four structural layers. The upper tree layer (canopy) has a low species richness, and is dominated by Blepharocalyx salicifolius accompanied by Parapiptadenia excelsa. Erythrina falcata and Tipuana tipu are present with a very low frequency. Although this layer is represented by very few species, they conform a high woodland with a fairly continuous forest cover that confers the physiognomy to the community. The arboreal understorey (lower tree layer) grows to a height of approximately 6 m and is richer in species; it is dominated by Sebastiania brasiliensis (leche-leche) and Scutia buxifolia (nocán), accompanied mainly by Xylosma pubescens (coronillo) and Allophylus edulis (chal chal), and less frequently by Condalia buxifolia (piquillín), Sebastiania commersoniana (blanquillo), Celtis ehrenbergiana var. discolor (tala), Schinus bumeloides, Celtis iguanaea (tala pispa), Jacaranda mimosifolia (jacarandá) and Vassobia breviflora (pucancho), among others. The low frequency of Acacia aroma, a heliophilous species that normally ingresses in cleared and luminous sites within the woodland, has a more closed and umbrophilous plant cover. The shrubby understorey does not exceed a height of 2 m. It is open and discontinuous, and dominated by Urera baccifera (ortiguilla), accompanied mainly

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by Capsicum chacoense (ají kitucho), Cestrum parqui (revienta caballos), Senna occidentalis, Baccharis capitalensis, Barnadesia odorata (clavillo), Dendrophorbium bomanii, Senecio rudbeckiifolius, Senna pendula var. eriocarpa and Rubus imperialis (zarzamora), among others. The herbaceous understorey has greater species richness, although the species have low abundance-cover values and are distributed in a discontinuous manner without any particular species dominating, although there is a frequent presence of Mirabilisjalapa, Elephantopus mollis, Parthenium hysteriophorus, Petiveria alliacea, Cantinoa mutabilis, Oenothera rosea, Tagetes terniflora and Verbena litoralis. The soil is covered by plant remains and various species of pteridophytes. The total species richness of this community is 92 species (Table 4.21), of which 20 are trees, 29 shrubs and 41 belong to the herbaceous layer. Of the seven characteristic or indicator species in this community, four belong to the tree layer: three behave as dominant (Blepharocalyx salicifolius, Sebastiania brasiliensis and Scutia buxifolia), one is preferential (Xylosma pubescens) and the three remaining species are in the shrub layer (Urera baccifera, Capsicum chacoense and Senna occidentalis). This community does not have any species that behave as differential. In terms of its structural characteristics, mainly due to the presence of four clearly defined vegetation layers—two of which are arboreal (upper and lower)—, this mesowoodland represents the mature stage of the forest community. Its distribution area may have been broader and have extended into the lower-lying areas currently occupied by Schino bumeloidis-Allophyletum edulis (Association 2), which—as indicated—probably represents the degradation stage of this mesowoodland. Both communities have floristic and phytosociological affinities; for example they share some characteristics species of the lower understorey and the shrub layer (Xylosma pubescens, Urera baccifera and Capsicum chacoense) and another species that is practically exclusive to these two communities (Senna occidentalis). This similarity can be seen in the dendrogram of the Hierarchical Classification Analysis (Fig. 4.6) and in the ordination diagram of the Canonical Correspondence Analysis, where both communities are very close together and have a similar response to the environmental gradient, essentially at low values of altitude and It (Fig. 4.9: Sector I). The community described here may not have been exploited for the purposes of timber extraction due to its location on higher and steeper slopes, which hindered its access and exploitation. In the upper limit of its altitudinal distribution, this community is replaced by another woodland where there is also a predominance of Blepharocalyx salicifolius (Zanthoxylo cocoi-Blepharocalycetum salicifolii; Association 8). Similarly, this species is once again dominant in another two communities that will be described below, highlighting the abundance of this species in the lower Mesotropical belt in the study area. These observations coincide with previous reports from the study of a plot located at 1150 m asl in a sector of the Serranías de Zapla, where this is the most abundant species (83 individuals in one hectare) (Cuyckens 2005). In synthesis, the communities analysed until now represent the vegetation in the lower-lying, warmer and less humid zones of the study area, as indicated by the relation between the abundance and distribution of its species with high thermicity

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indices and low altitudes and ombrothermic indices (Fig. 4.9: Sector I). The three communities show some floristic similarities as they share similar environmental conditions. However, the mesowoodland of Enterolobio contortisiliciAnadenantheretum cebilis, which is located in the southernmost limit of the Reserve under a drier ombrotype, presents certain particularities due to its proximity to the foothill woodland. There are more floristic similarities between the subhumid mesotropical mesowoodland of Xylosmo pubescentis-Blepharocalycetum salicifolii and the microwoodland of Schino bumeloidis-Allophyletum edulis. Both are located one following the other in the low-lying areas of the western part of the Reserve. The first represents a mature forest community and the second very probably constitutes a secondary woodland as a result of the disturbance of the first. 4. Jacarando mimosifoliae-Vassobietum breviflorae ass. nova hoc loco (Table 4.11, holotypus rel. 4; Cluster 2A; Appendix C). It is distributed in the western part of the study area, in the lower Mesotropicalupper subhumid belt (1192–1200 m asl; It ¼ 408–409; Io ¼ 4.9) (Appendix A: Cluster 2A; Fig. 4.9). It occupies higher gently sloping foothills on less-developed soils, with stony deposits as a result of landslides originating on the adjacent slopes located at higher altitudes (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open and deciduous microwoodland that grows to a height of approximately 6 m. Three structural layers can be differentiated. The tree layer is dominated by Vassobia breviflora (pucancho), Celtis iguanaea (tala pispa) and Acacia aroma (tusca); this last species is very frequent both on the margins and in the interior of the woodland. It is mainly accompanied by Schinus fasciculatus, Jacaranda mimosifolia (jacarandá), Xylosma pubescens (nocán), Allophylus edulis (chal-chal), Chloroleucon tenuiflorum (tatané) and Sebastiania commersoniana (blanquillo), which are found with high frequencies. Present in a lower proportion are Sebastiania brasiliensis (leche-leche), Acacia caven (espinillo), Schinus bumeloides, Condalia buxifolia (piquillín), Geoffroea decorticans (chañar) and Prosopis alba (algarrobo). Adult individuals of Enterolobium contortisiliquum (pacará), Erythrina falcata (ceibo jujeño), Tipuana tip (tipa blanca), Parapiptadenia excelsa (horco cebil), Anadenanthera colubrina var. cebil (cebil colorado) and Eucalyptus sp. appear above the crown level in this layer in an isolated manner. Due to their sporadic presence they do not form a defined stratum or contribute to the physiognomy of the woodland. The shrubby understorey grows to a height of approximately 1.5–2 m. It is open and dispersed and dominated by Barnadesia odorata (clavillo) and Dendrophorbium bomanii, accompanied mainly by Cestrum parqui, Mimosa polycarpa, Vernonanthura squamulosa, Solanum lorentzii, Baccharis capitalensis and Baccharis microdonta; appearing less frequently are Buddleja stachyoides, Budleja diffusa, Iresine diffusa, Jungia polita, Malvastrum coromandelianum, Rubus imperialis, Senna pendula var. eriocarpa and Sida rhombifolia, among others. The herbaceous understorey is very open, and most of the species have low abundancecover values and form a clear cover where no particular species is dominant. The most

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frequent are Leonurus japonicus, Modiolastrum malvifolium, Parthenium hysteriophorus, Mikania micrantha, Jungia pauciflora, Mimosa xanthocentra, Bromus catharticus, Cantinoa mutabilis and Bidens pilosa, among others. The total species richness in this community is 79 species (Table 4.21), of which 23 are trees, 26 shrubs and 30 belong to the herbaceous layer. Of the 11 characteristic or indicator species in this community, eight belong to the tree layer: three behave as dominant (Vassobia breviflora, Celtis iguanaea and Acacia aroma) and five are selective or preferential (Schinus fasciculatus, Jacaranda mimosifolia, Chloroleucon tenuiflorum, Geoffroea decorticans and Prosopis alba). Most of these species prefer drier environments and possibly exploit less developed soils with a stony structure to become established within the community; the three remaining species are in the shrub (Barnadesia odorata and Dendrophorbium bomanii) and herbaceous layer (Modiolastrum malvifolium). This community does not have any differential species. This is the most impacted community in the study area, as it is close to a zone occupied by small rural settlements. Timber is extracted on a regular basis and there is a presence of cattle. The isolated trees that emerge from the tree canopy are likely remnants of a mature and intervened woodland, so it certainly represents a secondary woodland. According to the dendrogram in the Hierarchical Classification Analysis (Fig. 4.6, this community shows greater floristic and phytosociological similarities with the neighbouring woodlands of Schino bumeloidis-Allophyletum edulis (Association 2) and Xylosmo pubescentis-Blepharocalycetum salicifolii (Association 3) located at a lower elevation towards the east; whereas the ordination diagram in the Canonical Correspondence Analysis (Fig. 4.5) shows it in an intermediate position between the previous associations and Erythrino falcatae-Tipuanetum (Association 5), with which it limits on higher slopes. This reveals the gradual rise in the terrain in the western limit of the Reserve, along with a drop in temperature and a slight increase in humidity, as indicated in its bioclimatic indices. 5. Erythrino falcatae-Tipuanetum tipi ass. nova hoc loco (Table 4.12, holotypus rel. 5; Cluster 1C; Appendix C). It is distributed in the western part of the study area in the lower Mesotropicalupper subhumid belt and reappears in the central part in upper Subhumid and lower Humid ombrotypes. It is the only community in this bioclimatic belt that ascends to the areas with a humid ombrotype (1115–1258 m asl; It ¼ 401–412; Io ¼ 5–7.4) (Appendix A: Cluster 1C; Fig. 4.9). It occupies hillsides with a moderate slope, generally with developed soils, whereas in the central zone it descends to the foothills adjacent to the river terraces; it shows no marked preference in terms of slope exposure, although it is better represented on south-facing slopes (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open, semi-deciduous mesowoodland that grows to a height of approximately 15 m. Four structural layers can be distinguished. The upper tree layer (canopy) is dominated by Tipuana tipu (tipa blanca), accompanied by Erythrina

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falcata (ceibo jujeño), Parapiptadenia excelsa (horco cebil), Anadenanthera colubrina var. cebil (cebil colorado), Blepharocalyx salicifolius (horco molle) and Juglans australis (nogal criollo), and less frequently by Enterolobium contortisiliquum (pacará), Cinnamomum porphyrium (laurel del cerro) and Cedrela angustifolia (cedro coya). The arboreal understorey is diverse and semi-open, although it forms a more continuous cover than the canopy layer, growing to a height of approximately 8 m. It is dominated by Sebastiania brasiliensis (leche-leche) and Allophylus edulis (chalchal), accompanied mainly by Acacia aroma (tusca), Celtis iguanaea (tala pispa), Condalia buxifolia (piquillín), Sapium haematospermum (lecherón), Xylosma pubescens (coronillo), Tecoma stans (guaranguay), Acacia caven (espinillo) and Sebastiania commersoniana (blanquillo); there is a less frequent presence of Chloroleucon tenuiflorum (tatané), Vassobia breviflora (pucancho), Schinus myrtifolius, Senna spectabilis (carnaval), Celtis ehrenbergiana var. discolor (tala), Zanthoxylum coco (cochucho) and Scutia buxifolia (nocán), among others. In drier sites there is a presence of isolated individuals of Geoffroea decorticans (chañar) and Prosopis alba (algarrobo blanco). The shrubby understorey is rich in species and grows to a height of 2–3 m. It is dominated by Urera baccifera (ortiguilla) accompanied mainly by Barnadesia odorata, Vernonanthura squamulosa, Jungia polita, Vernonanthura pinguis, Acalypha amblyodonta and Senecio rudbeckiifolius; there is a less frequent presence of Smilax campestris, Sida rhombifolia, Mimosa debilis, Chamissoa altissima, Buddleja stachyoides, Cestrum parqui, Dolichandra ungis-cati, Baccharis coridifolia, Clematis haenkeana, Dendrophorbium bomanii, Solanum palinacanthum, Senna pendula var. eriocarpa, Baccharis capitalensis, Carica glandulosa, Budleja diffusa, Baccharis microdonta and Croton saltensis, among others. The herbaceous understorey has a high species richness, but it forms a more open and discontinuous cover. The species are generally present with low abundance-cover values, and none behaves as dominant. Notable for their frequency are Elephantopus mollis, Cantinoa mutabilis, Modiolastrum malvifolium and Bidens pilosa; less frequent are Salvia personata, Justicia goudotii, Jungia pauciflora, Tagetes terniflora, Tagetes filifolia, Bidens subalternans, Bromus catharticus, Hypochaeris microcephala, Leonurus japonicus, Cuphea racemosa, Viguiera tucumanensis var. oligodonta, Praxelis clematidea, Anredera cordifolia, Desmodium subsericeum and Galinsoga caracasana, among others. This community is very diverse with a total species richness of 140 species (Table 4.21), of which 39 are trees, 45 shrubs and 56 belong to the herbaceous layer. Of the 12 characteristic or indicator species in this community, five belong to the tree layer: three behave as dominant (Tipuana tipu, Sebastiania brasiliensis and Allophylus edulis) and two are preferential (Erythrina falcata and Prosopis alba); the seven remaining species are from the shrub (Urera baccifera, Acalypha amblyodonta, Baccharis coridifolia and Solanum palinacanthum) and herbaceous layer (Cantinoa mutabilis, Modiolastrum malvifolium and Turnera sidoides); this last behaves as a differential species.

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Due to its structural characteristics this community represents a mature mesowoodland. However, there are also cleared sectors where there was probably selective cutting of tall trees, although there are no signs of intensive cutting. It is worth noting that within the set of representative relevés of this community, three (relevés 20–22) show a certain degree of heterogeneity, as they have high or significant phytosociological values of Blepharocalyx salicifolius (horco molle) and Anadenanthera colubrina var. cebil (cebil colorado), species who have their optimum in other communities; however, these relevés display the general floristic and phytosociological characteristics of the community. Specifically, these relevés are located on more humid hillsides in contact with the mesowoodland of Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8), in which the aforementioned species are dominant. This mesowoodland is located at a higher altitude but descends as edaphohygrophilous through humid gorges and comes into contact with this community in some sectors on hillsides where the soils have a good water balance. Based on this, these could be considered as differential species for the ecotone formed by both communities, and their presence would reflect a humid variant of the type community analysed here, represented by the relevés indicated previously. However, more detailed studies are required to enable a better floristic and phytosociological interpretation of the relevéd sites, given the complexity of the environment where they are located. It is also worth noting that the high phytosociological values of Tecoma stans in relevé 106 indicate its proximity to the river terraces of the Zapla river, where this species forms part of the riparian microwoodland of Salici humboldtiana-Acacietum aromae (variant with Tecoma stans; Association 13); this can be seen in the ordination diagram of the Canonical Correspondence Analysis, where it is located close to the cloud representing this microwoodland (Fig. 4.9: Sector I). In general terms, this community is the one with the closest floristicphytosociological relations with the aforementioned mesowoodland of Zanthoxylo cocoi-Blepharocalycetum salicifolii, as can be seen from the dendrogram obtained in the Hierarchical Classification Analysis (Fig. 4.6). In addition, both communities are close or overlapping in the ordination diagram in the Canonical Correspondence Analysis, and have a similar response to the environmental gradient, which is reflected in the general conditions of greater elevation and humidity and the drop in temperature (Fig. 4.9: Sector II). A mesowoodland dominated by Tipuana tipu has been described by Martín (2014) for the vegetation in the middle basin of the Chijra river, but at a higher altitude in the upper Mesotropical-upper Subhumid belt (1422–1461 m asl). This community is not represented in the Reserve, although it is close to its western limit and presents significant differences that indicate that it is a different community to the one analysed here. The comparison of both communities reveals replacements in the floristic composition and variations in the phytosociological values of the species they share: for example, Tipuana tipu is dominant along with Parapiptadenia excelsa, while Erythrina falcata appears as a companion species in the canopy along with Juglans australis, Blepharocalyx salicifolius and Cinnamomum porphyrium. Important tree species are also absent, such as Anadenanthera

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colubrina var. cebil, Enterolobium contortisiliquum and Cedrela angustifolia; and other species appear such as Eupatorium lasiophthalmum, Sambucus nigra ssp. peruviana, Solanum betaceum, Solanum delitescens, Acalypha communis, Hymenostephium debile, Smallanthus macroscyphus, Chamissoa maximiliani, Mirabilis jalapa, Chaptalia nutans, Exostigma notobellidiastrum, Seemannia nematanthodes, Cuphea calophylla, Galinsoga mandonii, Bidens mandonii, Salvia rypara, Cyclopogon congestus and Cyclopogon elegans. To determine whether both communities come into contact at some point outside the limits of the Reserve, it will be necessary to conduct studies focused on the altitudinal band that separates them (approximately 150 m), which has not yet been relevéd. 6. Schinetum myrtifolio-gracilipedis ass. nova hoc loco (Table 4.13, holotypus rel. 2; Cluster 3A; Appendix C). It is distributed in the western part of the study area, in the lower Mesotropicalupper Subhumid belt (1235–1243 m asl; It ¼ 402–403; Io ¼ 5,5) (Appendix A: Cluster 3A; Fig. 4.9). It occupies some sections in foothills at higher elevations and gently sloping hillsides with developed soils (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open to semi-closed deciduous microwoodland that grows to a height of approximately 5–6 m. Three different layers can be differentiated structurally. The tree layer is dominated by Schinus gracilipes, Allophylus edulis (chal-chal) and Sebastiania brasiliensis (leche-leche), accompanied mainly by Schinus myrtifolius, Sebastiania commersoniana (blanquillo), Vassobia breviflora (pucancho) and Condalia buxifolia (piquillín); there is a less frequent presence of Chloroleucon tenuiflorum (tatané), Scutia buxifolia (nocán) and Xylosma pubescens (coronillo); there is an isolated presence of Acacia aroma (tusca) and young individuals of Anadenanthera colubrina var. cebil (cebil colorado) emerging from the canopy. The shrubby understorey is open and grows to a height of approximately 1–1.5 m. No species is dominant, although there is a notable presence of Jungia polita, Rubus imperialis and Vernonanthura squamulosa, and a less frequent and disperse presence of Malvastrum coromandelianum, Baccharis microdonta, Buddleja stachyoides, Clematis haenkeana, Croton saltensis, Dendrophorbium bomanii, Senna pendula var. eriocarpa and Urera baccifera, among others. The herbaceous understorey is also open and forms a discontinuous cover; its most frequent species include Jungia pauciflora, Zinnia peruviana, Eleusine indica, Mirabilis jalapa and Praxelis clematidea, accompanied by Fleischmannia schickendantzii, Hypochaeris microcephala, Bidens subalternans, Justicia goudotii, Parthenium hysteriophorus and Tagetes terniflora, among others. This is the least represented and least diverse terrestrial community. Its total species richness is 46 (Table 4.21), of which 12 are trees, 17 belong to the shrub layer and a similar number are present in the herbaceous layer. It has four characteristic or indicator species and all belong to the tree layer: three species behave as dominant (e.g. Schinus gracilipes, Allophylus edulis and Sebastiania brasiliensis) and one as preferential (e.g. Schinus myrtifolius). In spite of its low species richness

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it conforms a clearly defined community with a dense tree cover that limits the entry of light and determines the development of a sparse and disperse shrubby and herbaceous understorey. Structurally it is the only community in the lower Mesotropical belt that has an extremely close and continuous forest cover. The dendrogram of the Numerical Classification Analysis (Fig. 4.6) shows the floristic-phytosociological similarities of this community with the mesowoodland of Juglandi australis-Blepharocalycetum salicifolii (Association 7) with which it coexists altitudinally in areas of high foothills; for example, they share some characteristic species (Schinus myrtifolius, Allophylus edulis and Sebastiania brasiliensis). The two last species are dominant and contribute to the physiognomy of this community, and in turn also dominate the lower tree layer of the mesowoodland of Juglandi australis-Blepharocalycetum salicifolii. The aforementioned similarities can be seen in the ordination diagram in the Canonical Correspondence Analysis, where both communities overlap and are similarly related with the environmental variables (Fig. 4.9, Sector II). This community also probably represents a secondary forest (preclimax) as a result of the intensive cutting of mature woodland, of which there are no long-lived or tall remnants; the presence of the few young individuals of Anadenanthera colubrina var. cebil (cebil colorado), a species that grows to a great height in its adult state, indicate the gradual ingression and development of tree species from the neighbouring mesowoodlands. It is therefore possible that as a result of secondary succession this community may have substituted the adjoining mesowoodland of Juglandi australis-Blepharocalycetum salicifolii in the foothills and gentle slopes that are more accessible to exploitation. 7. Juglandi australis-Blepharocalycetum salicifolii ass. nova hoc. loco (Table 4.14, holotypus rel. 9; Cluster 3B; Appendix C). It is distributed throughout the western edge of the study area, in the lower Mesotropical-upper Subhumid belt (1239–1253 m asl; It ¼ 401–402; Io ¼ 5.5) (Appendix A: Cluster 3B; Fig. 4.9). It occupies gentle slopes with different exposures, generally on deep soils. It has a high frequency and is more widespread in east-, northeast- and southeast-facing areas (Figs. 5.1 and 5.2); in some higher sectors it substitutes the previous association (Schinetum myrtifolio-gracilipedis), while in others it coexists altitudinally with it. From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open semi-deciduous seasonally evergreen mesowoodland that grows to a height of approximately 15–18 m. Four well-defined structural layers can be differentiated. The upper tree layer (canopy) is dominated by tall specimens of Juglans australis (nogal criollo) and Blepharocalyx salicifolius (horco molle), accompanied by isolated individuals of Anadenanthera colubrina var. cebil (cebil colorado), Erythrina falcata (ceibo jujeño), Parapiptadenia excelsa (horco cebil) and Tipuana tipu (tipa blanca). The arboreal understorey has a greater species richness and grows to a height of approximately 8 m. It is dominated by Sebastiania brasiliensis (leche-leche) and Allophylus edulis (chal-chal), mainly accompanied by Vassobia breviflora (pucancho), Scutia buxifolia (nocán), Condalia buxifolia (piquillín), Schinus

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gracilipes, Schinus myrtifolius, Celtis iguanaea (tala pispa), Sebastiania commersoniana (blanquillo) and Xylosma pubescens (coronillo); there are also several isolated individuals of Celtis ehrenbergiana var. discolor (tala), Senna spectabilis (carnaval), Acacia aroma (tusca), Morus alba, Chloroleucon tenuiflorum (tatané) and Sapium haematospermum (lecherón). The shrubby understorey is open to semi-open and attains a height of approximately 1.5–2 m; it is dominated by Jungia polita and Senna pendula var. eriocarpa, accompanied by Vernonanthura squamulosa, Chamissoa altissima, Dendrophorbium bomanii, Heteropterys sylvatica, Malvastrum coromandelianum and Rubus imperialis; there is a less frequent presence of Acalypha amblyodonta, Barnadesia odorata, Mimosa debilis, Smilax campestris, Urera baccifera, Baccharis microdonta, Clematis haenkeana, Baccharis coridifolia, Buddleja stachyoides and Croton saltensis, among others. The herbaceous understorey is open and dominated by Jungia pauciflora accompanied by Zinnia peruviana, Cantinoa mutabilis, Justicia goudotii, Mikania micrantha, Praxelis clematidea, Fleischmannia schickendantzii, Gorgonidium vermicidum, Hypochaeris microcephala, Galinsoga caracasana, Mirabilis jalapa, Parthenium hysteriophorus and Tagetes terniflora; there is also an abundant presence of ferns in this layer. The soil is scattered with plant detritus and the rocky outcrops are covered by various moss species. This community does not have a high diversity compared to other mesowoodlands in the area. Its total species richness is 74 species (Table 4.21), of which 22 are trees, 24 shrubs and 28 belong to the herbaceous layer. Of the eight characteristic or indicator species in this community, five belong to the tree layer: four behave as dominant (Juglans australis, Blepharocalyx salicifolius, Sebastiania brasiliensis and Allophylus edulis) and one is preferential (Schinus myrtifolius). The three remaining species are from the shrub (Jungia polita and Senna pendula var. eriocarpa) and herbaceous layer (Jungia pauciflora); no species behaves as differential. It is worth noting that despite the fact they contain practically none of the canopy species, relevés 41 and 38 have been assimilated to this community based mainly on the results obtained in the Hierarchical Classification Analysis (Fig. 4.6), which clusters them within the set of relevés that are representative of the community. However, it is debatable whether these relevés belong phytosociologically to this community; they thereefore have a preliminary character, as they have floristicphytosociological similarities with both the arboreal understorey in this community and with the adjoining microwoodland (Schinetum myrtifolio-gracilipedis; Association 6) described above. This community represents a mature woodland that has been relatively well conserved and certainly not been subjected to intensive cutting, although there has probably been clear-cutting of valuable forestry specimens. However, it contains tall trees that have not been extracted due to the more complex topography of the terrain. It is also probable that its distribution area was previously more extensive and reached as far as the sites that are currently occupied by the microwoodland of Schinetum myrtifolio-gracilipedis, with which it shares floristic and phytosociological affinities as indicated earlier.

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In the middle basin of the Chijra river, Martín (2014) described a mesowoodland of Blepharocalyx salicifolius-Juglans australis growing at a higher altitude in a band (between 1407–1453 m asl) on southeast- and east-facing slopes. This woodland shares floristic and phytosociological characteristics with the community analysed here; specifically, the most significant similarities occur in the tree layer (canopy and lower). It is thus very likely that this is a single community that has a broader and discontinuous distribution area: this would mark the lower altitudinal limit of its distribution in the Reserve, and it would be replaced at a higher altitude by the mesowoodland of Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8). Outside the Reserve and towards the west, it would reappear at a higher elevation in the Chijra river basin, where according to Martín (2014) the dominant species in the tree canopy (Blepharocalyx salicifolius and Juglans australis) are accompanied by Erythrina falcata, Parapiptadenia excelsa, Cinnamomum porphyrium and isolated specimens of Cedrela angustifolia. In the arboreal understorey there is a dominance of Allophyllus edulis, accompanied mainly by two species with an optimum at a higher altitude: Sambucus nigra ssp. peruviana and Eupatorium lasiophthalmum. Appearing with less frequency are Condalia buxifolia, Myrsine laetevirens, Sebastiania commersoniana, Solanum betaceum and Xylosma pubescens; in the shrubby understorey they are mainly present in Celtis ehrenbergiana, Senna pendula, Vernonanthura pinguis and Urera baccifera, whereas the herbaceous understorey is dominated by Mirabilis jalapa and Oplismenus hirtellus, accompanied by Acalypha communis, Chaptalia nutans, Cuphea calophylla, Elephantopus mollis, Exostigma notobellidiastrum, Galinsoga mandonii, Gorgonidium vermicidum, Jungia pauciflora, Rivina humilis and Sida cabreriana, among others (Martín 2014). 8. Zanthoxylo cocoi-Blepharocalycetum salicifolii ass. nova hoc loco (Table 4.15, holotypus rel. 11; cluster 1B; Appendix C). It is distributed in the central and western area of the study area, within the lower Mesotropical-upper Subhumid belt (1233–1275 m asl; It ¼ 399–403; Io ¼ 5–5,5) (Appendix A: Cluster 1B; Fig. 4.5). It occupies steeply sloping hillsides with different exposures, generally on developed soils (Figs. 5.1 and 5.2); it replaces the mesowoodland of Juglandi australis-Blepharocalycetum salicifolii (Association 7) on slopes with a higher elevation, gradient and insolation, whereas in other sectors it coexists altitudinally with it; this community also descends as edaphohygrophilous through humid gorges and terraces where it contacts with the mesowoodland of Erythrino falcatae-Tipuanetum tipi (Association 5). From the phytogeographical point of view it belongs to the mountain forest district (“basal forest”) (Cabrera 1994). This is a semi-open mesowoodland, although in some sectors it has a more continuous cover. It is semi-deciduous to seasonally evergreen, and grows to a height of approximately 15 m, with some taller emergent trees. Four well-defined structural layers can be differentiated: the upper tree layer (canopy) is dominated by tall specimens of Blepharocalyx salicifolius (horco molle) and Anadenanthera colubrina var. cebil (cebil colorado), accompanied by Tipuana tipu (tipa blanca),

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Juglans australis (nogal criollo), Cinnamomum porphyrium (laurel del cerro), Parapiptadenia excelsa (horco cebil) and Erythrina falcata (ceibo jujeño); isolated individuals of Cedrela angustifolia (cedro coya) and Enterolobium contortisiliquum (pacará) appear with a very low frequency. The arboreal understorey is semi-open and grows to a height of approximately 7–8 m. It has a high species richness but is not dominated by any particular species. Appearing with high frequencies are Celtis iguanaea (tala pispa), Sebastiania brasiliensis (leche-leche), Vassobia breviflora (pucancho), Allophylus edulis (chalchal), Zanthoxylum coco (cochucho), Scutia buxifolia (nocán), Sebastiania commersoniana (blanquillo), Schinus myrtifolius, Celtis ehrenbergiana var. discolor (tala), Acacia aroma (tusca), Acacia caven (espinillo) and Tecoma stans (guaranguay). The last three species are abundant in the luminous edges of the woodland; present with a lower frequency are Chloroleucon tenuiflorum (tatané), Condalia buxifolia (piquillín), Sapium haematospermum (lecherón), Senna spectabilis (carnaval), Myrsine laetevirens (palo San Antonio), Solanum riparium (fumo bravo), Cnicothamnus lorentzii (azafrán), Kaunia lasiophthalma (malvón), Schinus gracilipes, Randia micrantha, Escallonia millegrana and Xylosma pubescens (coronillo). The shrubby understorey is open to semi-open and grows to a height of 2–3 m, and has a high diversity; it is dominated by Barnadesia odorata (clavillo) and Urera baccifera (ortiguilla), accompanied mainly by Vernonanthura squamulosa, Acalypha amblyodonta, Boehmeria caudata, Clinopodium bolivianum (muñamuña), Jungia polita, Chamissoa altissima, Heimia montana, Abutilon grandifolium, Croton saltensis and Cestrum parqui; with a less frequent presence of Buddleja stachyoides, Carica glandulosa, Lantana canescens, Senecio rudbeckiifolius, Senna pendula var. eriocarpa, Smilax campestris, Verbesina macrophylla var. nelidae, Chromolaena laevigata, Hebanthe occidentalis, Manetia jorgensenii, Heteropterys sylvatica, Lantana trifolia, Mimosa polycarpa, Rubus imperialis, Vernonanthura pinguis, Achyrocline fláccida, Weddelia saltensis and Acalypha plicata, among others. The herbaceous understorey is also open to semiopen, but is highly diverse and has a greater species richness than the shrub layer; it is dominated by Jungia pauciflora, mainly accompanied by Tagetes terniflora, Elephantopus mollis, Cortaderia selloana and Justicia goudotii; present with less frequency are numerous species such as Salvia personata, Urtica chamaedryoides, Petiveria alliacea, Tibouchina paratropica, Fleischmannia schickendantzii, Adenostemma brasilianum, Rivinia humilis, Mikania micrantha, Anredera cordifolia, Cortaderia hieronymi, Axonopus compressus, Phytolacca bogotensis, Dicliptera squarrosa, Bidens pilosa, Begonia boliviensis var. boliviensis, Cajophora hibiscifolia, Conyza sumatrensis, Digitaria insularis, Solanum tenuispinum, Acalypha communis and Oenothera rosea, among many others. This is the community with the greatest diversity in the whole of the study area; its total species richness is 183 species (Table 4.21), of which 40 are trees, 44 shrubs and 99 belong to the herbaceous layer. Of the seven characteristic or indicator species in this community, three belong to the tree layer: two behave as dominant (Blepharocalyx salicifolius and Anadenanthera colubrina var. cebil) and one as

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preferential (Zanthoxylum coco); the remaining species come from the shrub (Barnadesia odorata and Urera baccifera) and herbaceous layer (Jungia pauciflora and Thalictrum venturii). As indicated at the start, this woodland descends as edaphohygrophilous through some gorges and sectors with greater availability of water (slopes and streams), and as a result the relevés taken on these sites include certain indicator species of these environments in the underlying floristic cluster in this community (e.g. Salix humboldtiana, Tessaria integrifolia, Baccharis salicifolius and Tessaria dodoneifolia). On low-lying differences in level on hillsides with outcrops of soil and rocks, or on accumulated small areas of scree, there is a frequent presence of two species that ingress from the adjoining ravines where they form dense azonal communities (e.g. Cortaderia selloana and Cortaderia hieronymi). However, until more detailed studies can be done, the presence of these species is not considered an indicator of a facie, as the relevés in which both were recorded do not reveal any particular floristic combination that deviates from the typical combination for this community; in view of this, it is only possible at present to link the appearance of these species with the exploitation of patches of more unstable soil in the interior of the woodland. This mesowoodland substitutes the woodland of Juglandi australisBlepharocalycetum salicifolii (Association 7) on slopes with a higher elevation and gradient. The replacement is more pronounced in sunny exposures where there is a decrease in the abundance-cover of Juglans australis (nogal criollo), which tends to prefer slopes with less insolation (generally east-, northeast- and southeast-facing). In parallel there is a significant increase in the frequency and phytosociological values of Anadenanthera colubrina var. cebil (cebil colorado), which along with Blepharocalyx salicifolius becomes dominant in the community.

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Terrestrial Vegetation in the Upper Mesotropical Bioclimatic Belt

9. Tecomo stantis-Anadenantheretum cebilis ass. nova hoc loco (Table 4.16, holotypus rel. 7; Cluster 5C); Appendix C). It is distributed in the central part of the study area, within the upper Mesotropicallower Humid belt and in the transition to the lower Mesotropical under the same ombrotype (1260–1310 m asl; It ¼ 389–395; Io ¼ 7,7–7.9) (Appendix A: Cluster 5C; Fig. 4.5). It occupies high ravines and hillsides with a gentle to moderate slope, in different exposures and generally on developed soils (Figs. 5.1 and 5.2). This community also descends as edaphohygrophilous through humid gorges and ravines until the lower Mesotropical belt (relevés 107–109), where it contacts with the mesowoodland of Erythrino falcatae-Tipuanetum tipi (Association 5), substituting it completely in the higher slopes in the upper Mesotropical belt. From the phytogeographical point of view it belongs to the mountain forest district (“high-mountain forest”) (Cabrera 1994).

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This is an open to semi-open semi-deciduous mesowoodland that grows to a height of approximately 10 m, with some taller emergent specimens. Four structural layers can be differentiated. The upper tree layer (canopy) has cleared areas and is poor in species, and is dominated by Anadenanthera colubrina var. cebil (cebil colorado) accompanied by Tipuana tipu (tipa blanca), Erythrina falcata (ceibo jujeño) and Parapiptadenia excelsa (horco cebil). There is an isolated presence of Cedrela angustifolia (cedro coya) and Enterolobium contortisiliquum (pacará). The arboreal understorey is open, although it is more disperse on some sites and semi-open on others, and grows to a height of approximately 6 m. It also has a low species richness and is not dominated by any particular species. It is characterised by the presence of Tecoma stans (guaranguay), Sebastiania brasiliensis (leche-leche) and Acacia aroma (tusca), which are accompanied less frequently by Allophylus edulis (chal-chal), Trema micrantha, Stillingia tenella, Aralia soratensis (sacha paraíso), Celtis iguanaea (tala pispa), Duranta serratifolia, Schinus gracilipes, Zanthoxylum petiolare (naranjillo), Chrysophyllum marginatum (aguaí), Cordia saccelia, Coutarea hexandra (dominguillo), Kaunia lasiophthalma (malvón) and Pisonia zapallo (zapallo caspi). The shrubby understorey is very open and disperse and grows to a height of approximately 2 m. It has a high species richness; it is not dominated by any particular species, but there is a high frequency of tall herbs such as Vernonanthura pinguis and Verbesina macrophylla var. nelidae, and other species such as Verbesina suncho, Acalypha plicata, Justicia kuntzei, Lantana canescens, Lantana trifolia, Vernonanthura squamulosa, Dendrophorbium bomanii, Solanum lorentzii, etc. There is a less frequent presence of Clematis haenkeana, Chamissoa altissima, Dolichandra ungis-cati, Heteropterys sylvatica, Budleja diffusa, Chromolaena laevigata, Hebanthe occidentalis, Jungia polita, Manetia jorgensenii, Mimosa debilis, Pavonia sepium, Phenax laevigatus, Rubus imperialis and Senecio rudbeckiifolius, among many others. The herbaceous understorey is open to semiopen and also rich in species; it is dominated by Rivinia humilis, Petiveria alliacea and Pharus lappulaceus, accompanied mainly by Oplismenus hirtellus, Adenostemma brasilianum, Salvia personata, Jungia pauciflora, Acalypha communis, Anredera cordifolia, Solanum aloysiifolium, Urtica chamaedryoides, Digitaria insularis, among others; and less frequently by Fleischmannia schickendantzii, Desmodium affine, Acalypha boliviensis, Duchesnea indica, Pseudechinolaena polystachya, Salpichroa origanifolia, Sida cabreriana, Dicliptera squarrosa, Justicia goudotii, Phytolacca bogotensis, Primula malacoides, Chaptalia nutans, Cuphea racemosa, Ruellia erythropus, Scoparia ericacea and Tragia volubilis, among others. This community has a high diversity represented in its shrub and herbaceous layer; its total species richness is 127 species (Table 4.21), of which 22 are trees, 52 shrubs and 53 belong to the herbaceous layer. Of the ten characteristic or indicator species in this community, three belong to the tree layer: one behaves as dominant (Anadenanthera colubrina var. cebil) and two as preferential (Tecoma stans and Trema micrantha); the remaining species are from the shrub (Verbesina suncho, Vernonanthura pinguis and Verbesina macrophylla var. nelidae) and

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herbaceous layer (Petiveria alliacea, Rivinia humilis, Pharus lappulaceus and Oplismenus hirtellus). This mesowoodland shows signs of anthropic impact and does not represent the mature stage of the forest community, although it can clearly be seen to be in recovery due to its legal protection status. This is indicated by the generally low abundance-cover values of the species in the canopy and the lower tree layer. As it has a more open structure that favours the entry of light in the understorey, there has been an ingression and propagation of heliophilous species that contribute to the physiognomy of the community. This can be seen mainly from the frequent presence of Tecoma stans (guaranguay) in the interior of the woodland. This species is widely distributed in several environments in the subtropical mountain woodlands in the region, and is frequent in the marginal vegetation associated to riparian environments, and on the edges and cleared areas of the woodlands; it also behaves as a pioneer, forming secondary woodlands in areas where the vegetation has been altered (Cabrera 1994; Navarro and Maldonado 2002; Grau et al. 2010; Martín 2014; Haagen Entrocassi 2014). 10. Myrciantho pseudomatoi-Blepharocalycetum salicifolii ass. nova hoc loco (Table 4.17, holotypus rel. 7; Cluster 5A; Appendix C). It is distributed in the central part of the study area, within the upper Mesotropical-lower Humid belt (1319–1360 m asl; It ¼ 382–388; Io ¼ 7,9–8) (Appendix A: Cluster 5A; Fig. 4.9). It coexists altitudinally with the mesowoodland of Cinnamomo porphyrium-Blepharocalycetum salicifolii (Association 11), substituting it on shadier and more steeply-sloping sites preferentially oriented towards the east, northeast and southeast (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (“high-mountain forest”) (Cabrera 1994). It constitutes a dense seasonally evergreen mesowoodland, completely closed in some sectors and growing to a height of approximately 10 m. Four well-defined structural layers can be differentiated: the upper tree layer (canopy) is dominated by Blepharocalyx salicifolius (horco molle) and Cinnamomum porphyrium (laurel del cerro), although the first predominates totally in some sites; they are accompanied by Parapiptadenia excelsa (horco cebil), and there is a scarce presence of Cedrela angustifolia (cedro coya) and Anadenanthera colubrina var. cebil (cebil colorado), whereas Erythrina falcata (ceibo jujeño), Juglans australis (nogal criollo) and Tipuana tipu (tipa blanca) occur in an isolated manner. The arboreal understorey is semi-open and grows to a height of approximately 5–6 m; it is dominated only by Myrcianthes pseudomato (mato arrayán), accompanied by Myrsine laetevirens (palo San Antonio), Allophylus edulis (chal-chal) and Sebastiania brasiliensis (leche-leche); there is a scarce presence of Prunus tucumanensis (palo luz), Cnicothamnus lorentzii (azafrán), Kaunia lasiophthalma (malvón), Myrcianthes pungens, Solanum riparium (fumo bravo), Xylosma pubescens (coronillo), Pisonia zapallo (zapallo caspi) and Schinus gracilipes. The shrubby understorey is disperse and stands no higher than one metre, except in a few isolated cases. It has low species richness and is not dominated by any

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particular species, although there is a greater frequency of Smilax campestris, Acalypha plicata and Justicia mandonii, accompanied mainly by Justicia kuntzei, Rubus imperialis and Verbesina suncho; present with a lower frequency are Dolichandra ungis-cati, Vernonanthura pinguis, Chamissoa altissima, Lantana canescens, Verbesina macrophylla var. nelidae, Acalypha amblyodonta, Acalypha lycioides, among others. There is an isolated presence of Vernonanthura squamulosa, Baccharis microdonta, Boehmeria caudata, Chiropetalum boliviense, Hebanthe occidentalis, Heteropterys sylvatica, Mimosa polycarpa, Muehlenbeckia sagittifolia, among others. The herbaceous understorey is very open and has a lower species richness than the shrubby understorey; it is characterised by the frequent presence of Justicia goudotii and Bromelia serra, accompanied by Petiveria alliacea, Acalypha communis, Pharus lappulaceus, Rivinia humilis, Cortaderia hieronymi, Jungia pauciflora, Mimosa xanthocentra, Deyeuxia polígama and Panicum trichanthum; there is an isolated presence of Acalypha boliviensis, Adenostemma brasilianum, Fleischmannia schickendantzii Urtica chamaedryoides, Cantinoa mutabilis, Galium hypocarpium, Muhlenbergia schreberi and Seemannia gymnostoma, among others. This community is not very diverse; its total species richness is 77 species (Table 4.21), of which 20 are trees, 32 shrubs and 25 herbaceous. It has four characteristic or indicator species, of which three belong to the tree layer and behave as dominant (Cinnamomum porphyrium, Blepharocalyx salicifolius and Myrcianthes pseudomato), and one to the herbaceous layer (Justicia mandonii); it has no differential species. This community is very little represented in the study area, probably because its distribution is limited preferentially to steeper east-, northeast- and south-facing hillsides. It has floristic similarities with the mesowoodland of Cinnamomo porphyrium-Blepharocalycetum salicifolii (Association 11), with which it shares some characteristic species, as can be seen mainly in the ordination diagram in the Canonical Correspondence Analysis (Fig. 4.9: Sector III); however the dominance of Blepharocalyx salicifolius and Myrcianthes pseudomato—which occur with high abundance-cover values—and the absence of many of the characteristic species of that mesowoodland (Aralia soratensis, Bougainvillea stipitata, Cedrela saltensis, Duranta serratifolia, Stillingia tenella, Elephantopus mollis, Onoseris alata, Phytolacca bogotensis, Piper hieronymi and Solanum betaceum) determine that this is a different community to the woodland of Cinnamomo porphyriumBlepharocalycetum salicifolii. Particularly Myrcianthes pseudomato is a preferential species which, based on its high phytosociological values, behaves as dominant in the community, and participates in its physiognomy. As a result, in spite of having many common species and being dominated in the canopy by the same species, the two communities are different due to their topographic location, as evidenced by the abundance and dominance of these two mirtaceae on shadier hillsides. In view of this, this community represents the humid mirtaceae woodlands characteristic of the upper belts of the mountain forest (Cabrera 1994). Woodlands of this type have been described in the Sierra de San Javier (Tucumán, Argentina), dominated by Blepharocalyx salicifolius, Myrcianthes pseudomato and Myrcianthes

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mato; however, they are at a higher altitude and replace the woodlands dominated by Cinnamomum porphyrium and Blepharocalyx salicifolius (Grau et al. 2010). No mirtaceae woodlands have been described in the vegetation of the Chijra (Martín 2014) and Caulario river basins (Haagen Entrocassi 2014). 11. Cinnamomo porphyrium-Blepharocalycetum salicifolii ass. nova hoc loco (Table 4.18, holotypus rel. 6; Cluster 5B–5D; Appendix C). It is distributed in the central part of the study area, within the upper Mesotropical-lower Humid belt (1320–1433 m asl; It ¼ 374–387; Io ¼ 7.9–8.2) (Appendix A: Cluster 5B–5D; Fig. 4.9). It occupies gentle to moderately sloping hillsides with deep soils in different exposures (Figs. 5.1 and 5.2). At higher altitudes it substitutes the mesowoodlands of Tecomo stantis-Anadenantheretum cebilis (Association 9) and Zanthoxylo cocoi-Blepharocalycetum salicifolii (Association 8) on hillsides located more towards the north. This community coexists altitudinally with the mesowoodland of Myrciantho pseudomatoi-Blepharocalycetum salicifolii (Association 10), which preferentially occupies more steeply sloping and shadier hillsides. From the phytogeographical point of view it belongs to the mountain forest district (“high-mountain forest”) (Cabrera 1994). This is a semi-open and seasonally evergreen mesowoodland that grows to a height of approximately 20 m. Four well-defined structural layers can be differentiated. The upper tree layer (canopy) is dominated by tall specimens of Cinnamomum porphyrium (laurel del cerro) and Blepharocalyx salicifolius (horco molle), accompanied mainly by Parapiptadenia excelsa (horco cebil), Anadenanthera colubrina var. cebil (cebil colorado), Tipuana tipu (tipa blanca) and Cedrela angustifolia (cedro coya); there is a lower frequency of Juglans australis (nogal criollo) and Enterolobium contortisiliquum (pacará), and an isolated presence of Erythrina falcata (ceibo jujeño) and Cedrela saltensis. The arboreal understorey is disperse and grows to a height of approximately 8 m. It has a high species richness, although it is not dominated by any particular species. It is characterised mainly by the presence of Allophylus edulis (chal-chal), Myrcianthes pseudomato (mato arrayán), Kaunia lasiophthalma (malvón) and Sebastiania brasiliensis (leche-leche), and with a lower frequency by Solanum riparium (fumo bravo), Tecoma stans (guaranguay), Vassobia breviflora (pucancho), Myrcianthes pungens, Aralia soratensis (sacha paraíso), Bougainvillea stipitata (alfilerillo), Stillingia tenella, Sapium haematospermum (lecherón), Celtis iguanaea (tala pispa), Schinus gracilipes and Duranta serratifolia; also appearing in an isolated manner are Prunus tucumanensis (palo luz), Alnus acuminata (aliso del cerro), Pisonia zapallo (zapallo caspi), Myrsine laetevirens (palo San Antonio), Trema micrantha, Acacia aroma (tusca), Coutarea hexandra (dominguillo), Ilex argentina (palo yerba), Chrysophyllum marginatum (aguaí) and Citrus sp. The shrubby understorey is richer in species but has a diverse cover growing to a height of approximately 2 m; it is not dominated by any particular species, and as in the previous community, it has a high frequency of tall herbs such as Verbesina macrophylla var. nelidae and Vernonanthura pinguis, along with other species like Acalypha plicata, Boehmeria caudata, Baccharis latifolia, Justicia mandonii,

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Phenax laevigatus, Vernonanthura squamulosa, Cestrum parqui, Justicia kuntzei and Weddelia saltensis; there is a less frequent presence of Verbesina suncho, Dendrophorbium bomanii, Rubus imperialis, Urera baccifera, Aphelandra hieronymi, Solanum lorentzii, Heimia montana, Croton saltensis, Clinopodium bolivianum, Dolichandra ungis-cati, Chromolaena laevigata, Koanophyllon solidaginoides, Piper hieronymi, Abutilon grandifolium, Ludwigia peruviana, Cnidoscolus tubulosus, Solanum betaceum, Solanum confusum, Heteropterys sylvatica, Pavonia sepium, among many others; and an isolated presence of Ophryosporus lorentzii, Tournefortia paniculata, Thalictrum venturii, Aldama mollis, Buddleja stachyoides, Chiropetalum boliviense and Cnidoscolus vitifolius, among others. The herbaceous understorey is open to semi-open, although it forms a denser cover than the shrub layer and has a high species richness; it is dominated by Elephantopus mollis, accompanied mainly by Acalypha communis, Pharus lappulaceus, Petiveria alliacea, Rivinia humilis, Phytolacca bogotensis, Justicia goudotii, Tibouchina paratropica, Desmodium subsericeum, Adenostemma brasilianum, Salvia personata and Cuphea racemosa; there is a less frequent presence of Dicliptera squarrosa, Solanum aloysiifolium, Duchesnea indica, Onoseris alata, Anredera cordifolia, Urtica chamaedryoides, Pseudechinolaena polystachya, Panicum trichanthum, Conyza sumatrensis, Primula malacoides, Desmodium affine, Chaptalia nutans, Galinsoga caracasana, Muhlenbergia schreberi, Ruellia ciliatiflora, Tragia volubilis, Digitaria insularis, Solanum tenuispinum, Samolus valerandi, Plantago australis and Mikania micrantha, among many others. This community has a high diversity that is represented mainly in its shrub and herbaceous layer; its total species richness is 165 (Table 4.21), of which 35 are trees, 58 shrubs and 72 belong to the herbaceous layer. It also has the highest number of characteristic or indicator (22) species, of which ten belong to the tree layer: two behave as dominant (Cinnamomum porphyrium and Blepharocalyx salicifolius) and eight as selective or preferential (Cedrela angustifolia, Cedrela saltensis, Kaunia lasiophthalma, Myrcianthes pungens, Aralia soratensis, Bougainvillea stipitata, Stillingia tenella and Duranta serratifolia); the remaining species come from the shrub (Acalypha plicata, Baccharis latifolia, Justicia mandonii, Phenax laevigatus, Aphelandra hieronymi, Piper hieronymi, Solanum betaceum and Elephantopus mollis) and herbaceous layer (Acalypha communis, Phytolacca bogotensis, Onoseris alata and Petunia occidentalis); this last species behaves as differential. This is the community with the largest area of distribution within the upper Mesotropical belt. It is a mature mesowoodland with tall long-lived tall trees and a shady humid understorey with abundant epiphyte flora mainly comprising species of bromeliaceae, orchidiaceae, cactaceae, piperaceae, lichens and ferns. There is also an abundance of creepers and climbers; the soil is covered by a dense layer of plant detritus, and there are numerous species of pteridophytes that form extensive colonies. This community shares greater floristic and phytosociological similarities with the woodlands of Tecomo stantis-Anadenantheretum cebilis (Association 9) and Myrciantho pseudomatoi-Blepharocalycetum salicifolii (Association 10), as can be seen in the Hierarchical Classification Analysis (Fig. 4.6), whereas the Canonical Correspondence Analysis (Fig. 4.9: Sector III) shows greater affinity with the second

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woodland, and both respond positively to high values of Io. In the ordination diagram the community is represented by a cluster of sciophilous species located very close to the direction of the maximum variation of the Io vector, indicating primarily that its distribution responds to the more humid conditions (Sector III; relevés 43–45, 47–51). However, a small cluster of species can be distinguished (transition between sectors III and IV) that is also associated to high Io but at greater altitudes than the previous cluster; this cluster is close to the direction of maximum variation of the altitude vector and reflects the upper distribution limit of this community in the mountain forest (relevés 115, 117, 118 and 120). It shows a significant increase in the values of Cedrela angustifolia and the presence of Juglans australis, Alnus acuminata and Prunus tucumanensis, indicating the start of the ecotone with the mesowoodland of Pruno tucumanensis-Podocarpetum parlatorei (variant with Cedrela angustifolia; Association 12), located at a higher elevation and already belonging to the mountain woodland. Of the three mountain forest communities in the upper Mesotropical belt, this is the most humid, concurring with the reports in the vegetation studies in the Sierra de San Javier (Tucumán, Argentina) which distinguish one “dry” and another “humid” mountain forest, this last also dominated by Cinnamomum porphyrium and Blepharocalyx salicifolius (Grau et al. 2010). These two species also appear with a high frequency in several communities (52% and 76% respectively) in the vegetation in the Chijra river basin, growing under the upper Subhumid ombrotype; the first as a companion species and the second as dominant in one community (Martín 2014). In the less humid Caulario river basin (lower Subhumid), Cinnamomum porphyrium has a lower frequency (44.4%) and Blepharocalyx salicifolius is absent (Haagen Entrocassi 2014). 12. Pruno tucumanensis-Podocarpetum parlatorei Navarro and Maldonado 2002 subass. cedreletosum angustifoliae subass. nova hoc loco (Table 4.19, holotypus rel. 4; Cluster 4; Appendix C). It is distributed in the central part of the study area, within the Mesotropical-lower Humid belt (1488–1620 m asl; It ¼ 352–367; Io ¼ 8.3–8.7) (Appendix A: Cluster 4; Fig. 4.9). It occupies the edges and borders of the Serranías de Zapla in different exposures. Due to its location in the highest levels of the Reserve, this community only contacts with the mesowoodland of Cinnamomo porphyriumBlepharocalycetum salicifolii (Association 11) located immediately below, and with which it forms a broad ecotone (Figs. 5.1 and 5.2). From the phytogeographical point of view belongs to the mountain woodland district (Cabrera 1994). This is a seasonally evergreen mesowoodland which forms a semi-open woodland in the lower layer and is dense and closed at higher altitudes. It grows to a height of approximately 10 m, and four clearly defined strata can be differentiated in its structure. In its lower section (between 1488–1545 m asl), the presence of species that have their optimum in other communities an ecotone variant to be differentiated (with Cedrela angustifolia; relevés 96–99). The type community grows between 1577–1620 m asl (relevés 101–104). It is a closed woodland dominated solely by Podocarpus parlatoei (pino del cerro),

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accompanied by Cedrela angustifolia (cedro coya) and Juglans australis (nogal criollo), and in a very isolated manner by Blepharocalyx salicifolius (horco molle). The arboreal understorey grows to a height of 6–8 m and is dominated by Prunus tucumanensis (palo luz) and Alnus acuminata (aliso), accompanied by Sambucus nigra ssp. peruviana (sauco), Allophylus edulis (chal-chal), Schinus gracilipes and Cnicothamnus lorentzii (azafrán). Present in a disperse manner are Myrsine laetevirens (palo San Antonio), Stillingia tenella, Duranta serratifolia, Escallonia millegrana, Ilex argentina (palo yerba) and Zanthoxylum coco (cochucho). The only species confined to the type community is Berberis jobii, and there is also a presence of Rubus imperialis, Austroeupatorium inulifolium, Baccharis latifolia, Solanum aligerum, Clinopodium bolivianum, Campovassouria cruciata, Boehmeria caudata, Lepechinia vesiculosa, Muehlenbeckia sagittifolia, Ophryosporus lorentzii, Phenax laevigatus, Solanum confusum, Pavonia sepium, Senecio hieronymi, Senecio rudbeckiifolius, Baccharis dracunculifolia and Barnadesia odorata, among others; all these species also appear in the variant of the community with a different frequency and phytosociological values. In the herbaceous layer the species that only appear in the type community are Calceolaria teucrioides, Muhlenbergia schreberi, Cortaderia hieronymi, Mimosa xanthocentra and Veronica arvensis, whereas the remainder are shared with the variant of the community. The species found with greatest frequency include Duchesnea indica, Tibouchina paratropica, Stevia yaconensis var. subeglandulosa, Sibthorpia conspicua, Elephantopus mollis and Urtica chamaedryoides; present with a lower frequency are Axonopus compressus, Galium hypocarpium, Conyza sumatrensis, Pilea jujuyensis, Tradescantia boliviana, Cenchrus latifolius, Cajophora hibiscifolia, Desmodium affine, Deyeuxia polígama, Mikania micrantha, Phytolacca bogotensis, Festuca hieronymi, Anredera cordifolia and Begonia micranthera var. micranthera, among others. It also has an abundant pteridophyte cover. The variant of the ecotone is characterised by the presence of Cedrela angustifolia with high abundance-cover values, accompanied mainly by Cinnamomum porphyrium, Blepharocalyx salicifolius and Juglans australis, and, with a lower frequency, by Podocarpus parlatoei, Parapiptadenia excelsa and isolated individuals of Erythrina falcata (ceibo jujeño). Also frequent in the lower tree layer, although with low to moderate phytosociological values, are Prunus tucumanensis, Alnus acuminata, Sambucus nigra ssp. peruviana and Allophylus edulis, accompanied by Schinus gracilipes, Cnicothamnus lorentzii, Myrsine laetevirens, Stillingia tenella, Duranta serratifolia, Escallonia millegrana and Ilex argentina. The differential species of the variant in this stratum are: Anadenanthera colubrina var. cebil, Myrcianthes pseudomato, Kaunia lasiophthalma, Scutia buxifolia, Myrcianthes pungens, Bougainvillea stipitata and Vassobia breviflora. The differential species of the ecotone in the shrub layer are: Bidens squarrosa, Koanophyllon solidaginoides, Acalypha plicata, Heimia montana, Manetia jorgensenii, Solanum betaceum, Vernonanthura squamulosa, Ophryosporus piquerioides, Abutilon grandifolium, Aphelandra hieronymi, Cestrum parqui, Cnidoscolus tubulosus, Croton saltensis, Dendrophorbium bomanii, Justicia kuntzei, Justicia mandonii and Piper hieronymi. In the herbaceous layer the

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differential species are: Jungia pauciflora, Dicliptera squarrosa, Acalypha communis, Adenostemma brasilianum, Chaptalia nutans, Galinsoga caracasana, Solanum tenuispinum, Desmodium subsericeum, Digitaria insularis, Galium lilloi, Justicia goudotii, Onoseris alata and Tragia volubilis. This community has a high diversity; its total species richness is 111 species (Table 4.21), of which 26 are trees, 45 shrubs and 40 belong to the herbaceous layer. Of the 18 characteristic or indicator species, six belong to the tree layer: one behaves as dominant (Podocarpus parlatorei), three as selective (Prunus tucumanensis, Alnus acuminata and Ilex argentina), one as exclusive (Sambucus nigra ssp. peruviana) and one as preferential of the ecotone variant (Cedrela angustifolia). Of the remaining species, eight belong to the shrub layer (Austroeupatorium inulifolium, Solanum aligerum, Clinopodium bolivianum, Campovassouria cruciata, Lepechinia vesiculosa, Ophryosporus lorentzii, Solanum confusum and Berberis jobii); this last behaves as a differential species. Four belong to the herbaceous layer (Tibouchina paratropica, Stevia yaconensis var. subeglandulosa, Sibthorpia conspicua and Calceolaria teucrioides); this last is also differential. The characteristic or indicator species that represent the type community form a clearly defined cluster that is distributed in the direction of maximum variation of the altitude vector (Fig. 4.9: Sector IV); these include Podocarpus parlatorei (Podo), Prunus tucumanensis (Prun), Alnus acuminata (Alnu), Ilex argentina (Ilex), Sambucus nigra ssp. peruviana (Samb), Austroeupatorium inulifolium (Aust), Solanum aligerum (Sali), Campovassouria cruciata (Camp), Lepechinia vesiculosa (Lepe), Ophryosporus lorentzii (Oplo), Berberis jobii (Berb), Sibthorpia conspicua (Sibc) and Calceolaria teucrioides (Calt). The ecotone variant is indicated in the ordination diagram in the Canonical Correspondence Analysis by the segregation of one cluster of species that also responds positively to the increase in altitude (Fig. 4.9: Sector IV). Many of them are differentiating species for the variant such as Cedrela angustifolia (Cean), Myrsine laetevirens (Myrs), Bidens squarrosa (Bisq), Koanophyllon solidaginoides (Koan), Heimia montana (Heim), Solanum betaceum (Sbet), Ophryosporus piquerioides (Oppi), Abutilon grandifolium (Abut) and Galium lilloi (Gali); and other species are shared with the type community such as Rubus imperialis (Rubu), Clinopodium bolivianum (Clino), Boehmeria caudata (Bohe), Muehlenbeckia sagittifolia (Mueh), Solanum confusum (Scon), Tibouchina paratropica (Tibo), Stevia yaconensis var. subeglandulosa (Stey), Axonopus compressus (Axon), Pilea jujuyensis (Pile), Cenchrus latifolius (Cenc), Deyeuxia polígama (Deye), Festuca hieronymi (Fehi) and Begonia micranthera var. micranthera (Begm). According to Cabrera (1994), the woodlands of Podocarpus parlatorei (Gymnospermae, Podocarpaceae) or “pino del cerro” as they are known, constitute one of the characteristic vegetation types of the mountain woodland and are frequently associated with Juglans australis (nogal criollo) and Alnus acumunata (aliso). Until 1962 these woodlands occupied 110,000 hectares in the province of Jujuy; however, due to their value as timber they have been intensely exploited and there are therefore very few sites on which mature woodlands can be found today. According to Grau et al. (2010), in the early-intermediate successional stages, the

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characteristic tree species in these woodlands in addition to Podocarpus parlatorei are Alnus acuminata and Crinodendron tucumanum, whereas in the late successional stages and in mature woodlands there is an incorporation of Ilex argentina, Prunus tucumanensis, Cedrela lilloi, Juglans australis and diverse species from the Myrtaceae family, all present in the community analysed here (except Crinodendron tucumanum which was not relevéd). This mesowoodland therefore probably represents the late or mature stage of the forest community. Due to its location and difficult access it is relatively well conserved; the main disturbance it currently suffers is the temporary transit of livestock. However, the presence of traces of manufacturing indicates that selective extraction of some of its valuable forestry species has taken place in the past. The general floristic composition of this community concurs with the flora recorded in other mountain woodlands in northwest Argentina. In the locality of Los Toldos (Salta, Argentina) these woodlands are dominated by Podocarpus parlatorei, together with Juglans australis (Pinazo et al. 2003). Three Podocarpus parlatorei communities have been described in the Sierra de Medina (Tucumán, Argentina), with varying species richness and structure. In one community it is dominant with Alnus acuminata (at 1493 m asl; 115 species), in another with Juglans australis (at 1500 m asl; 110 species) and in the third with Allophylus edulis (at 1540 m asl; 94 species) (Perea et al. 2008). Podocarpus parlatorei woodlands are located at a higher altitude (1800 m asl) in the Chijra river basin, at the limit of the upper Mesotropical belt under an upper Subhumid ombrotype. The floristic composition of this woodland is very similar to the one seen here, with a tree layer that includes Blepharocalyx salicifolius, Cinnamomum porphyrium, Erythrina falcata, Duranta serratifolia, Allophyllus edulis, Kaunia lasiophthalma, Sambucus nigra ssp. peruviana, Vassobia breviflora, Sebastiania brasiliensis, Sebastiania commersoniana and Schinus myrtifolius. The three last species are absent from the community in the Reserve. The characteristic species in the shrub and herbaceous layer include Boehmeria caudata, Tibouchina paratropica, Pavonia sepium, Solanum confusum, Phenax laevigatus, Elephantopus mollis and Desmodium affine. These species can also be found in the community analysed here, in addition to others such as Oplismenus hirtellus, Solanum microdontum, Bidens mandonii, Jungia pauciflora, Dicliptera jujuyensis, Seemannia nematanthodes, Iresine diffusa, Sida cabreriana, Sida rhombifolia and Begonia micranthera, among others (Martín 2014). Finally, the study by Cuyckens (2005) on a plot located at 1600 m asl in the Serranías de Zapla makes no reference to the existence of woodlands dominated by Podocarpus parlatorei, and includes these species as part of a community dominated by Allophylus edulis, which is the most common species in the sampled plot. These differences may have a methodological basis, as the author does not apply a phytosociological approach in her study and only considers the greater or lesser species abundance. It is probable however that due to its altitudinal location this community corresponds to a Podocarpus parlatorei woodland like the one described in the present study.

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5.10

Riparian Vegetation

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Final Considerations on Terrestrial Plant Communities in the Lower and Upper Mesotropical Bioclimatic Belt

The above diagnosis describes the main characteristics that define the identity of these communities, and highlights the floristic and phytosociological similarities between some of them in response to similar environmental conditions, or for dynamic successional reasons. The ordination diagram in the Canonical Correspondence Analysis (Fig. 4.9) shows the species distribution in the communities based on the gradients of altitude, It and Io in the study area; this chart shows that the clouds that conform the communities are very close together, overlapping or separate according to the species’ response to these variables and whether or not they have certain floristic and phytosociological affinities with each other. As a result, the plant communities that grow in the lower Mesotropical belt (basal mountain forest) are clearly separated from those that occupy the upper Mesotropical belt (high-mountain forest and mountain woodland). In turn, the first communities include those that occupy the lower, warmer areas with less humidity in the southern and western area of the Reserve (Associations 1, 2 and 3; Sector I: 1032–1127 m asl; It ¼ 417–429; Io ¼ 4.7–4.8) and those that are distributed successively at greater heights in the western (Associations 4–8; Sector II: 1192–1275 m asl; It ¼ 399–409; Io ¼ 4.9–5.5) and central zone (Association 8: 1115 m asl; It ¼ 412; Io ¼ 7.4) under conditions of lower temperature and greater humidity. The set of communities in the upper Mesotropical belt include those that belong to the upper belt of the mountain forest (“high-mountain forest”) and which are distributed at higher elevations along the altitudinal gradient, and in more temperate and humid conditions (Associations 9, 10 and 11; Sector III and transition to IV: 1260–1433 m asl; It ¼ 374–395; Io ¼ 7.7–8.2); while the community belonging to the mountain woodland is clearly segregated and located at the upper end of the altitudinal gradient, and is therefore the most humid and least warm in the study area (Association 12; Sector IV: 1488–1620 m asl; It ¼ 352–367; Io ¼ 8.3–8.7). Some of these communities have a generally good state of conservation and constitute mature mesowoodlands that have been relatively unimpacted or are clearly in the process of recovery, with long-lived tall trees, many with hard and valuable wood. Other communities reveal past interventions including intensive cutting, and form secondary microwoodlands without any valuable forestry species.

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Riparian Vegetation

13. Salici humboldtianae-Acacietum aromae ass. nova hoc loco (Table 4.20, holotypus rel. 1; Cluster 6B; Appendix C).

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It is distributed in the centre-south and southern part of the study area, within the lower Mesotropical-lower Subhumid belt, extending towards the lower Humid belt (1015–1030 m asl; It ¼ 422–429; Io ¼ 4.7–7.2) (Appendix A: Cluster 6B; Fig. 4.9). It occupies low-lying floodable terraces and the banks and beaches of the Zapla river and the Pacará stream (Figs. 5.1 and 5.2). From the phytogeographical point of view it belongs to the mountain forest district (Cabrera 1994). It constitutes an open to semi-open deciduous microwoodland growing to a height of approximately 4 m, where the type community (relevé 52) and a variant with Tecoma stans (relevé 105) can be distinguished. The first grows on the banks and beaches of the Zapla river, in an unstable environment subject to severe flooding and the inflow of alluvial sediment (stones and mud) in summer. This woodland is very poor floristically, which may be due to the fact that the representative relevé for this woodland has a smaller area (500 m2) and to the geomorphological and soil characteristics of its location. Structurally, three layers can be differentiated: the tree layer is characterised by Acacia aroma (tusca), Salix humboldtiana (sauce criollo), Tessaria integrifolia (pájaro bobo), Sebastiania brasiliensis (leche-leche), and juvenile specimens of Parapiptadenia excelsa (horco cebil). The companion species present include Celtis iguanaea (tala pispa), Allophylus edulis (chal-chal), Trema micrantha, Vassobia breviflora (pucancho) and Acacia caven (espinillo). The shrubby understorey contains Urera baccifera (ortiguilla), Baccharis salicifolius and Tessaria dodoneifolia (chilca), and in the herbaceous understorey includes Dicliptera squarrosa, Polygonum punctatum, Paspalum distichum, Alternanthera philoxeroides, Asclepias curassavica, Galinsoga caracasana and Nicandra physalodes. Even when the species richness of these layers is very low, most of the species present are indicators of riparian environments and their presence is therefore significant. The variant with Tecoma stans (guaranguay) grows at a higher altitude under a humid ombrotype and is distributed primarily on relatively more stable river terraces, which explains its greater species richness. Its tree layer also contains the typical species of the community (except Trema micrantha, Vassobia breviflora and Acacia caven) and some emergent species of Anadenanthera colubrina var. cebil (cebil colorado) and Enterolobium contortisiliquum (pacará), which appear in an isolated manner. In the shrub layer there is a predominance of Urera baccifera, accompanied by Baccharis salicifolius, Tessaria dodoneifolia, Clematis haenkeana, Acalypha amblyodonta, Carica glandulosa, Sida rhombifolia and Smilax campestris, whereas the herbaceous layer contains mainly Jungia pauciflora, Dicliptera squarrosa, Mimosa xanthocentra, Polygonum punctatum, Parthenium hysteriophorus, Rivinia humilis, Hydrocotyle bonariensis, Paspalum distichum, Tradescantia boliviana, Alternanthera philoxeroides and Asclepias curassavica, among others. Many of these species have not been relevéd in the type community. This variant contains species from the type community and therefore indicators of the riparian environment, but also has some species from nearby foothills and hillsides that descend towards the river terraces and become established. The presence of species from the adjoining woodlands is fairly common in all these subtropical riparian communities in northwest Argentina (Sirombra and Mesa 2010).

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Riparian Vegetation

155

The total species richness in this community is 45 species (Table 4.21), of which 13 are trees, eight shrubs and 24 belong to the herbaceous layer. Of the 11 characteristic or indicator species in this community, four belong to the tree layer: one behaves as dominant (Acacia aroma), two are selective (Salix humboldtiana and Tessaria integrifolia) and one behaves as preferential of the community variant (Tecoma stans). Of the remaining characteristic species, three are from the shrub layer: one behaves as dominant of the variant (Urera baccifera) and two as selective (Baccharis salicifolius and Tessaria dodoneifolia), whereas four species in the herbaceous layer also behave as selective (Polygonum punctatum, Paspalum distichum, Alternanthera philoxeroides and Asclepias curassavica). This community does not have any differential species. The classic work by Cabrera (1994) does not describe any woodlands of SalixHumboldtiana and Acacia aroma for the mountain forest. However it does mention the small riparian woodland of “guaranguay” (Tecoma stans) in the foothill woodland, which probably corresponds to the variant of the type community analysed here, given that its floristic composition, in addition to Tecoma stans, also reveals a predominance of Salix humboldtiana, Tessaria integrifolia, Baccharis salicifolius and Tessaria dodoneifolia. In the riparian vegetation of the Caulario river basin, microwoodlands of Tecoma stans-Acacia aroma were described in the lower Mesotropical-lower Subhumid belt (between 958–998 m asl), with a similar composition to the variant identified in the Reserve, except for the absence of the “sauce criollo” (Salix humboldtiana) (Haagen Entrocassi 2014). In the vegetation of the middle basin of the Chijra river, only Acacia aroma forms part of the riparian shrub formations of Acacia caven-Sapium haematospermum (between 1424–1455 m asl), whereas Salix humboldtiana and Tecoma stans are absent from the riparian environment. This microwoodland constitutes the edaphohygrophilous vegetation associated to the Zapla river basin, of which the Pacará stream is also a part, and thus represents the azonal vegetation in the Reserve. This can be seen in the ordination diagram in the Canonical Correspondence Analysis (Fig. 4.9), where the species that characterise this woodland do not respond significantly to the environmental variables analysed—basically the altitude and Io gradient—, which is to be expected as this is a community that depends directly or indirectly on the water accumulated in the soil, and is not substantially affected by the changes in elevation and humidity that occur within this altitudinal interval. Its location in the ordination diagram places it close to the lowest community in the upper Mesotropical belt (Tecomo stantisAnadenantheretum cebilis; Association 9). Both communities share the same diagnostic species (Tecoma stans), in addition to other species that, as indicated, come from the surrounding woodlands and extend into the riparian environment; the mesowoodland of Tecomo stantis-Anadenantheretum cebilis is relatively near this riparian community, as it descends as edaphohygrophilous through humid gorges to the lower Mesotropical belt. Similarly, in the dendrogram of the Hierarchical Classification Analysis (Fig. 4.2), this community is clustered with the mountain forest communities in the upper Mesotropical belt, although it belongs to a lower bioclimatic belt.

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5 Biodiversity Analysis: A Geobotanic Interpretation

Finally, it should be clarified that from a floristic, phytosociological, geomorphological and soil characteristics point of view this woodland are clearly recognized as community in spite of the number of representative relevés presented in this work. However, it is worth noting the need for future relevé collection sampling to expand the information on these riparian communities.

Chapter 6

Final Remarks

The floristic composition of the vegetation is represented by 257 species belonging to 194 genera and 66 botanical families. 65 species were recorded in the tree layer, representing 32% of the arboreal flora of Las Yungas, while 85 species were identified in the shrub layer and 107 species in the herbaceous layer, which was the layer showing the highest richness in terms of species and families. The study area contains a representation of 39% of the families, 20% of the genera and 9.1% of the species in the province of Jujuy. Of the total species recorded, 216 are native, 25 endemic, 13 exotic and three cosmopolitan. The most represented families were Asteraceae, Fabaceae, Poaceae, Solanaceae and Euphorbiaceae. The family with the greatest number of species in the tree layer was Fabaceae, with Asteraceae in the shrub and herbaceous layers. There are a few species that occur with a high frequency (between 60 and 100%) and many species with a low frequency (less than 20%). Although the abundance (number of individuals) was not determined for each of the species recorded, it was generally observed in the field that the species with high frequency tend to be the most abundant. The floristic composition, abundance-dominance, frequency and species richness vary along the environmental gradient, indicating that the species are most abundant and frequent within an optimum interval of ecological conditions along this gradient, and that outside this optimum they decrease or disappear. One cluster of species is distributed throughout the whole gradient, generally reflecting a wide range of ecological tolerance that enables them to occupy various environments. Another cluster of species appears exclusively within a particular interval of this gradient. 41 species are found only in the lower Mesotropical belt and 35 in the upper Mesotropical belt, revealing a more restricted distribution interval. Similarly, within the upper Mesotropical belt, one cluster of species is only found in the upper belt of the mountain forest (“high-mountain forest”), whereas another is practically exclusive to the mountain woodland. Two species richness peaks occur in mountain forest communities: one in the lower Mesotropical belt (with 183 species) and another in the upper Mesotropical © Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3_6

157

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6 Final Remarks

belt (with 165 species), coinciding with other studies that indicate the mountain forest as the most diverse vegetation layer within the subtropical mountain woodlands. The greatest exchange of species is found in the tree layer and occurs in the transition between the lower Mesotropical and upper Mesotropical bioclimatic belt (at 1260 m asl), and in the ecotone between the mountain forest and the mountain woodland (above 1488 m asl approximately) in the upper Mesotropical belt, indicating a significant change in environmental conditions linked to the decrease in temperature and increase in humidity, which in turn determines changes in the floristic composition of the dominant tree layer. Phanerophytes constitute the most abundant biotype (57%), in consonance with the type of woodland formation growing in the study area, followed by hemicryptophytes (26%), therophytes (8%), geophytes (7%) and chamaephytes (2%). The vegetation in the study area is distributed in an altitudinal belt of 600 m (between 1015 and 1620 m asl) within the lower and upper Mesotropical bioclimatic belt, with a lower Subhumid, upper and lower Humid pluviseasonal tropical bioclimate. Thirteen plant communities were identified and characterised, corresponding to the subtropical mountain woodlands or Yungas characteristic of the mountain forest and mountain woodland districts in the phytogeographical province of Las Yungas (Amazonian Domain, Neotropical Region). These woodlands can in turn be attributed to the Bolivian-Tucuman sub-Andean pluviseasonal mountain vegetation in the Bolivian-Tucuman biogeographical province (Tropical South-Andean Region, Neotropical Sub-Kingdom, Neotropical-Austroamerican Kingdom; Rivas-Martínez et al. 2011). To recognise and delimit the plant communities, 103 diagnostic (characteristic or indicator) species were selected, of which 29 behave as dominant, 67 are distributed between exclusive, selective, preferential and differential, and seven are stenoic. The tree layer has the highest percentage of diagnostic species (46.6%), which would explain the diversity of woodland species identified. The communities identified form seasonal, mainly semi-deciduous and evergreen micro- and mesowoodlands growing on foothills, hillsides, ravines, gorges and the edges of mountain ranges (terrestrial communities), and on river terraces and beaches (riparian communities). The former have a greater species richness, probably because they are located in more stable areas than riparian environments, which tend to be more fluctuating and heterogeneous. The species richness of the plant communities underwent variations along the altitudinal gradient, but did not follow the general pattern observed for altitudinal gradients in the Neotropical, which indicates a decrease in species richness in relation to the increase in altitude. This could be due to the fact that the altitudinal interval in the study is not sufficiently pronounced to reflect a decrease in species richness in relation to the increase in altitude; other factors may also have influenced the richness of the communities, such as the number and area of the representative relevés, their environmental location and state of conservation. The composition and distribution of the plant communities in the study area are mainly determined by the altitudinal gradient prevailing in the area, and by the gradients of the thermicity (It) and ombrothermic (Io) indexes as a result of the variations in temperature and precipitation that occur due to the altitude. However, the influence of the altitudinal

6 Final Remarks

159

gradient is associated in some situations with the effect of other geophysical factors such as the topographic orientation of the slopes, the geomorphology and the soils, which exert their effects in sites with specific environmental characteristics (e.g. ravines, gorges, river terraces and beaches). The plant communities growing in the lower Mesotropical bioclimatic belt (1015–1275 m asl; It ¼ 399–429) represent the micro- and mesowoodlands that occupy the low-lying, warm and subhumid areas in the southern and western zones of the study area, under lower and upper Subhumid ombrotypes (Io ¼ 4.7–5.5). However, one of these communities reappears in the central zone with a lower Humid ombrotype. These communities belong to the lower altitudinal belt of the mountain forest (“basal forest”) and are characterised by more thermophilous species. The plant communities that are distributed in the upper Mesotropical bioclimatic belt (1260–1620 m asl; It ¼ 352–395) represent the mesowoodlands growing at higher altitudes under more temperate and humid conditions in the central zone of the study area, under a lower Humid ombrotype (Io ¼ 7.7–8.7). These communities belong to the upper belt of the mountain forest (“high-mountain forest”) and to the mountain woodland, and are generally characterised by sciophilous and less thermophilous species. The communities that belong to the mountain forest (“basal forest” and “high-mountain forest”) have a greater percentage of diagnostic species (82.5%) than the mountain woodland community (17.5%). These differences are correlated with the greater species richness found in the mountain forest. The state of conservation of the communities is linked to their location in the territory; those distributed in the lower Mesotropical belt show a higher degree of anthropic intervention and generally constitute more open and luminous woodlands containing a larger number of heliophilous, exotic and cosmopolitan species. Some of these communities form secondary microwoodlands with only three layers, with young trees with very little or no value to forestry. The communities located in the upper Mesotropical belt are in a better state of conservation due to their location at higher altitudes and in areas that are difficult to access, which has limited or completely prevented their exploitation in certain sites. They form mature mesowoodlands with very little intervention or in evident recovery, with tall, longlived trees with very hard and valuable wood. The communities identified and described in the present study were typified from the phytosociological point of view as the following associations: Enterolobio contortisilici-Anadenantheretum cebilis; Schino bumeloidis-Allophyletum edulis; Xylosmo pubescentis-Blepharocalycetum salicifolii; Jacarando mimosifoliaeVassobietum breviflorae; Erythrino falcatae-Tipuanetum tipi; Schinetum myrtifoliogracilipedis; Juglandi australis-Blepharocalycetum salicifolii; Zanthoxylo cocoi-Blepharocalycetum salicifolii; Tecomo stantis- Anadenantheretum cebilis; Myrciantho pseudomatoi-Blepharocalycetum salicifolii; Cinnamomo porphyriumBlepharocalycetum salicifolii; Pruno tucumanensis-Podocarpetum parlatorei; Salici humboldtianae-Acacietum aromae. These are new associations that are described for the first time for the province of Jujuy and have a provisional character, except that of Pruno tucumanensis-Podocarpetum parlatorei which has been described for

160

6 Final Remarks

Bolivia, and represents a geographic variant of the same (subass. cedreletosum angustifoliae nova). Serranías de Zapla Multiple Use Ecology Reserve has a high environmental and biological diversity. It is in the sector with the highest biodiversity in Las Yungas and forms part of a biological corridor that connects several national and foreign protected areas. An area of 7730 hectares (21%) located at the northern extreme of the zone are part of the Las Yungas Biosphere Reserve (RBYungas). It protects the basin of the Zapla and Las Capillas rivers, both of which are tributaries of the San Francisco river. They have permanent water regimes and belong to the great hydrographic network of the upper Bermejo river basin, which in turn is part of the Guaraní aquifer, one of the largest underground water reserves on the planet. It is included within Category II (Yellow) of the Territorial Ordination Plan for the Native Woodlands in the province of Jujuy, as an area of woodland maintenance subject to conservation and/or sustainable management plans. It is also within the territory declared as the “Jujuy Model Forest”, framed within the “National Model Forest Programme”. It has a high conservation potential due to the presence of plant species with biological value, due either to their scarcity or to their vulnerable status in Argentina, or because they are a source of food and refuge for many resident and migratory animal species. The existence of secondary forests containing trees of value for forestry is evidence of the current state of recovery of some sectors of woodland that had previously been overexploited. The present research work is part of a series of studies that began in the province of Jujuy in 2012, in which for the first time the phytosociological methodology of Braun-Blanquet is applied to delimit and describe the plant communities in the subtropical mountain woodlands, and to analyse their relationship with the bioclimate. The bioclimatic maps for the province of Jujuy are used as a support tool, and adequately represent the correspondence between the climate and the vegetation plant distribution in this province. The results obtained in this study contribute to a greater knowledge of the subtropical mountain woodlands or Yungas in northwest Argentina, and particularly in the province of Jujuy. They reveal for the first time a series of woodland communities whose composition and distribution expand the current ecological information on these woodlands, while at the same time establishing the first precedent for their phytosociology and syntaxonomy by recording the communities identified as new associations with a provisional character. This research did not undertake the study of the vegetation belt located in the far northeast of the Reserve which, due to its location and for reasons of cost, was not sampled; new relevé campaigns in this area will probably reveal a greater influence of the foothill woodland and allow comparison with the vegetation in the southern

6 Final Remarks

161

zone of the Reserve, which as indicated earlier, reveals the encroachment of floristic elements from the foothills. Similarly, and for the same reasons, the work does not include the study of the riparian communities in the upper Mesotropical belt, which will be taken into account in future research. It would also be advisable to increase the number of relevés in the association Salici humboldtianae-Acacietum aromae in order to optimise its representation. The sampling must be repeated on sites close to those occupied by the relevés in Cluster 6A, which has yet to be typified, in order to determine whether it belongs to any of the communities described, as due to the floristic and phytosociological characteristics of its relevés it was not recognised as a community with its own identity. The information obtained in this research was sufficient to achieve the objectives and hypothesis proposed initially. It allowed us to identify and delimit the plant communities and determine their floristic composition, species richness, abundancedominance, the frequency of the species that form them, and the biological types that characterise this woodland formation; we were also able to establish their distribution and the bioclimatic characteristics of the areas in which they are found; and finally, we analysed the influence of the environmental gradient as a factor controlling the vegetation, and the main environmental factors (altitude, It and It) responsible for the variations observed in the floristic composition and distribution of the vegetation in the subtropical mountain woodlands of the Serranías de Zapla Multiple Use Ecology Reserve (Jujuy, Argentina). Finally, following Navarro and Maldonado (2002) we can propose the following syntaxonomical scheme: Podocarpo parlatorei-Tipuanetea tipi Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Tipuano tipi-Podocarpetalia parlatorei Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Juglandi australis-Phoebion porphyrae Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Enterolobio contortisilici-Anadenantheretum cebilis ass. nova Schino bumeloidis-Allophyletum edulis ass. nova Xylosmo pubescentis-Blepharocalycetum salicifolii ass. nova Jacarando mimosifoliae-Vassobietum breviflorae ass. nova Erythrino falcatae-Tipuanetum tipi ass. nova Schinetum myrtifolio-gracilipedis ass. nova Juglandi australis-Blepharocalycetum salicifolii ass. nova Zanthoxylo cocoi-Blepharocalycetum salicifolii ass. nova Myrciantho callicomae-Podocarpion paralatorei Rivas-Martínez et Navarro in Navarro and Maldonado 2002 Tecomo stantis-Anadenantheretum cebilis ass. nova Myrciantho pseudomatoi-Blepharocalycetum salicifolii ass. nova Cinnamomo porphyrium-Blepharocalycetum salicifolii ass. nova Pruno tucumanensis-Podocarpetum parlatorei Navarro and Maldonado 2002 subass. cedreletosum angustifoliae subass. nova

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6 Final Remarks

Salici humboldtiani-Prosopietea albae Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Salici humboldtiani-Prosopietalia albae Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Prosopion albae Rivas-Martínez et Navarro in Navarro et Maldonado 2002 Salici humboldtianae-Acacietum aromae ass. nova

Appendix A

Bioclimatic characterization of study area Rel. 52 105 53 54 56 55 57 60 59 58 61 62 65 63 64 66 67 68 69 70 71 72 80 73 74 75

Alt 1015 1030 1032 1032 1034 1035 1035 1035 1036 1037 1085 1086 1087 1090 1090 1091 1093 1095 1099 1102 1105 1107 1108 1109 1112 1115

T 17.5 17 17.4 17.4 17.4 17.4 17.4 17.4 17.39 17.39 17.27 17.27 17.26 17.25 17.25 17.24 17.23 17.22 17.2 17.19 17.17 17.16 17.16 17.15 17.14 17.12

P 992 1472 992 992 992 992 992 992 992 992 991 991 991 991 991 991 991 991 991 991 991 991 991 991 991 991

M 20.2 20.14 20.17 20.17 20.16 20.16 20.16 20.16 20.16 20.16 20.03 20.03 20.03 20.02 20.02 20.02 20.01 20.01 20 19.99 19.99 19.98 19.98 19.98 19.97 19.97

M 5.2 5.1 5.11 5.11 5.1 5.1 5.1 5.1 5.09 5.09 4.87 4.87 4.86 4.85 4.85 4.84 4.83 4.82 4.8 4.79 4.77 4.76 4.76 4.75 4.74 4.72

Tp 2100 2040 2088 2088 2088 2088 2088 2088 2086.8 2086.8 2072.4 2072.4 2071.2 2070 2070 2068.8 2067.6 2066.4 2064 2062.8 2060.4 2059.2 2059.2 2058 2056.8 2054.4

It 429 422 427 427 427 427 427 427 426 426 422 422 421 421 421 421 421 420 420 420 419 419 419 419 418 418

Io 4.7 7.2 4.7 4.7 4.7 4.7 4.7 4.7 4.7 4.7 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8 4.8

Th Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr

Om Low Shu Low Hum Low Shu Low Shu Low Shu Low Shu Low Shu Low Shu Low Shu Low Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu

Gr 6B 6B 1A

As 13 13 1

2B

2

2B

2

2C

3

(continued)

© Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3

163

164

Rel. 79 76 77 78 81 82 83 84 85 86 106 1 2 3 4 22 5 21 6 7 20 8 42 39 40 36 37 41 32 33 34 38 31 35 30 28 29 27

Appendix A

Alt 1116 1118 1121 1127 1192 1195 1195 1199 1198 1200 1115 1208 1215 1219 1228 1230 1235 1243 1247 1250 1255 1258 1235 1240 1240 1243 1243 1239 1240 1240 1241 1241 1242 1244 1245 1250 1252 1253

T 17.12 17.11 17.1 17.06 16.74 16.72 16.72 16.7 16.71 16.7 16.6 16.7 16.62 16.6 16.56 16.5 16.52 16.43 16.46 16.45 16.37 16.41 16.47 16.45 16.45 16.43 16.43 16.45 16.45 16.45 16.45 16.45 16.44 16.43 16.42 16.4 16.39 16.38

P 991 991 991 991 991 991 991 991 991 991 1472 991 991 991 991 1.079 991 1.079 991 991 1.079 991 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079 1.079

M 19.97 19.96 19.96 19.95 19.82 19.81 19.81 19.8 19.8 19.8 19.97 19.78 19.77 19.76 19.74 19.75 19.73 19.72 19.7 19.7 19.7 19.68 19.74 19.73 19.73 19.72 19.72 19.73 19.73 19.73 19.69 19.69 19.72 19.72 19.72 19.7 19.7 19.7

M 4.72 4.71 4.69 4.66 4.34 4.32 4.32 4.3 4.31 4.3 4.67 4.26 4.22 4.2 4.16 4.1 4.12 4.035 4.06 4.05 3.97 4.01 4.07 4.05 4.05 4.03 4.03 4.05 4.05 4.05 4.04 4.04 4.04 4.03 4.02 3.98 3.99 3.98

Tp 2054.4 2053.2 2052 2047.2 2008.8 2006.4 2006.4 2004 2005.2 2004 1992 2004 1994.4 1992 1987.2 1980 1982.4 1971.6 1975.2 1974 1964.4 1969.2 1976.4 1974 1974 1971.6 1971.6 1974 1974 1974 1974 1974 1972.8 1971.6 1970.4 1965.6 1966.8 1965.6

It 418 418 417 417 409 408 408 408 408 408 412 407 406 406 405 403 404 402 402 402 400 401 403 402 402 402 402 402 402 402 402 402 402 402 402 401 401 401

Io 4.8 4.8 4.8 4.8 4.9 4.9 4.9 4.9 4.9 4.9 7.4 4.9 5 5 5 5.4 5 5.4 5 5 5.5 5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5

Th Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr

Om Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Low Hum Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu

Gr

As

2A

4

1C

5

1C 3A

5 6

3B

7

(continued)

Appendix A

165

Rel. 23 24 25 26 19 18 9 16 17 15 10 14 13 12 11 107 108 109 110

Alt 1233 1235 1249 1250 1255 1257 1259 1260 1260 1263 1265 1265 1270 1273 1275 1180 1222 1249 1260

T 16.48 16.47 16.4 16.4 16.37 16.36 16.4 16.4 16.4 16.38 16.37 16.37 16.35 16.33 16.32 16.25 16.04 15.9 15.85

P 1.079 1.079 1.079 1.079 1.079 1.079 991 991 991 991 991 991 991 991 991 1472 1472 1472 1472

M 19.74 19.74 19.71 19.71 19.7 19.69 19.68 19.68 19.68 19.67 19.67 19.67 19.66 19.65 19.65 19.84 19.76 19.7 19.68

M 4.08 4.07 4 4 3.97 3.96 4 4 4 3.98 3.97 3.97 3.95 3.93 3.92 4.35 4.14 4 3.95

Tp 1977.6 1976.4 1968 1968 1964.4 1963.2 1968 1968 1968 1965.6 1964.4 1964.4 1962 1959.6 1958.4 1950 1924.8 1908 1902

It 403 402 401 401 400 400 401 401 401 400 400 400 400 399 399 404 399 396 395

Io 5.4 5.4 5.5 5.5 5.5 5.5 5 5 5 5 5 5 5 5 5.1 7.5 7.6 7.7 7.7

111 113 114 95 94 93 92 91 89 88 87 43 44 45 47 48 49 50 51 115 117 118 120

1278 1300 1310 1319 1322 1325 1331 1337 1342 1355 1360 1320 1335 1344 1351 1359 1360 1360 1365 1367 1393 1400 1433

15.76 15.65 15.6 15.55 15.54 15.5 15.49 15.46 15.4 15.37 15.3 15.5 15.5 15.4 15.4 15.3 15.3 15.3 15.3 15.3 15.2 15.15 14.98

1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472 1472

19.64 19.6 19.6 19.56 19.5 19.5 19.5 19.5 19.5 19.5 19.48 19.56 19.5 19.5 19.5 19.48 19.48 19.48 19.47 19.46 19.4 19.4 19.33

3.86 3.75 3.7 3.65 3.6 3.6 3.59 3.56 3.54 3.47 3.45 3.65 3.6 3.5 3.5 3.45 3.45 3.45 3.42 3.41 3.28 3.25 3.08

1891.2 1878 1872 1866 1864.8 1860 1858.8 1855 1848 1844 1836 1860 1860 1848 1848 1836 1836 1836 1836 1836 1824 1818 1797.6

392 390 389 388 386 386 385 385 384 383 382 387 386 384 384 382 382 382 382 381 379 378 374

7.8 7.8 7.9 7.9 7.9 7.9 7.9 7.9 8 8 8 7.9 7.9 8 8 8 8 8 8 8 8 8.1 8.2

Th Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr Low Mtr- Sup Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr

Om Gr Upp Shu 1B Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Upp Shu Low Hum 5C Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum

5A

As 8

9

10

5B- 11 5D 5B- 11 5D

(continued)

166

Rel. 96 97 98 99 101 102 103 104 112 90 46 116 119 100

Appendix A

Alt 1488 1519 1528 1545 1577 1585 1600 1620 1285 1338 1347 1375 1417 1561

T 14.7 14.56 14.51 14.43 14.27 14.23 14.15 14.05 15.72 15.46 15.4 15.27 15.06 14.35

P 1472 1472 1472 1472 1472 1472 1472 1472 142 1472 1472 1472 1472 1472

M 19.22 19.16 19.14 19.11 19.05 19.03 19 18.96 19.63 19.5 19.5 19.45 19.36 19.08

M 2.81 2.66 2.61 2.53 2.37 2.33 2.25 2.15 3.82 3.56 3.5 3.37 3.16 2.45

Tp 1764 1747.2 1741.2 1731.6 1712.4 1707.6 1698 1686 1886.4 1855 1848 1832.4 1807 1722

It 367 364 363 361 357 356 354 352 391 385 384 381 376 359

Io 8.3 8.4 8.4 8.5 8.6 8.6 8.7 8.7 7.8 7.9 8 8 8.1 8.5

Th Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr Upp Mtr

Om Gr Low Hum 4 Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum Low Hum 6A Low Hum Low Hum Low Hum Low Hum Low Hum

As 12

     

Rel. Number of relevé, T Annual mean temperature, P Annual mean precipitation, Pp Annual positive precipitation, M Monthly mean máximum temperature of the coldest month, m Monthly mean minimum temperature of the coldest month, Tp Annual positive temperature, It Themicity index, Io Ombrothermic index, Low Mtr Lower Mesotropical, Upp Mtr Upper Mesotropical, Low Shu Lower SubHumid, Upp Shu Upper Subhumid, Low Hum Lower Humid, Gr. HCA groups (Fig 4.2), As Plant association number  Indicates non-adscribed relevés. Table is organized according to Fig 4.2

Appendix B

Floristic composition of the study area Species Abutilon grandifolium Acacia aroma Acacia caven Acalypha amblyodonta Acalypha boliviensis Acalypha communis Acalypha lycioides Acalypha plicata Achyrocline flaccida Adenostemma brasilianum Agalinis genistifolia Aldama mollis Allophylus edulis Alnus acuminata Alternanthera philoxeroides Anadenanthera colubrina var. cebil Anagallis arvensis Anredera cordifolia Aphelandra hieronymi Aralia soratensis Asclepias curassavica Austroeupatorium inulifolium Axonopus compressus Baccharis capitalensis Baccharis coridifolia Baccharis dracunculifolia Baccharis latifolia

Family Malvaceae Fabaceae Fabaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Asteraceae Asteraceae Orobanchaceae Asteraceae Sapindaceae Betulaceae Amaranthaceae Fabaceae Primulaceae Bassellaceae Acanthaceae Araliaceae Apocinaceae Asteraceae Poaceae Asteraceae Asteraceae Asteraceae Asteraceae

Str a A A a h h a a a h a a A A h A h h a A h a h a a a a

Lf Naf Mif Mif Naf Th Hc Naf Naf Ca Th Naf Ca Mif Mif Hc Mef Th Hc Naf Mif Hc Naf Geo Naf Naf Naf Naf

Stat Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Ex Na Na Na Ex Na Na Na Na Na Na

Freq 16 56 28 32 8 26 8 31 9 24 6 6 84 9 5 56 6 31 7 7 3 8 20 22 11 8 18 (continued)

© Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3

167

168

Species Baccharis microdonta Baccharis salicifolius Baccharis trimera Barnadesia odorata Begonia boliviensis var. boliviensis Begonia micranthera var. micranthera Berberis jobii Bidens pilosa Bidens squarrosa Bidens subalternans Blepharocalyx salicifolius Boehmeria caudata Bomarea edulis Borreria spinosa Bougainvillea stipitata Bromelia serra Bromus catharticus Buddleja stachyoides Budleja diffusa Budleja iresinoides Caiophora hibiscifolia Calceolaria chelidonioides Calceolaria elatior Calceolaria teucrioides Campovassouria cruciata Cantinoa mutabilis Capsicum chacoense Carica glandulosa Carica quercifolia Cedrela angustifolia Cedrela saltensis Celtis ehrenbergiana var. discolor Celtis iguanaea Cenchrus latifolius Cestrum parqui Chamissoa altissima Chamissoa maximiliani Chaptalia nutans Chiropetalum boliviense Chloroleucon tenuiflorum Chromolaena laevigata

Appendix B

Family Asteraceae Asteraceae Asteraceae Asteraceae Begoniaceae Begoniaceae Berberidaceae Asteraceae Asteraceae Asteraceae Myrtaceae Urticaceae Alstroemeriaceae Rubiaceae Nyctaginaceae Bromeliaceae Poaceae Budlejaceae Budlejaceae Budlejaceae Loasaceae Calceolariaceae Calceolariaceae Calceolariaceae Asteraceae Lamiaceae Solanaceae Caricaceae Caricaceae Meliaceae Meliaceae Celtidaceae Celtidaceae Poaceae Solanaceae Amaranthaceae Amaranthaceae Asteraceae Euphorbiaceae Fabaceae Asteraceae

Str a a a a h h a h a h A a h h A h h a a a h h h h a h a a A A A A A h a a a h a A a

Lf Naf Naf Naf Naf Ge Ge Naf Th Naf Th Mef Naf Hc Hc Mif Ca Hc Naf Naf Naf Hc Hc Hc Hc Naf Hc Naf Naf Mif Mef Mef Mif Mif Geo Naf Naf Naf Hc Naf Mif Naf

Stat Na Na Na Na Na Na End Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na Na End Na Na Na Na Na Na Na Na Na

Freq 29 5 5 48 13 5 2 22 10 18 58 29 7 2 6 13 19 28 12 12 17 2 5 2 8 31 12 19 5 28 2 60 24 6 40 43 1 14 3 30 13 (continued)

Appendix B

Species Chrysophyllum marginatum Cinnamomum porphyrium Citrus sp. Clematis haenkeana Clinopodium bolivianum Cnicothamnus lorentzii Cnidoscolus tubulosus Cnidoscolus vitifolius Collaea argentina Condalia buxifolia Conyza sumatrensis Conyza tunariensis Cordia saccelia Cortaderia hieronymi Cortaderia selloana Coutarea hexandra Croton saltensis Cuphea racemosa Dendrophorbium bomanii Desmodium affine Desmodium subsericeum Deyeuxia polygama Dicliptera squarrosa Digitaria insularis Dolichandra ungis-cati Duchesnea indica Duranta serratifolia Elephantopus mollis Eleusine indica Enterolobium contortisiliquum Erythrina falcata Escallonia millegrana Eucalyptus sp. Festuca hieronymi Festuca superba Fleischmannia schickendantzii Galinsoga caracasana Galium hypocarpium Galium lilloi Galium richardianum Gamochaeta pensylvanica

169

Family Sapotaceae Lauraceae Rutaceae Ranunculaceae Lamiaceae Asteraceae Euphorbiaceae Euphorbiaceae Fabaceae Rhamnaceae Asteraceae Asteraceae Boraginaceae Poaceae Poaceae Rubiaceae Euphorbiaceae Lythraceae Asteraceae Fabaceae Fabaceae Poaceae Acanthaceae Poaceae Bignoniaceae Rosaceae Verbenaceae Asteraceae Poaceae Fabaceae Fabaceae Escalloniaceae Myrtaceae Poaceae Poaceae Asteraceae Asteraceae Rubiaceae Rubiaceae Rubiaceae Asteraceae

Str A A A a a A a a a A h h A h h A a h a h h h h h A h A h h A A A A h h h h h h h h

Lf Mif Mef Mif Lia Naf Mif Naf Naf Naf Mif Th Th Mif Hc Hc Mif Naf Hc Naf Hc Hc Geo Geo Geo Lia Hc Mif Geo Th Mef Mef Mif Mef Hc Hc Hc Th Hc Hc Th Hc

Stat Na Na Ex Na Na Na Na Na Na Na Na Na Na Na Na Na End Na Na Na Na End Na Na Na Ex Na Na Ex Na Na Na Ex Na End Na Na Na End Na Na

Freq 2 38 3 28 28 18 8 1 8 46 15 8 1 18 13 2 24 25 43 12 21 7 20 18 29 25 9 52 13 25 31 10 4 8 3 30 27 13 8 2 11 (continued)

170

Species Geoffroea decorticans Glandularia tweedieana Gorgonidium vermicidum Hebanthe occidentalis Heimia montana Heteropterys sylvatica Hydrocotyle bonariensis Hypochaeris microcephala Ilex argentina Iresine diffusa Jacaranda mimosifolia Juglans australis Jungia pauciflora Jungia polita Justicia goudotii Justicia kuntzei Justicia mandonii Kaunia lasiophthalma Koanophyllon solidaginoides Lantana canescens Lantana trifolia Leonurus japonicus Lepechinia vesiculosa Leptochloa virgata Lippia suffruticosa Ludwigia peruviana Malvastrum coromandelianum Manetia jorgensenii Manihot grahami Mikania micrantha Mimosa xanthocentra Mimosa debilis Mimosa polycarpa Modiolastrum malvifolium Morus alba Muehlenbeckia sagittifolia Muhlenbergia schreberi Mirabilisjalapa Myrcianthes pseudomato Myrcianthes pungens Myroxylon peruiferum

Appendix B

Family Fabaceae Verbenaceae Araceae Amaranthaceae Lythraceae Malpighiaceae Apiaceae Asteraceae Aquifoliaceae Amaranthaceae Bignoniaceae Juglandaceae Asteraceae Asteraceae Acanthaceae Acanthaceae Acanthaceae Asteraceae Asteraceae Verbenaceae Verbenaceae Lamiaceae Lamiaceae Poaceae Verbenaceae Onagraceae Malvaceae Rubiaceae Euphorbiaceae Asteraceae Fabaceae Fabaceae Fabaceae Malvaceae Moraceae Polygonaceae Poaceae Nyctaginaceae Myrtaceae Myrtaceae Fabaceae

Str A h h a a a h h A a A A h a h a a A a a a h a h a a a a A h h a a h A a h h A A A

Lf Mif Hc Geo Naf Naf Naf Hc Hc Mif Naf Mif Mef Hc Naf Hc Naf Naf Mif Naf Naf Naf Hc Naf Hc Naf Naf Naf Naf Mif Hc Hc Naf Naf Hc Mif Naf Hc Geo Mif Mif Mef

Stat Na Na Na Na End Na Na Na Na Na Na Na Na Na Na Na End Na Na End End Ex Na Na Na Na Cosmo End Na Na Na Na Na Na Ex Na Na Ex Na Na Na

Freq 8 10 8 16 25 21 3 15 3 7 13 33 54 37 54 18 17 20 11 32 10 19 6 3 4 7 19 13 8 28 21 19 17 25 5 16 8 28 25 9 4

(continued)

Appendix B

Species Myrsine laetevirens Nicandra physalodes Oenothera rosea Onoseris alata Ophryosporus lorentzii Ophryosporus piquerioides Oplismenus hirtellus Orthopappus angustifolius Panicum trichanthum Parapiptadenia excelsa Parthenium hysterophorus Paspalum distichum Pavonia sepium Petiveria alliacea Petunia occidentalis Pharus lappulaceus Phenax laevigatus Phytolacca bogotensis Pilea jujuyensis Piper hieronymi Pisonia zapallo Plantago australis Podocarpus parlatorei Polygonum punctatum Praxelis clematidea Primula malacoides Prosopis alba Prunus tucumanensis Pseudechinolaena polystachya Randia micrantha Rivinia humilis Rubus imperialis Ruellia ciliatiflora Ruellia erythropus Salix humboldtiana Salpichroa origanifolia Salvia personata Sambucus nigra ssp. peruviana Samolus valerandi Sapium haematospermum Schinus bumeloides

171

Family Myrsinaceae Solanaceae Onagraceae Asteraceae Asteraceae Asteraceae Poaceae Asteraceae Poaceae Fabaceae Asteraceae Poaceae Malvaceae Phytolaccaceae Solanaceae Poaceae Urticaceae Phytolaccaceae Urticaceae Piperaceae Nyctaginaceae Plantaginaceae Podocarpaceae Polygonaceae Asteraceae Primulaceae Fabaceae Rosaceae Poaceae Rubiaceae Phytolaccaceae Rosaceae Acanthaceae Acanthaceae Salicaceae Solanaceae Lamiaceae Adoxaceae Samolaceae Euphorbiaceae Anacardiaceae

Str A h h h a a h h h A h h a h h h a h h a A h A h h h A A h A h a h h A h h A h A A

Lf Mif Th Hc Hc Naf Naf Geo Hc Geo Mef Th Geo Naf Hc Th Geo Naf Hc Hc Naf Mif Hc Mef Th Th Hc Mif Mif Geo Mif Hc Naf Hc Hc Mif Ge Hc Mif Hc Mif Mif

Stat Na Na End Na End Na Na Na Na Na Na Na Na Na End Na Na Na End Na Na Na Na Na Na Ex Na Na Cosmo Na Na Na Na Na Na Na Na Na Na Na End

Freq 19 9 14 9 6 9 8 4 13 61 33 3 16 40 3 22 18 23 7 6 3 13 6 8 31 8 2 12 14 6 40 47 10 8 4 17 29 7 19 28 23

(continued)

172

Species Schinus fasciculatus Schinus gracilipes Schinus myrtifolius Scoparia ericacea Scutia buxifolia Sebastiania brasiliensis Sebastiania commersoniana Seemannia gymnostoma Senecio hieronymi Senecio rudbeckiifolius Senna occidentalis Senna pendula var. eriocarpa Senna spectabilis Setaria parviflora Sibthorpia conspicua Sibthorpia repens Sida cabreriana Sida rhombifolia Sinningia warmingii Smilax campestris Solanum abutiloides Solanum aligerum Solanum aloysiifolium Solanum betaceum Solanum confusum Solanum lorentzii Solanum palinacanthum Solanum riparium Solanum sisymbriifolium Solanum tenuispinum Stevia jujuyensis Stevia yaconensis var. subeglandulosa Stillingia tenella Tagetes filifolia Tagetes terniflora Tecoma stans Terminalia triflora Tessaria dodoneifolia Tessaria integrifolia Thalictrum venturii Tibouchina paratropica

Appendix B

Family Anacardiaceae Anacardiaceae Anacardiaceae Plantaginaceae Rhamnaceae Euphorbiaceae Euphorbiaceae Gesneriaceae Asteraceae Asteraceae Fabaceae Fabaceae Fabaceae Poaceae Plantaginaceae Plantaginaceae Malvaceae Malvaceae Gesneriaceae Smilacaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Solanaceae Asteraceae Asteraceae Euphorbiaceae Asteraceae Asteraceae Bignoniaceae Combretaceae Asteraceae Asteraceae Ranunculaceae Melastomataceae

Str A A A h A A A h a a a a A h h h h a h a a a h a a a a A h h h h A h h A A a A a h

Lf Mif Mif Mif Hc Mif Mif Mif Hc Naf Naf Naf Naf Mif Geo Hc Hc Hc Naf Hc Ca Naf Naf Hc Mif Naf Naf Naf Mif Hc Hc Hc Hc Mif Th Th Mif Mef Naf Mif Naf Ca

Stat Na End Na End Na Na Na Na Na Na Na End Na Na Na Na Na Cosmo Na Na Na End Na Na End Na End Na Na End End End Na Na Na Na Na Na Na Na Na

Freq 7 32 25 7 35 79 50 6 13 38 8 38 23 3 8 1 18 18 4 38 5 8 20 7 9 36 8 18 5 15 3 10 8 13 39 32 3 4 3 8 25

(continued)

Appendix B

Species Tipuana tipu Tournefortia paniculata Tradescantia boliviana Tragia volubilis Trema micrantha Trixis grisebachii Turnera sidoides Urera baccifera Urtica chamaedryoides Valeriana effusa Vassobia breviflora Verbascum virgatum Verbena litoralis Verbesina macrophylla var. nelidae Verbesina suncho Vernonanthura pinguis Vernonanthura squamulosa Veronica arvensis Veronica persica Viguiera tucumanensis var. oligodonta Wedelia saltensis Xylosma pubescens Zanthoxylum coco Zanthoxylum petiolare Zinnia peruviana

173

Family Fabaceae Boraginaceae Commelinaceae Euphorbiaceae Celtidaceae Asteraceae Turneraceae Urticaceae Urticaceae Valerianaceae Solanaceae Scrophulariaceae Verbenaceae Asteraceae Asteraceae Asteraceae Asteraceae Plantaginaceae Plantaginaceae Asteraceae Asteraceae Salicaceae Rutaceae Rutaceae Asteraceae

Str A a h h A a h a h h A h h a a a a h h h a A A A h

Lf Mef Naf Hc Hc Mif Naf Hc Naf Hc Th Mif Hc Hc Naf Ca Naf Naf Th Th Hc Naf Mif Mif Mif Th

Stat Na Na Na Na Na Na Na Na Na Na Na Ex Na Na Na Na Na Ex Ex Na Na Na Na Na Na

Freq 47 4 16 10 6 6 3 59 26 3 61 3 13 28 18 30 66 6 5 14 17 48 16 11 22

Str Stratum, Lf Life form, Stat Status, Freq Frequency (%). Layers: A tree, a shrub, h herbaceous. Biotypes: Th therophyte, Geo geophyte, Hc hemicryptophyte, Ca chamaephyte, Naf nanophanerophyte, Mif microphanerophyte, Mef mesophanerophyte, Lia creeper. Status: Na native species, Ex exotic species, Cosmo cosmopolitan species, End endemic species

Appendix C

General comparative synthetic table Associations N Cluster N N relevés N species N characteristic species Altitude range (m asl)

1 1A 8 93 16 10321037

2 2B 10 99 11 10851102

3 2C 10 92 7 11051127

4 2A 6 79 11 11921200

5 1C 12 138 12 11151258 MISS MISS MISS MISS HI

6 3A 5 46 4 12351243

7 3B 11 74 8 12391253

8 1B 15 183 7 12331275

9 5C 7 127 10 12601310

10 5A 8 77 4 13191360

11 5B-5D 12 165 22 13201433

12 4 8 111 18 14881620

MISS MISS MISS MSHI

MSHI

MSHI

MSHI

13 6B 2 45 11 10151030 MISI HI

Bioclimatic belt

MISI

Altitudinal vegetation belt

SMb

SMb

SMb

SMb

SMb

SMb

SMb

SMb

SMa

SMa

SMa

BM

SMrip

V III V IV IV V V V III IV V V V II . V II . . . . II . III V . . . . II I

II . V IV . IV V III . I V V V V V V . II V II II . . I II I . . . . . . . . . . . . .

. . III . . V V IV . II II II IV V . V V V I I II . . I I . . . . . . . . . . . . . .

I . V V . I III V . . V V II V . V . . V II V V I II I . . . . II . . . . . . . . .

III I V II . IV V IV . . V III . V . IV III III III I I . I IV V III III III III IV . I II . . . . . .

2 . . . . . 5 5 . . 1 5 . 5 . 2 2 . 2 . . . . . . 5 5 . . . . . . . . . . . .

I . V . . I V IV . . I V . V . IV V V I . . . . I I V V V . . . . . . . . . . .

V . V I . IV V IV . I V V . V . II IV V III I . . . IV V II IV IV IV IV . II IV . . . I II .

V . II I . II V . . II V . . III . . . . . . . . . II IV II . . . V III . . II . . II I .

II . . . . V IV . . .

V . II II . V IV . . . I III . V . . . V . . . . . I IV II . II . III I IV V III III I II IV III

III . . . . II . . . .

1 . 2 1 . 2 2 . . . 2 1 . 1 . . . . . . . . . . . . . . . 1 1 . . . . . . . .

Characteristic trees: Anadenanthera colubrina var. cebil1 Carica quercifolia2 Celtis iguanaea1 Enterolobium contortisiliquum2 Myroxylon peruiferum4 Parapiptadenia excelsa1 Sebastiania brasiliensis1 Sebastiania commersoniana1 Terminalia triflora4 Zanthoxylum petiolare2 Acacia aroma1 Vassobia breviflora1 Schinus bumeloides2 Allophylus edulis1 Manihot grahami3 Xylosma pubescens2 Scutia buxifolia1 Blepharocalyx salicifolius1 Chloroleucon tenuiflorum2 Geoffroea decorticans2 Jacaranda mimosifolia2 Schinus fasciculatus2 Prosopis alba2 Erythrina falcata2 Tipuana tipu1 Schinus gracilipes1 Schinus myrtifolius2 Juglans australis1 Zanthoxylum coco2 Tecoma stans2 Trema micrantha2 Myrcianthes pseudomato1 Cinnamomum porphyrium1 Aralia soratensis2 Bougainvillea stipitata2 Cedrela saltensis2 Duranta serratifolia2 Kaunia lasiophthalma2 Myrcianthes pungens2

I . I . . . .

. . IV . II . V . . . . . I I I . I . . . V V . . . . II II

I . V . . II IV . . . . . I . IV . V I . . III III . I . II III II

(continued) © Springer Nature Switzerland AG 2020 G. S. Entrocassi et al., Subtropical Mountain Forests of Las Yungas: Vegetation and Bioclimate, Geobotany Studies, https://doi.org/10.1007/978-3-030-25521-3

175

176 Association N Stillingia tenella2 Cedrela angustifolia Alnus acuminata2 Ilex argentina2 Podocarpus parlatorei1 Prunus tucumanensis2 Sambucus nigra ssp. peruviana3 Salix humboldtiana5 Tessaria integrifolia5 Characteristic shrubs and herbs: Rivinia humilis1 Solanum aloysiifolium1 Elephantopus mollis1 Urera baccifera1 Mirabilisjalapa1 Samolus valerandi1 Capsicum chacoense2 Senna occidentalis2 Barnadesia odorata1 Dendrophorbium bomanii1 Modiolastrum malvifolium2 Acalypha amblyodonta2 Baccharis coridifolia2 Cantinoa mutabilis2 Solanum palinacanthum2 Turnera sidoides4 Jungia polita1 Senna pendula var. eriocarpa1 Jungia pauciflora1 Thalictrum venturii2 Oplismenus hirtellus2 Petiveria alliacea1 Pharus lappulaceus1 Verbesina macrophylla var. nelidae2 Verbesina suncho2 Vernonanthura pinguis2 Justicia mandonii2 Acalypha communis2 Acalypha plicata2 Aphelandra hieronymi2 Baccharis latifolia2 Onoseris alata2 Petunia occidentalis4 Phenax laevigatus2 Phytolacca bogotensis2 Piper hieronymi2 Solanum betaceum2 Austroeupatorium inulifolium2 Berberis jobii4 Calceolaria teucrioides4 Campovassouria cruciata2 Clinopodium bolivianum2 Lepechinia vesiculosa2 Ophryosporus lorentzii2 Sibthorpia conspicua2 Solanum aligerum2 Solanum confusum2 Stevia yaconensis var. subeglandulosa2 Tibouchina paratropica2 Alternanthera philoxeroides5 Asclepias curassavica5 Baccharis salicifolius5 Paspalum distichum2 Polygonum punctatum5 Tessaria dodoneifolia5 Other species:

Appendix C 1 . II . . . . . . .

2 . . . . . . . . .

3 . . . . . . . . .

4 . . . . . . . . .

5 . I . . . . . . .

6 . . . . . . . . .

7 . . . . . . . . .

8 . II . . . . . I I

9 II I . . . . . . .

10 . III . . . II . . .

11 II IV I I . I . . .

12 III V V II IV V V . .

13 . . . . . . . 2 2

V V V V I I . . V II I . . I . . . III II . . IV II III II . . II I . . . . II I . . . . . . . . . . . . . . . . . . . .

II . V V IV IV III I III IV IV II I II . . I III III . . II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

II I III V IV . IV III III III I II . III . . . III II . . III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

I .

II . IV V . III . . V III IV IV III IV III II V III III . . II . I I IV . . . . . . . . . . . . . . . I . . . . . II . . . . . I .

. . . 2 3 . . . 1 2 1 . 1 . 1 . 4 2 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . II II . . . II III I II I III I . V V V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

III I IV V II I . . V II I V I II . . IV III V II I III . III . II . II II . . . . . II

V IV

. . IV I . . . . II I

I . II . . III II IV . V V V V V V II IV V . II . . III III . . . . . . II . . . . II . II . . . . . .

II . I . . . I II . . III II II IV III V II V . . . . II . . . . . . II II . . . . . . . . . . . . .

V III V III II II . . . III I . . I . . . II I I . V V V III IV V V V III V III II IV V III II I . . . III . I II II II I IV I . . . I .

1 . . 2

II . . . IV

II . . . . . . . . II

. I . I I V V V II . IV I . III III IV . . . . . . II . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

I . . . . IV . . . . . . . I I II I I I

II

. . . . . . . II II . . . . . . I II II I IV I IV II I II V II II IV IV IV IV IV IV IV V V . . . . . .

1 . . . . . 1 . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 2 2 2 2

(continued)

Appendix C Association N Abutilon grandifolium Acacia caven Acalypha lycioides Achyrocline flaccida Adenostemma brasilianum Aldama mollis Anredera cordifolia Axonopus compressus Baccharis capitalensis Baccharis microdonta Bidens pilosa Bidens squarrosa Bidens subalternans Boehmeria caudata Bromus catharticus Buddleja stachyoides Budleja diffusa Budleja iresinoides Cajophora hibiscifolia Carica glandulosa Celtis ehrenbergiana var. discolora Cenchrus latifolius Cestrum parqui Chamissoa altissima Chaptalia nutans Chiropetalum boliviense Chromolaena laevigata Clematis haenkeana Cnicothamnus lorentzii Cnidoscolus tubulosus Condalia buxifolia Conyza sumatrensis Conyza tunariensis Cortaderia hieronymi Croton saltensisa Cuphea racemosa Desmodium affine Desmodium subsericeum Dicliptera squarrosa Digitaria insularis Dolichandra ungis-cati Duchesnea indica Eleusine indica Fleischmannia schickendantzii Galinsoga caracasana Galium hypocarpium Galium lilloia Gamochaeta pensylvanica Glandularia tweedieana Gorgonidium vermicidum Hebanthe occidentalis Heimia montanaa Heteropterys sylvatica Hypochaeris microcephala Justicia goudotii Justicia kuntzei Koanophyllon solidaginoides Lantana canescensa Leonurus japonicus Malvastrum coromandelianum

177 1 . III . . I . II . II . . . II II . . . . . I . I IV II II . . II . I . . . . . I I . I . II II . II III . . . . . I . . . IV . . . I .

2 . IV . . . . I I II II II . I . II II . II II I I . III V . . . III III . V . . . II I . II . I . . I III II . . III II . I . . III III . . III II II

3 . . . . . . II II III II I . I . II II . I II II IV . III II . . . I . . IV . . . . II . II I I . . I II II . . II II . II II I . II . . II I .

4 . II . . . . I . IV IV III . III . IV III III . . . . . V I . . . III . . III . III . . III I . . I . . III . II . . II III . . . . . I . . . V III

5 II IV . I . I III . III II IV . III I III III II II I II III . III III . . I III I . IV I II I II III . III . . III . I I III . . I II I . . II III III . . II III II

6 . . . . . . . . . 2 1 . 2 . . 2 . . . . . . . 1 . . . 2 . . 5 . . . 2 . . . . 1 . . 3 2 . . . . . . . . . 2 2 . . . . 3

7 . . . . . . . I . II I . . . I I . . I . II . . III . . . II . . V . I . I . . . . I I . I II II . . . . II . . III II III . . . I III

8 IV IV I II III I III III II II II II I IV I III II II II III IV . IV IV II I II I III I III II I III IV II . I II II III II I III I I I I . II II IV II I IV . II III II II

9 II . II I V II IV II . II . I . II . I III II . I . . . III III I III III . II . II . II II III III II III IV III III . III I II I . . . III II III I III V I V . .

10 . . II . I . . . . I . . . I . . . . . . . I . II . I . . II . . . . II . . . . . . III . . I . I . . . . I . I . V IV . II . .

11 II . II I IV I III II . II . II . V . I . . II I . I IV II III I III . . II . III . I II IV III IV III II III III . I III II II . . I II III II . V IV III I . I

12 I . . II II II II III . II . II . IV . II . . II . . II I . II . . . IV I . III . I I . II I II I II V . . II III I . . . . II . . I I II II . .

13 . 1 . . . . 1 . . . . . . . 1 . . . . 1 . . . . . . . 1 . . . . . . . 1 . . 2 . . 1 . . 1 . . . . . . . . . 1 . . . . .

(continued)

178

Appendix C

Association N 1 2 3 4 5 6 7 8 9 10 11 12 13 Manetia jorgenseniia . . . . . . . II III . II II . Mikania micrantha III I II V . . III III . . II II . Mimosa xanthocentra . II I IV II 1 I I II II I I 1 Mimosa debilis . III I . III . II I III . I . . Mimosa polycarpa II . . V . . . II . I II . . Morus alba II . . . I . I I . . . . . Muehlenbeckia sagittifolia . III I . . . . I . I II IV . Muhlenbergia schreberi . . . . . . . I . I III II . Myrsine laetevirens II . . . I . . III . IV I III . Nicandra physalodes I . . III II 1 I . . . . . 1 . II III . I . I II . . I . . Oenothera roseaa Ophryosporus piquerioides . . I . I . . . . I II II . Panicum trichanthum . . I . . . I II . II III . . Parthenium hysteriophorus II IV III V II 2 II I II . I II 1 Pavonia sepium IV . I . I . . . III . II III . Plantago australis . II I . II . . I II I II . . Praxelis clematidea II V II III III 3 III I I . . . 1 Pseudechinolaena polystachya . I I . . . . I III I III . . Rubus imperialis II III III III I 4 III II III IV III V . Ruellia ciliatiflora . I II . . . . . I I III . . Ruellia erythropus III . . . I . . . III . I . . Salpichroa origanifolia . III II . II . . . III I II . . Salvia personata II I I . III . . III V . IV . . Sapium haematospermum V I I . IV . I III . . II . . Senecio hieronymi . . . . . . I II I I I III . Senecio rudbeckiifolius . III III . IV 1 I III III II II III . Senna spectabilis . III I I III . II III . . . . . Sida cabreriana III I I . . . . II III I II . . Sida rhombifolia . III . III III . I I I . . . 1 Smilax campestris II IV I III III 1 II III I V . . 1 Solanum lorentzii V II II IV I . . II IV II III II . Solanum riparium II . . . I . . III . II III . . Solanum sisymbriifolium . I . I II . I . . . . . . . I II II . . . II II . II I . Solanum tenuispinuma Tagetes filifolia . . . . III 1 I I II . . . 1 Tagetes terniflora IV IV III . III 2 II V II . I . 1 Tradescantia boliviana II III . . I . . I II . I III 1 Tragia volubilis II . . . . . . . III . III I . Urtica chamaedryoides . . . . II . . III IV I III IV . Verbena litoralis II III III II . . . . . . . . . Vernonanthura squamulosa V IV II V V 4 IV V IV II IV II . Veronica arvensis . . . . I . . . I . II I . Bromelia serra II . . . I . . II . V . . . Veronica persica . . . . I . . I II I . . 1 Viguiera tucumanensis var. oligodonta . II II . III . . II . . . . . Weddelia saltensis . . II . . . . II III I IV . . Zinnia peruviana . II . II III 3 V I I . . . 1 Other species: Orthopappus angustifolius II in 1, I in 4, 11; Randia micrantha I in 1, 5, 8; Eucalyptus sp. I in 2-4; Anagallis arvensis and Iresine diffusa II in 4, 5, I in 8; Trixis grisebachii II in 4, 8, 12; Begonia micranthera, Escallonia millegrana and Festuca hieronymi II in 5, 8, 12; Begonia boliviensis var. boliviensis I in 5, II in 8, 11; Tournefortia paniculata I in 5, 9, 11; Baccharis dracunculifolia I in 7, II in 8, 12; Deyeuxia polygamaa II in 8, 10, 12; Primula malacoides, Scoparia ericaceaa and Setaria parviflora I in 8, 11, III in 9; Bomarea edulis, Ludwigia peruviana and Seemannia gymnostoma I in 8, 10, 11; Pisonia zapallo I in 9-11; Collaea argentina I in 4, II in 8; Baccharis trimera I in 5, 8; Cortaderia selloana I in 5, IV in 8; Lantana trifoliaa II in 8, IV in 9; Borreria spinosa, Calceolaria elatior, Leptochloa virgate and Stevia jujuyensisa I in 8, 11; Pilea jujuyensisa I in 8, III in 12; Hydrocotyle bonariensis and Salix humboldtiana I in 8, 13; Chrysophyllum marginatum, Coutarea hexandra, Solanum abutiloides and Valeriana effuse I in 9, 11; Acalypha boliviensis III in 9, 11; Agalinis genistifolia, Citrus sp. and Galium richardianum I, Verbascum virgatum II in 5; Calceolaria chelidonioides, Chamissoa maximiliani, Festuca superbaa and Lippia suffruticosa I in 8; Cordia saccelia I in 9; Cnidoscolus vitifolius I, Sinningia warmingii II in 11. Associations and clusters (in brackets): 1: Enterolobio contortotisilici-Anadenantheretum cebilis (1A); 2: Schino bumeloidis-Allophyletum edulis (2B); 3: Xylosmo pubescentis-Blepharocalycetum salicifolii (2C); 4: Jacarando mimosifoliae-Vassobietum breviflorae (2A); 5: Erythrino falcatae-Tipuanetum tipi (1C); 6: Schinetum myrtifolio-gracilipedis (3A); 7: Juglandi australis-Blepharocalycetum salicifolii (3B); 8: Zanthoxylo cocoi-Blepharocalycetum salicifolii (1B); 9:Tecomo stantis-Anadenantheretum cebilis (5C); 10: Myrciantho pseudomatoi-Blepharocalycetum salicifolii (5A); 11: Cinnamomo porphyrium-Blepharocalycetum salicifolii (5B-5D); 12: Pruno tucumanensis-Podocarpetum parlatorei (4); 13: Salici humboldtianae-Acacietum aromae (6B). Bioclimatic belts: MISI: Lower Mesotropical-lower Subhumid; MISS: Lower Mesotropical-upper Subhumid; MSHI: Upper Mesotropical-lower Humid; MISI and HI: Lower Mesotropical-lower Subhumid and lower Humid. Altitudinal vegetation belt: SMb: Basal mountain forest; SMa: High mountain forest; BM: Mountain woodland; SMrip: Riparian mountain forest. a

Endemic spec.; Characteristic or indicator spec.: 1Dominant, 2Selective or preferential, 3Exclusive, 4Differential, 5Stenoic.

Associations and clusters of relevés: 1: Enterolobio contortotisilici-Anadenantheretum cebilis (1A); 2: Schino bumeloidis-Allophyletum edulis (2B); 3: Xylosmo pubescentis-Blepharocalycetum salicifolii (2C); 4: Jacarando mimosifoliae-Vassobietum breviflorae (2A); 5: Erythrino

Appendix C

179

falcatae-Tipuanetum tipi (1C); 6: Schinetum myrtifolio-gracilipedis (3A); 7: Juglandi australisBlepharocalycetum salicifolii (3B); 8: Zanthoxylo cocoi-Blepharocalycetum salicifolii (1B); 9: Tecomo stantis- Anadenantheretum cebilis (5C); 10: Myrciantho pseudomatoi-Blepharocalycetum salicifolii (5A); 11: Cinnamomo porphyrium-Blepharocalycetum salicifolii (5B–5D); 12: Pruno tucumanensis-Podocarpetum parlatorei (4); 13: Salici humboldtianae-Acacietum aromae (6B) Bioclimatic belts: MI-SI Lower Mesotropical - lower Subhumid, MI-SS Lower Mesotropicalupper Subhumid, MS-HI Upper Mesotropical- lower Humid, MI-SI and HI Lower Mesotropical-lower Subhumid and lower Humid. Altitudinal vegetation belt: SM b Basal mountain forest, SM a High mountain forest, BM Mountain woodland, SM rib Riparian mountain forest Characteristic or indicator species: 1: dominant; 2: selective or preferential; 3: exclusive; 4: differential; 5: stenoic; *: endemic

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Web Sites of Interest Fundación ProYungas. http://www.proyungas.org.ar SIGA Proyungas. http://siga.proyungas.org.ar/mapas/