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Land Use Changeand Mountain Biodiversity Maximo Liberman, and Christian Körner A CRC title, part of the Taylor & Francis imprint, a member of the Taylor & Francis Group, the academic div

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Part I Introduction

3523_book.fm Page 1 Tuesday, November 22, 2005 11:23 AM

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Land Use Change

and Mountain Biodiversity

Maximo Liberman, and Christian Körner

A CRC title, part of the Taylor & Francis imprint, a member of the Taylor & Francis Group, the academic division of T&F Informa plc.

Boca Raton London New York

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Published in 2006 by

CRC Press

Taylor & Francis Group

6000 Broken Sound Parkway NW, Suite 300

Boca Raton, FL 33487-2742

© 2006 by Taylor & Francis Group, LLC

CRC Press is an imprint of Taylor & Francis Group

No claim to original U.S Government works

Printed in the United States of America on acid-free paper

10 9 8 7 6 5 4 3 2 1

International Standard Book Number-10: 0-8493-3523-X (Hardcover)

International Standard Book Number-13: 978-0-8493-3523-5 (Hardcover)

Library of Congress Card Number 2005047213

This book contains information obtained from authentic and highly regarded sources Reprinted material is quoted with permission, and sources are indicated A wide variety of references are listed Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials

or for the consequences of their use.

No part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers

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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for

identification and explanation without intent to infringe.

Library of Congress Cataloging-in-Publication Data

Land use change and mountain biodiversity / [edited by] Eva Spehn, Maximo Liberman, and Christian Körner.

p cm.

Selected papers from 2 workshops, the first held in Moshi, Tanzania, Aug 19-24, 2002 and the second held in La Paz, Bolivia, Aug 20-23, 2003.

Includes bibliographical references.

ISBN 0-8493-3523-X (alk paper)

1 Mountain ecology Congresses 2 Land use Environmental aspects Congresses 3 Biological diversity Congresses I Körner, Christian, 1949- II Spehn, E M Eva M.) III Liberman, Máximo.

QH541.5.M65L36 2005

Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com

Taylor & Francis Group

is the Academic Division of Informa plc.

3523_Discl.fm Page 1 Wednesday, November 23, 2005 3:10 PM

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Preface

SUSTAINABLE USE AND BIODIVERSITY OF SUBTROPICAL AND TROPICAL

HIGHLANDS

Within the worldwide biodiversity program of

DIVERSITAS, the Global Mountain Biodiversity

Assessment (GMBA) seeks to assess the biological

richness of high-elevation biota around the world

Mountains provide an excellent opportunity for a

global biodiversity research network, as they exist

in every climatic zone GMBA has a high-elevation

focus, including the uppermost forest regions or

their substitute rangeland vegetation, the treeline

ecotone, and the alpine and the nival belts

Although acknowledging the significance of

lower-montane biota, they fall outside the GMBA

agenda Beyond description, GMBA aims at

explaining the causes of biological richness in

mountains and its change over time Given that

changes in biodiversity most often result from

human land use, one specific GMBA agenda is the

assessment of land use impacts Such assessments

have priority in low-latitude regions, where land

use pressure on upland biota is greatest Upland

grazing, often facilitated by fire management, is

the most widespread utilization of mountain

ter-rain, often followed by erosion and enhanced risk

for valley and foreland environments

High-eleva-tion forests have disappeared in most regions, and

the few relicts are under intense use Cultivation

of formerly pristine areas and intensification of

agriculture in montane areas are often associated

with a loss of mountain biodiversity Both

prob-lems are most severe in the tropics and subtropics

This book is the second volume produced by

the Global Mountain Biodiversity Assessment

(GMBA) of DIVERSITAS, following Mountain

Biodiversity: A Global Assessment (eds Ch

Körner and E.M Spehn), published by Parthenon

in 2002 The chapters of this volume have been

selected in a peer-reviewing process from the

pre-sentations offered at two GMBA workshops, one

in Africa (Moshi, Tanzania, August 19 to August

24, 2002) and the other one in the Andes (La Paz,Bolivia, August 20 to August 23, 2003) More than

50 researchers actively participated, sharingknowledge from all major mountain regions, with

a particular focus on the Andes and the Africanmountains The two workshops profited greatlyfrom the hospitality of the African Mountain Asso-ciation (AMA), which hosted the African work-shop at its sixth international conference on sus-tainable mountain development in Africa Wewould like to cordially thank Prof Salome Misana

of the Department of Geography, University of Dar

es Salaam, Tanzania, for the organization of theconference and for her local support and inputduring the first workshop The second workshop

in the Andes was locally organized by MaximoLiberman, SERNAP, in Huarina at the shore ofLake Titikaka in Bolivia, under the auspices of theAndean Mountain Association (AMA)

Under the patronage of, and with support from,DIVERSITAS, these workshops have been under-written by various agencies The workshops andthe synthesis process were generously funded bythe Swiss Agency for Development and Coopera-tion The Swiss Federal Office for Agricultureenabled the cooperation with the Swiss FederalResearch Station of Agroecology and Agriculture(Agroscope Zürich–Reckenholz) on this project.The Food and Agriculture Organization (FAO) ofthe United Nations supported the preparation ofthis publication through the FAO/NetherlandsPartnership Programme “Assessment of Agricul-tural Biodiversity.” SERNAP (Servicia Nacional

de Areas Protegidas de Bolivia)/ II Bolivia ported the Spanish edition of this volume, printed

sup-in Bolivia (SERNAP, La Paz, 2005)

We wish to thank the following persons whohelped with the editing of this volume: AndreasGrünig of the Swiss Federal Research Station for

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vi Land Use Change and Mountain Biodiversity

Agroecology and Agriculture (Agroscope

Zürich–Reckenholz) for his valuable help in the

process of editing submitted manuscripts;

Anne-marie Brennwald, Sylvia Martinez, and Susanna

Pelaez-Riedl of the Institute of Botany, University

of Basel, for text editing and graphic support;

Emma Sayer, who translated chapters to English,

and Cecile Belpaire (La Paz, Bolivia), who

trans-lated chapters to Spanish in the Spanish edition

Under the auspices of the Swiss Academy of

Natural Sciences, the GMBA office in Basel,

Swit-zerland (Eva Spehn and Sylvia Martinez) weresupported by the Swiss Federal Office of Scienceand Education 2001–2003 and the Swiss NationalScience Foundation (SNF) (2004– )

Eva Spehn, Maximo Liberman, and

Christian Körner

Basel, Switzerland and La Paz, Bolivia

January 2005

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Contributors

Bhupendra Singh Adhikari

Wildlife Institute of India

Dehradun, India

Khukmatullo Akhmadov

Tajik Forestry Research and Development Institute

Dushanbe, Tajikistan

Humberto Alzérreca Angelo

Programa Estralégico de Acción para la Cunca del

Rio Bermejo (PEA-Bolivia)

Tarija, Bolivia

Roxana Aragón

Facultad de Agronomía

Universidad de Buenos Aires

Buenes Aries, Argentina

Yoseph Assefa

Department of Biology

Addis Ababa University

Addis Ababa, Ethiopia

Jorge Alberto Bustamante Becerra

Department of Ecology, Biosciences InstituteUniversity of São Paulo

São Paulo, Brazil

Konrad Fiedler

Population EcologyInstitute for Ecology and Conservation BiologyUniversity of Vienna

Vienna, Austria

Menassie Gashaw

Ethiopian Wildlife OrganizationAddis Ababa, Ethiopia

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viii Land Use Change and Mountain Biodiversity

Roger B Good

National Parks and Wildlife Service

Queanbeyan, New South Wales, Australia

Instituto de Ciencias Ambientales y Ecológicas

Universidad de Los Andes

School of Resources, Environment and Society

Australian National University

Andrea Corinna Mayer

Swiss Federal Institute for Snow and Avalanche Research

Davos, Switzerland

Marcelo Fernando Molinillo

Instituto de Ciencias Ambientales y EcológicasUniversidad de Los Andes

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Range and Forage Institute

Agricultural Research Council

Pietermaritzburg, South Africa

Jesus Orlando Rangel Churio

Instituto de Ciencias Naturales

Universidad Nacional de Colombia

Instituto de Ciencias Ambientales y Ecologicas

Universidad de los Andes

Núcleo la Hechicera, Facultad de Ciencias

Julia K Smith

Instituto de CienciasAmbiental y EcologicasUniversidad de los AndesMenda, Venezuela

Eva M Spehn

Global Mountain Biodiversity AssessmentInstitute of Botany

University of BaselBasel, Switzerland

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x Land Use Change and Mountain Biodiversity

New Haven, Connecticut

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Table of Contents

PART I Introduction 1

Christian Körner, Gia Nakhutsrishvili, and Eva M Spehn

PART II Effects of Fire on Mountain Biodiversity 23

in the Bale Mountains, Ethiopia and the Influence of Fire 25

Masresha Fetene, Yoseph Assefa, Menassie Gashaw, Zerihun Woldu, and Erwin Beck

Karsten Wesche

Vegetation on Mt Kilimanjaro 51

Andreas Hemp

Jan C Axmacher, Ludger Scheuermann, Marion Schrumpf, Herbert V.M Lyaruu, Konrad Fiedler, and Klaus Müller-Hohenstein

Their Small-Mammal Communities in Madagascar 77

Bernardin P.N Rasolonandrasana and Steven M Goodman

Grasslands of Northwest Argentina 89

Roxana Aragón, Julietta Carilla, and Luciana Cristóbal

PART III Effects of Grazing on Mountain Biodiversity 101

Anthropogenic Impact 103

Jesus Orlando Rangel Churio

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xii Land Use Change and Mountain Biodiversity

Richness in an Old-Field Succession of the Venezuelan Páramos 119

Lina Sarmiento

Systems in Punas and Páramos 137

Marcelo Molinillo and Maximina Monasterio

the Southeastern Andes of Peru (Palccoyo, Cusco) 153

Jorge Alberto Bustamante Becerra

of Key High-Andean Puna Rangelands (Bofedales) in Ulla Ulla, Bolivia 167

Humberto Alzérreca, Jorge Laura, Freddy Loza, Demetrio Luna, and Jonny Ortega

Zulimar Hernández and Maximina Monasterio

in Rangeland Vegetation 199

Gopal S Rawat and Bhupendra S Adhikari

and Stabilization of the Ecosystem 211

Ken Green, Roger B Good, Stuart W Johnston, and Lisa A Simpson

Siegmar-W Breckle and Walter Wucherer

Pastures in Tajik Mountains 239

Khukmatullo M Akhmadov, Siegmar W Breckle, and Uta Breckle

PART IV Effects of Grazing on Mountain Forests 249

Chapter 18 Plant Species Diversity, Forest Structure, and Tree Regeneration in

Subalpine Wood Pastures 251

Andrea C Mayer, Christine Huovinen, Veronika Stoeckli, and Michael Kreuzer

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Table of Contents xiii

Chapter 19 Patterns of Forest Recovery in Grazing Fields in the Subtropical Mountains

of Northwest Argentina 261

Julietta Carilla, H Ricardo Grau, and Agustina Malizia

Chapter 20 Climatic and Anthropogenic Influences on the Dynamics of Prosopis ferox

Forests in the Quebrada de Humahuaca, Jujuy, Argentina 275

Mariano Morales and Ricardo Villalba

Chapter 21 Conservation of Biodiversity in the Maloti–Drakensberg Mountain Range 285

Terry M Everson and Craig D Morris

Chapter 22 Effects of Anthropogenic Disturbances on Biodiversity: A Major Issue

of Protected-Area Management in Nepal 293

Khadga Basnet

Chapter 23 Agricultural Development and Biodiversity Conservation in the Páramo

Environments of the Andes of Mérida, Venezuela 307

Maximina Monasterio, Julia K Smith, and Marcelo Molinillo

Chapter 24 Multidimensional (Climatic, Biodiversity, Socioeconomic), Changes in

Land Use in the Vilcanota Watershed, Peru 319

Stephan Halloy, Anton Seimon, Karina Yager, and Alfredo Tupayachi

PART VI Synthesis 335

Chapter 25 Fire and Grazing — A Synthesis of Human Impacts on Highland Biodiversity 337

Eva M Spehn, Maximo Liberman, and Christian Körner

Chapter 26 The Moshi-La Paz Research Agenda on “Land Use Effects on Tropical

and Subtropical Mountain Biodiversity” 349

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1 High-Elevation Land Use,

Biodiversity, and Ecosystem Functioning

Christian Körner, Gia Nakhutsrishvili, and Eva Spehn

ANTHROPOGENIC HIGHLAND

ECOSYSTEMS

Humans have shaped much of the world’s lands over millennia Landscapes of sustainableproductivity, high biodiversity, and aestheticattractiveness have developed through livestockgrazing These landscapes also exhibit highecosystem stability, a key requisite for erosioncontrol and catchment quality (Körner, 2000,2004; Figure 1.1)

high-As a cultural heritage associated with tional-knowledge-based land management,many of these high-elevation pasture landscapes,hayfields, marginal crop fields, and rangelandsare of significant conservational and historicalvalue In some parts of the world, however, high-land management had no tradition (e.g., NewZealand and Australia), and when abruptly intro-duced to an unadapted flora, often had disastrousconsequences (e.g Costin 1958)

tradi-Over the last 50 years, these anthropogenichighland biota have undergone dramatic changesassociated with even more dramatic societal andeconomic changes, in addition to the atmo-spheric (climatic) changes underway In the morewealthy parts of the world, much of the high-lands have undergone extensivation of use orabandonment In the less economically privi-leged parts, population growth and land use pres-sure have often caused an expansion of agricul-tural land use into less suitable regions andabandonment of traditional land use practices

Both of these facets of global change have haddrastic influences on highland integrity andbiodiversity Unfortunately, both these depar-tures from the traditional middle ground of sus-tainable land use have caused a loss of biological

richness, and both tend to incur land degradation,though this is only a transitory risk in the case

of abandonment (e.g Tasser et al 2003) but isoften terminal in the case of overusing when soilsare washed away

In this overview of the ecological dimensions

of highland grazing, we will follow the simpleand common biogeographic nomenclature of ele-vational belts We will use the altitudinal position

of the natural upper-climatic treeline, defined asthe line connecting the uppermost pockets oftrees (i.e below the tree species line but abovethe forest line; Körner, 2003), as a reference (irre-spective of whether such forest patches arelocally present or not) We will define the moun-tain slopes below as montane and the naturallytreeless land above as alpine In this sense

“alpine” does not refer to the Alps but appliesglobally (following from its preIndo-Germanicmeaning of “steep slopes”), with “Andean” and

“Afroalpine” as synonyms The climatic vation treeline correlates worldwide with a sea-sonal mean temperature of 6.7 ± 0.8°C (indepen-dent of season length; Körner and Paulsen, 2004).Somewhat lower threshold temperatures (5 to6°C) can be found at the equator (treelines at

high-ele-3800 to 4100 m), but the thresholds in the tropics match with those at higher latitudes Inthe humid and semihumid tropics and subtropics,much of the high-elevation pastureland is foundbetween 500 and 800 m below and between 300and 400 m above the treeline elevation (i.e.between 3000 and 4400 m), with lower elevationscommonly used for crop production and higherelevations commonly carrying too little vegeta-tion and not regularly grazed In the NorthernHemisphere temperate zone, with treeline posi-tions varying widely between 1500 and 3500 m3523_book.fm Page 3 Tuesday, November 22, 2005 11:23 AM

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sub-4 Land Use Change and Mountain Biodiversity

depending on latitude and oceanic climate, the

corresponding amplitudes are wider, namely,

from at least 1000 m (1500 m in continental

ranges) below to 400 m above treeline These are

the elevational ranges considered here and in the

remainder of this volume when the term highland

is used As the focal elevations of GMBA are the

upper-montane, treeline ecotone, and alpine belts,

most of the contributions refer to these higher

parts of what could be considered highlands in

the widest sense

According to an assessment by Kapos et al

(2000; cf Körner, 2004), the global land area

above 1000 m and below 4500 m represents

14.3% of the terrestrial area Given that (1) in

the subtropics and tropics, much of the lower

part of this topography-based assessment falls

outside the climatic range of interest here, and

that (2) a great fraction of mountains falls in the

largely bare polar and subpolar regions, a

realis-tic estimate of the global land area fraction

suit-able for agricultural use in the highlands will be

somewhere around 8%, with 3% falling in the

alpine belt (Körner 1995), and the remaining

(around 5%) in the montane belt About 25% of

the montane land area is still forested according

to Kapos et al (2000), and a similar area may

be arid or barren, so that the nonforested, tially grazed montane and alpine highlands willcover roughly 5% of the global terrestrial area,

poten-an area as large as the polar tundra region(Körner, 1995) Approximately half of this arealies in the tropics and subtropics

As small as this area may look on a globalscale, it covers a very critical mountain zone Ithas been estimated that nearly half of humanitydepends directly or indirectly on the water yieldfrom mountain catchments (Messerli and Ives,1997; Messerli, 2004), with the vegetation-cov-ered upper-catchment regions playing a key rolefor clean and steady discharge In this sense,highlands control much of the so-called watertowers of the globe, and the functional integrity

of these highlands matters for land areas (andpopulations) by far exceeding their actual size(Figure 1.2) The slopes of these catchments areonly as stable as their green cover This coverneeds a high functional diversity of plants tofulfill its protective role under all sorts of unpre-dictable environmental conditions Thus theecology and richness of highland biota are inti-mately linked to the welfare of a large fraction ofhuman population, beyond their significance forlocal livelihoods (Körner and Spehn, 2002;

FIGURE 1.1 Fingerprints of millennia of land use in the highlands Examples of anthropogenic grassland from (a) Bolivian altiplano, 4100 m; (b) Cayambe region, 3700 m, Ecuador; (c) Sajama region bofedales, Bolivia,

4100 m; and (d) Spiti Valley, the Himalayas, 3700 m, India.

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High-Elevation Land Use, Biodiversity, and Ecosystem Functioning 5

Körner, 2004) Given that much of the tropical

and subtropical mountain forelands is rather dry,

this interdependency is even larger at

low-lati-tude regions The teleconnection between

high-land grazing grounds and metropolitan areas

may be thousands of kilometers as, for instance,

exists between the upper-Nile catchments and

Cairo or between eastern Anatolia and what was

Mesopotamia Sustainable highland

manage-ment, thus, has significant economic impact on

people living far outside the mountains

In this introductory chapter, we (1) aim to

the functioning of highland biota with specialreference to low latitudes, (2) will provide a briefsummary of previous observations on highlandpasture systems, and (3) will, then, open thearena for the global change implications forbiodiversity and ecosystem functioning in sub-tropical and tropical highlands, the main theme

of this volume

DRIVERS OF HIGHLAND ECOLOGY (WITH SPECIAL REFERENCE TO THE TROPICS AND SUBTROPICS)

The following is a brief reconsideration of themajor forces that shape upland biota These fallinto topography-related and climatic driversand biological determinants

Compression of climatic zones Mountainsare inhabited by more species of plants, animals,and microbes as one would estimate from theirland area and have often been called “hot spots”

of biodiversity (Körner, 2004) This has severalreasons intrinsically linked to topography andgravity Due to the elevational range covered,mountains encapsulate several climatic lifezones that would otherwise be separated by thou-sands of kilometers at low elevation (Barthlott et

al 1996) Hence, nowhere else on land can morebiological richness be encountered on a 100-km2scale than on the slopes of a high tropical moun-tain In relative terms, this effect also holds formountains in extratropical regions

Habitat diversity The second importantfactor at smaller scales is topographic diversity.Exposure, steepness of slope, variation of sub-strate, and microclimate over short distancescreate a multitude of microhabitats, each nest-ing a different set of organisms This habitatdiversity again permits aggregation of ratherdiverse biota over otherwise short distances.Gravity is the primary force behind this geodi-versity; where it lacks action, as in plains, irre-spective of elevation, biological diversitydeclines Because preferred grazing grounds areoften flat and smooth, their biological inventory

is commonly smaller than is found on the rounding slopes However, the species pool insuch plains could be even lower without grazingbecause grazing often creates “structure” bypatchy disturbance, dung deposition, food pref-erence, etc (Edwards et al 2004) Such effects

sur-FIGURE 1.2 Nearly half of mankind depends in one

way or the other on mountain water Highland

vegeta-tion is the safeguard of catchment quality and yield It

cleans, stores, and channels water to the lowlands.

Land use in these regions has far-ranging economic

consequences From top to bottom: upper catchment,

Bolivia, 4000 m; montane transgression, Sichuan, west

China; irrigation canal, lowland California.

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