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Full text of "World Atlas of Biodiversity: earth's living resources in the 21st century"

WORLD ATLAS OF 

BIODIVERSITY 

EARTH'S LIVING RESOURCES IN THE 21st CENTURY 



^ 



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UNEP WCMC 



BRIAN GROOMBRIDGE and MARTIN D. JENKINS 



World Atlas of Biodiversity addresses the remark- 



ible growth in concern at all levels for living things 



and the environment and the increased appreciation 



' the links between the state of ecosystems and 



the state of humankind. Building on a wealth of re- 



search and analysis by the conservation community 
worldwide, this book provides a comprehensive 
and accessible view of key global issues in biodiver- 
sity. It outlines some of the broad ecological 
relationships between humans and the rest of the 



iterial world and summarizes information on the 



health of the planet. Opening with an outline of 



some fundamental aspects of material cycles and 



energy flow in the biosphere, the book goes on to 



discuss the expansion of this diversity through geo- 
logical time and the pattern of its distribution over 



the surface of the Earth, and analyzes trends in the 



condition of the main ecosystem types and the 



species integral to them. 



Digitized by tine Internet Arciiive 

in 2010 witii funding from 

UNEP-WCIVIC, Cambridge 



Iittp://www.arcliive.org/details/worldatlasofbiod02groo 



World Atlas of Biodiversity 



Published in association witli 
UNEP-WCMC by the University of 
California Press 

University of California Press 
Berl<eley and Los Angeles, California 
University of California Press, Ltd. 
London, England 

© 2002 UNEP World Conservation 
Monitoring Centre 

UNEP-WCMC 
219 Huntingdon Road 
Cambridge CB3 DDL, UK 
Tel; *Uk 101 1223 277 3U 
Fax:+a(01 1223 277 136 
E-mail: info0unep-wcmc.org 
Website: www.unep-wcmc.org 

World Atlas of Biodiversity: 

Earth's Living Resources in the 21st Century 

IS a revised and updated edition of 

Global Biodiversity: 

Earth's Living Resources in the 21st Century 

No part of this book may be reproduced by 
any means or transmitted into a machine 
language without the written permission of 
the publisher 



Ihe contents of this volume do not 
necessarily reflect the views or policies of 
UNEP-WCfvIC, contributory organizations, 
editors or publishers. The designations 
employed and the presentations do not imply 
the expression of any opinion whatsoever on 
the part of UNEP-WCIvIC or contributory 
organizations, editors or publishers 
concerning the legal status of any country, 
territory, city or area or its authority, or 
concerning the delimitation of its frontiers or 
boundaries or the designation of its name or 
allegiances. 



Cloth edition ISBN 

0-520-23668-8 

Cataloging-in-publication data is on file with 
the Library of Congress 

Citation Groombndge B. and Jenkins M.D. 
120021 World Atlas of Biodiversity. Prepared 
by the UNEP World Conservation Monitoring 
Centre. University of California Press, 
Berkeley, USA. 






UNEP WCMC 




World Atlas of Biodiversity 

Earth's Living Resources in the 21st Century 



Brian Groombridge & Martin D. Jenkins 



UNIVERSITY OF CALIFORNIA PRESS 

Berkeley Los Angeles London 



World Atlas of Biodiversity 



Prepared by 

UNEP World Conservation 

Monitoring Centre 

219 Huntingdon Road 
Cambridge CB3 ODL, UK 
Tel: +M 101 1223 277 3U 
Fax: +44 (01 1223 277 136 
E-mail: info0unep-wcmc.org 
Website: www.unep-wcmc.org 

Director 

Marl< Collins 

Authors 

Brian Groombndge 
Martin D Jenkins 

Additional contributors 

Adrian C. Newton (Project manager) 

Rachel Cool< 

Neil Cox 

Victoria Gaillard 

Edmund Green 

Janina Jakubowska 

Thomas Kaissl 

Valerie Kapos 

Charlotte Lusty 

Anna Morton 

Mark Spalding 

Christoph Zockier 







UNEP WCMC 



The UNEP World Conservation Monitoring 
Centre is the biodiversity information and 
assessment arm of the United Nations 
Environment Programme, the worlds 
foremost intergovernmental environmental 
organization. UNEP-WCMC aims to help 
decision-makers recognize the value of 
biodiversity to people everywhere, and to 
apply this knowledge to all that they do. The 
Centre's challenge is to transform complex 
data into policy-relevant information, to build 
tools and systems for analysis and 
integration, and to support the needs of 
nations and the international community as 
they engage in joint programs of action. 



Production of maps 

Simon Blyth 
with the assistance of 
Igor Lysenko 
Corinna Ravilious 
Jonathan Rhind 

Layout 

Yves Messer 



A Benson production 

27 Devonshire Road 
Cambridge CBl 2BH, UK 



Color separations 

Swaingrove 

Printed in the UK 



Acknowledgments 



First and foremost we would [ike to express our deepest ttianks to the Aventis Foundation, 
without whose generous funding the research and production work for this book could not 
have been undertaken. Preparation of the book was also generously supported by the 
Department of Environment, Food and Rural Affairs IDEFRAI of the UK Government. The 
Owen Family Trust is also acknowledged for financial support to the first edition of this text. 

We also acknowledge with thanks the generous assistance extended by the following, listed 
approximately in the same sequence as the chapters in which their material appears: 

Christopher Field and George tvlerchant. Department of Global Ecology, Carnegie Institution of 
Washington for use of data from a global model of net primary production. 

Robert Lesslie of the Department of Geography, Australian National University, Canberra, for 
allowing us to use data resulting from his global wilderness analysis. 

BirdLife International, of Cambridge, UK, for allowing use of spatial data on endemic bird 
areas and on threatened bird species. 

Gene Carl Feldman, Oceanographer at NASA/Goddard Space Flight Center, Greenbelt, 
Maryland, for approving use of material from the SeaWiFS Project of NASA/Goddard Space 
Flight Center and ORBIMAGE. 

The University of Maryland Global Land Cover Facility, for facilitating use of land cover data. 

Professor Wilhelm Barthlott of the Botanisches Institut und Botanischer Garten, Rheinischen 
Fnedrich-Wilhelms-Universitat, Bonn, for kindly allowing use of a map showing contours of 
global plant species diversity 

Jonathan Loh, responsible for the WWF Living Planet Report, for kindly approving use of 
global trend indices from the Living Planet Report 2000. 

John E.N. Veron, Chief Scientist at the Australian institute of Marine Sciences, Townsville, 
Queensland for allowing use of coral generic diversity data. 

Several biologists associated with IUCN/S5C specialist groups on fishes, mollusks and inland 
water Crustacea, for providing data and expertise on important areas for freshwater 
biodiversity collated in an earlier publication: Gerald R. Alien (Western Australian Museum); 
the late Denton Belk [TexasI; Philippe Bouchet ILaboratoire de Biologie des invertebres 
marins et matacologie. Museum National d'Histoire Naturelle, Paris); Keith Crandall 
(Department of Zoology, Brigham Young University); Neil Cumberlidge (Department of 



Biology, Northern Michigan University); Olivier Gargominy iLaboratoire de Biologie des 
invertebres marins et nnalacologie, Museum National d'Histoire Nalurelle, Pans); Maurice 
Kottelat (Cornol, Switzerland); Sven 0. Kullander IDepartment of Vertebrate Zoology, Swedish 
Museum ot Natural History, Stockholm); Christian Leveque (ORSTOM. Pans); R. von Sternberg 
(Center for Intelligent Systems, State University of New York at Binghamton); Guy Teugels 
ILaboratoire Ichthyologie, Musee Royal de I'Afnque Centrale. Tervuren). 

Ben ten Brink, Jan Bakkes and Jaap van Woerden for facilitating use of material illustrating 
work on scenarios earned out at the Rijksinstituut voor Volksgezondheid en Milieu IRIVM), 
BiUhoven, the Netherlands. 

Christian Nellemann (Norwegian Institute for Nature Research), Hugo Ahlenius (UNEP GRID- 
Arendal) and the Secretariat of GLOBIO (Global methodology for mapping human impacts on 
the biosphere), for material applying this approach to scenario development. 



Photographs 

Pages: 6, L. Olesen/UNEP/Still Pictures; 7, L.L. Hock/UNEP/Topham; 

U, B. Groombndge; 15. M. Wakabayashi/UNEP/Topham; 37. Giotto Castelli; 

38, M. Fnedlander/UNEP/Topham; 39. UNEP/Topham; 72. G. Bluhm/UNEP/Topham; 

73. K. Kaznaki/UNEP/Topham; 86. M. Schneider/UNEP/Topham; 

93, M.R. Andrianavalona/UNEP/Still Pictures; 96, H. Mundell/UNEP/Topham; 

98, R. Fana/UNEP/Topham; 99. UNEP/Topham; 103 J. Nuab/UNEP/Topham; 

110 top. R. delRosanon/UNEP/Topham;110 bottom. Mazinsky/UNEP/Topham; 

118. E. Green; U3 top, E. Green; bottom. M. Spalding; U4, M. Garcia Blanco/UNEP/Topham; 

151. E. Green; 152. D. Nayak/UNEP/StiU Pictures; 15i. E. Green; 

155. D. Seifert/UNEP/StiU Pictures; 165, UNEP/Topham; 169. S.W. Mmg/UNEP/Stilt Pictures; 

174, F. Colombini/UNEP/Topham; 178. C.K. Au/UNEP/Still Pictures; 

185. K. Lohua/UNEP/Still Pictures; 194. C. Petersen/UNEP/Topham; 

212. P Garside/UNEP/Topham; 216, C. Senanunsakul/UNEP/Still Pictures 



Contents 



Foreword xi 

Preface xii 

Introduction 1 

Chapter 1: The biosphere 3 

Maps 

1.1 Physical geography of the Earth 4 

1.2 Primary production in the 
biosphere 8 

Figure 

1.1 Hypsographic curve 5 

Tables 

1.1 Global annual net primary 
production 10 

1.2 Estimated global carbon 

budget and biomass totals 11 

Chapter 2: The diversity of organisms 13 

Figure 

2.1 The phylogenetic tree 19 

Tables 

2.1 Estimated numbers of 
described species, and 

possible global total 18 

2.2 Key features of the major 

groups of living organisms 20 

Boxes 

2.1 New species discoveries 16 

2.2 Improving taxonomic 
knowledge and capacity 17 



Chapter 3: Biodiversity through time 

Figures 

3.1 The four eons of the 
geological timescale 

3.2 Periods and eras of the 
Phanerozoic 

3.3 Animal family diversity 
through time 

3.4 Plant diversity through time 



23 



3.5 Freguency of percent extinction 
per million year period 29 

3.6 Number of family extinctions 
per geological interval through 

the Phanerozoic 29 

Table 
3.1 The principal mass extinctions 

in the Phanerozoic fossil 

record 30 



Chapter 4: Humans, food and 






biodiversity 


33 


Maps 






4,1 


Early human dispersal 


34 


4.2 


Livestock breeds: numbers 






and status 


42 


4.3 


FAO world diet classes 


48 


4.4 


Human population density 


52 


4,5 


Terrestrial wilderness 


54 


4.6 


Vertebrate extinctions since 






AD1500 


56 


4.7 


Threatened mammal species 


58 


4.8 


Critically endangered 






mammals and birds 


62 


4.9 


Threatened bird species density 


64 


Figures 




4.1 


Human population 


47 


4.2 


Vertebrate extinctions by 






period since ADl 500 


59 


Tables 





4.1 



24 


4.2 




4.3 


26 






4.4 


27 




28 


4.5 



Top ten food commodities, 
ranked by percentage 
contribution to global food 
supply 41 

World diet classes 44 

Examples of diversity in 
agricultural systems 45 

Number of individuals and 
biomass, selected organisms 46 
Land converted to cropland 49 



A. 6 Estimated large herbivore 
numbers and biomass in 
Mesolithic and modern Britain 

4.7 Late Pleistocene extinct and 
living genera of large animals 

4.8 Numbers of extinct animal 
species according to lUCN 

4.9 Island diversity at risk: birds 

4.10 Threatened species 

4.11 Number of threatened animal 
species in major biomes 

Boxes 

4.1 Loss of diversity in agricultural 
genetic resources 

4.2 'Lazarus species' 

Chapter 5: Terrestrial biodiversity 

Maps 

5.1 Photosynthetic activity on land 

5.2 Global land cover 

5.3 Diversity of vascular plant 
species 

5.4 Biodiversity at country level 

5.5 Flowering plant family density 

5.6 Terrestrial vertebrate family 
density 1 

5.7 Current forest distribution 1 

5.8 Non-forest terrestrial 
ecosystems 1 

Figure 

5.1 A typical species-area plot 

Tables 

5.1 Global distribution of land area, 
by latitude bands 

5.2 Different definitions of forest 
cover 

5.3 Sample effects on forest area 
estimates of different forest 
definitions 

5.4 Global area of five mam forest 
types 

5.5 Important families and genera, 
and numbers of species, in four 
areas of temperate broadleaf 
deciduous forest 

5.6 Biomass and carbon storage in 
the world's major forest types 

5.7 Tree species richness in tropical 
moist forests 

5.8 Estimated annual change in 
forest cover 1990-2000 



5.9 Global protection of forests 
within protected areas in 

50 lUCN categories l-VI 97 

5.10 Estimated plant species 

51 richness in the five regions of 
Mediterranean-type climate 105 

57 Boxes 

60 5.1 Defining ecosystems 74 



61 


5.2 


Species and energy 


78 




5.3 


Fire in temperate and boreal 




63 




forest 


82 




5.4 


Temperate forest bird trends 


85 




5.5 


Grassland bird trends 


104 


40 








51 


Chapter 6: Marine biodiversity 

Maps 


117 


71 


6.1 


Coral reef hotspots 


126 




6 


2 


Shark family diversity 


128 


74 


6 


3 


Marine turtle diversity 


130 


76 


6 


4 


Mangrove diversity 


134 




6 


5 


Seagrass species diversity 


136 


88 


6 


6 


Coral diversity 


140 


90 


6 


7 


Marine fisheries catch and 




94 




discards 


US 




Figures 




00 


6.1 


Species contributing most to 




06 




global marine fisheries 


145 




6.2 


Marine fisheries landings by 




08 




major group 


146 




6.3 


Global trends in the state 




77 




of world stocks since 1 974 


U7 




6.4 


Trends in global fisheries 








catch since 1970 


147 


71 


6.5 


Marine aquaculture 








production 


150 


80 


6.6 


Marine population trends 


158 




Tables 






6.1 


Area and maximum depth of 




81 




the world's oceans and seas 


117 




6.2 


Relative areas of continental 




81 




shelves and open ocean 


118 




6.3 


Marine diversity by phylum 


122 




6.4 


Diversity of craniates in the 








sea by class 


123 


83 


6.5 


Diversity of fishes in the 








seas by order 


124 


84 


6.6 


Marine tetrapod diversity 


125 




6.7 


Regional distribution of 




87 




breeding in seabirds 


129 




6.8 


Diversity of mangroves 


132 


97 


6 


9 


Current mangrove cover 


133 



6.10 Diversity of stony corals in 
ttie order Scleractmia 

6.1 1 Coral reef area 

6.12 Taxonomic distribution and 
status of tfireatened marine 
animals 

Boxes 

6.1 Life in sediments 

6.2 Marine introductions 



7.2 



7.3 



7.4 



7.5 



7.2 

7.3 
lA 



138 
139 



156 

119 
153 



Chapter 7: Inland water biodiversity 163 

Maps 

7.1 Freshwater fisfi family 

diversity 170 

Major areas of diversity of 

inland water fish 176 

Major areas of diversity of 

inland water mollusks 180 

Major areas of diversity of 

selected inland water 

crustacean groups 182 

Priority river basins 190 

Figures 

7.1 Reported global inland 
fisheries production 
Inland water fish 
introductions 1 

Freshwater population trends 1 
River basin richness and 
vulnerability 1 

7.5 Changes in condition of a 
sample of freshwater lakes 
between 1950s and 198Ds 1 

Tables 

7.1 Components of the hydrosphere 1 

7.2 



179 



Physical and biodiversity features 
of major long-lived lakes 

7.3 Partial list of global hotspots 
of freshwater biodiversity 

7.4 Insects of inland waters 

7.5 Fish diversity in inland 
waters, by order 

7.6 Tetrapod diversity in inland 
waters 

7.7 Major inland fishery countries 

7.8 Numbers of threatened 
freshwater fishes in selected 
countries 

7.9 Taxonomic distribution and 
status of threatened inland 
water vertebrates 



91 

63 

66 

68 
72 

173 

175 
179 

187 



7.10 Thirty high-priority river basins 190 
Boxes 

7.1 Saline and soda lakes 164 

7.2 Wetland loss in Asian drylands 184 

Chapter 8: Global biodiversity: 

responding to change 195 

Maps 

8.1 World protected areas 200 

8.2 Centers of plant diversity 202 

8.3 Major areas of amphibian 
diversity 204 

8.4 Endemic bird areas 206 

8.5 Marine protected areas 208 

8.6 International protected area 
agreements 210 

Figures 

8.1 Development of the global 
network of protected areas 198 

8.2 Possible future scenarios 
from GEO 3. evaluated with 

RIVM IMAGE 218 

8.3 Possible future scenarios, 
GLOBIO 220 

Table 

8.1 Major global conventions 

relevant to biodiversity 

maintenance 214 

Boxes 

8.1 Pioneering NGOs in 
biodiversity conservation 197 

8.2 The precautionary principle 199 

8.3 Systematic conservation 
planning 203 

8.4 Negotiating a multilateral treaty 213 

APPENDICES 

1 The phyla of living organisms 225 



2 Important food crops 

3 Domestic livestock 



244 
271 



4 Recent vertebrate extinctions 278 

5 Biodiversity at country level 295 



6 Important areas for 
freshwater biodiversity 



INDEX 



306 
329 



Foreword 



Klaus Topfer, Executive Director, United Nations Environment Programme 

It is a great pleasure for me to introduce tliis important new bool< from tlie UNEP World 
Conservation Monitoring Centre. Building on the analyses it carried out for tfie Eartii Summit 
in Rio de Janeiro In 1992 and for the new millennium just two years ago, UNEP-WCMC has 
once again updated and revised its important overview of life on Earth in time for a major 
global event. 

In tune with the message of the Johannesburg World Summit on Sustainable Development, 
this new atlas places humankind firmly in the context of the species and ecosystems 
upon which we all depend for our livelihoods. We are a part of biodiversity and as such 
we should treasure it, use It wisely and share its benefits In our own enlightened self-interest. 

I commend this bool< to all who seel< a greater appreciation of the Inter-dependency between 
our own future and that of global biodiversity. 



In closing I should like on behalf of UNEP to thank most warmly the Aventis Foundation and 
the UK Department for Environment, Food and Rural Affairs for their support In the 
preparation of the World Atlas of Biodiversity. 



Preface 



Mark Collins, Director, UNEP World Conservation Monitoring Centre 



The diversity of life is tfie defining feature of planet Earth. It is unique - as far as we know - in 
the infinity of the universe. For 11 000 years since agriculture began, humankind has 
increasingly appropriated the biological resources and natural productivity of lands and seas 
to support the expansion of civilizations and technologies. Everything that we have achieved 
has its origins in living animals, plants and the communities and ecosystems of which they are 
a part. But it is only in the past 30 years, since the United Nations Conference on the Human 
Environment in Stockholm in 1972, that we have begun to recognize the limits to natures gifts. 
We now know that our own success is placing strain on nature's ability to evolve, diversify, 
cleanse our air and water and provide us with the raw materials we need for food, fuel, fiber 
and health. 

Just ten years ago we began to take integrated and holistic action to ensure conservation 
and sustainable use ot biological resources. The UN Conference on Environment and 
Development lUNCEDI saw the signing of the Convention on Biological Diversity, the first 
global agreement on biodiversity that clearly positioned humankind as an integral part of the 
complex of life on Earth, rather than a special case somehow separate from nature and 
immune from its laws. The 'ecosystem approach' espoused by the Biodiversity Convention 
acknowledges that our relations with the rest of the living world are truly interactive, and that 
what we do to nature will in turn reflect on nature's ability to respond to our own needs. The 
Convention foresaw a careful balance in the management of the Earth's living wealth through 
conservation, sustainable use and equitable sharing of costs and benefits. 

There could be no better time to launch a fresh assessment of the living world. This World 
Atlas of Biodiversity is published to coincide with the World Summit on Sustainable 
Development in Johannesburg, Republic of South Africa. The focus of the meeting is once 
again on sustainable development, but this time the emphasis is clearly on poverty alleviation. 
The message is clear: harmonized economic, social and environmental development will be 
but a dream while so many of the world's people have no choices and no opportunities to take 
a planned approach to their lives. What is the relevance of this book in the context of the 
Johannesburg message? 

The reality is that this World Atlas of Biodiversity is of greater relevance now than at any 
time in the past. The world's living wealth remains the cornerstone of sustainable livelihoods 
and quality lifestyles in both the industrialized and developing worlds. Recognition of this fact 
is spreading, and the value of biodiversity in people's lives, socially, economically and 
environmentally, has never been more apparent than it is today. 

This Is not a textbook, it is a resource pack and a survival kit for the future. I hope that all 
who read it will find new insights into the significance of life on Earth to their own lives. 
And that they will take steps within their homes, communities and nations to utilize and 
enjoy living resources wisely, share' the benefits and hold the capital in trust for 
future generations. 



Introduction 



OBJECTIVES 

The past ten years have seen a remarkable growth in concern for wildlife and the 
environment, with an increased appreciation of the linl<s between the state of ecosystems 
and the state of humankind. Many analysts have concluded that achieving sustainable and 
equitable human development will require, among other measures, taking a more effective 
approach to managing human impacts on the biosphere. This was reinforced by the 1992 
United Nations Conference on Environment and Development Ithe Earth Summit), at which 
the Convention on Biological Diversity ICBD) was opened for signature. Many conservation 
and management Initiatives worldwide have arisen from efforts to meet the objectives 
framed by the CBD text. 

In principle, any kind of variation at any level of biological organization - encompassing 
genes, populations, species and communities - is biological diversity. The text of the CBD 
defines biological diversity as 'the variability among living organisms from all sources 
including, inter alia, terrestrial, marine and other aquatic ecosystems and the ecological 
complexes of which they are part; this Includes diversity within species, between species 
and of ecosystems'. In practice, the term is often contracted to biodiversity', and used to 
refer collectively to all such variation: In effect, as a convenient shorthand for the total 
complex of life In some given area, or on the Earth as a whole. 

In the present volume we aim to use the data now available to provide an overview of 
the current state of global biodiversity, using maps where helpful, and to ensure that 
this information is accessible to a wide readership. While biodiversity has many dimensions, 
attention is here focused on the diversity of living organisms and their populations, and on 
major aquatic and terrestrial ecosystem types. Far more space is given to the macro-scale 
organisms and landscape elements that may be subject to planning and management 
intervention than to microorganisms, despite the Immense metabolic diversity of the 
latter, and their pivotal role in driving biosphere cycles. 



STRUCTURE OF THIS BOOK 

The eight chapters fall informally Into four thematic sections. The first section opens with an 
outline of some fundamental aspects of material cycles and energy flow in the biosphere 
(Chapter 1). This Is followed by a synopsis of the diversity of living organisms (Chapter 2) and 
of change In this diversity through geological time (Chapter 3). The second section (Chapter M 
Is largely concerned with relationships between humankind and biodiversity, noting the 
Increasing human impact on the environment from early modern humans onward, the use of 
biodiversity in human nutrition, and reviewing trends in recent time, focusing on depletion and 
extinction of species. The third section alms to characterize communities and biodiversity 
trends in the three basic blome types: terrestrial, marine and inland waters (Chapters 5, 6 and 
7, respectively). Finally, Chapter 8 introduces some of the management and planning 
responses that have been implemented with a view to maintaining ecosystem health and 
putting human development on a sustainable foundation. 



The biosphere 3 



1 



The biosphere 



rrr— he BIOSPHERE IS THE THIN AND IRREGULAR ENVELOPE around and including the 
Earth's surface that contains all living organisms and the elements they exchange with 
the non-living environment. Water makes up about tw/o thirds of an average living cell, 
and organic molecules based on hydrogen, carbon, nitrogen and oxygen make up the remaining 
one third. These and other elements of living cells cycle repeatedly betw^een the soil, sediment, 
air and water of the environment and the transient substance of living organisms. 

The energy to maintain the structure of organisms enters the biosphere when sunlight is 
used by bacteria, algae and plants to produce organic molecules by photosynthesis, and all 
energy eventually leaves the biosphere again in the form of heat. Photosynthetic organisms 
themselves use a proportion of the organic material they synthesize; net primary production 
Is the amount of energy-rich material left to sustain all other life on Earth. 

Humans now appropriate a large proportion of global net primary production, and have 
caused planetary-scale perturbations In cycling of carbon, nitrogen and other elements. 



THE LIVING PLANET 

The defining characteristic of the planet Earth 
Is that it supports life, and has done so for at 
least 70 percent of its history (see Chapter 31. 
The position of the Earth relative to the sun, 
its size and composition appear to be the 
main factors that have allowed life to develop 
here. Most importantly, these factors have 
combined to ensure the permanent presence 
of a large amount of liquid water on the 
planets surface, and this is the fundamental 
prerequisite of life as we know it. 

The space occupied by living organisms 
and the part of the planet that supports them 
is called the biosphere'. The non-living 
biosphere comprises the hydrosphere (the 
waters), the soil and upper part of the 
lithosphere (the solid matter that forms the 
rocky crust of the Earth], and the lower part of 
the atmosphere (the thin layer of gas coating 
the planet's surface). These domains interact 
in ways critical to the operation of the 
biosphere, and are linked in particular by 
the properties of water as a solvent and 
medium that fosters the chemical reactions 
basic to life. 



While providing the conditions necessary 
for life, the structure and composition of the 
non-living parts of the biosphere have them- 
selves been profoundly affected through time 
by living organisms. Most clearly, and from 
the human viewpoint most importantly, the 
presence of significant quantities of free 
oxygen in the atmosphere is entirely the 
product of oxygen-releasing photosynthesis 
by cyanobacteria starting more than 2 000 
million years ago. The idea that living organ- 
isms do not merely influence conditions in the 
biosphere but in some way regulate them to 
maintain the conditions conducive to life has 
received considerable attention in recent 
years, chiefly in terms of the Gaia hypothesis' 
proposed in the 1970s. 

The extent of the biosphere 

At planetary scale, the biosphere can be 
pictured as a thin and irregular envelope 
around the Earth's surface, just a few kilo- 
meters deep on the globe's 6 371 -kilometer 
radius. Because most living organisms depend 
directly or indirectly on sunlight, the regions 
reached by sunlight form the core of the 



k WORLD ATLAS OF BIODIVERSITY 



Map 1.1 

Physical geography of 
the Earth 

The relative areas occupied 
by dry land and by water, 
and the general distribution 
of areas of extreme height 
or depth. 












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biosphere: i.e. the land surface, the top few 
millimeters of the soil, and the upper waters 
of lakes and the ocean through which sunlight 
can penetrate. 

The biosphere is not homogenous, be- 
cause actively metabolizing living organisms 
are sparse or absent where liquid water is 
absent, such as in the permanent ice at the 
poles and on the very highest mountain 
peaks, but abundant where conditions are 
favorable. Nor are its boundaries sharply 
defined, because bacterial spores and other 
dormant forms of life passively disperse 
virtually everywhere, from polar icecaps to 
several tens of kilometers above the surface 
of the Earth [approaching the upper limit of 
the stratosphere], and living microorganisms 
occur^ within rocks more than 3 kilometers 
deep in the lithosphere. 



The whole of the sea is theoretically 
capable of supporting active life and con- 
stitutes therefore the vast majority of the 
volume of the biosphere [Figure 1.1|. 
Depending on water clarity, the sunlit (photic) 
zone may reach just a few centimeters to a few 
hundred meters in depth, but the marine 
biosphere is extended into regions of total 
darkness, down to more than 10 000 meters in 
the ocean depths, by organisms that subsist 
on the rain of organic debris falling from the 
upper waters, in addition, there are animal 
communities on the sea floor based on 
microorganisms deriving their energy from 
hydrogen sulfide emitted from hydrothermal 
vents. Overall, however, the amount of living 
material in most of the sea - that part of the 
open ocean below the upper hundred or so 
meters - is relatively low. 



/: 



i 



/" 




The biosphere 5 




The atmosphere plays a vital role in 
the biosphere, not only in providing a source 
of essential gases, but also in buffering 
conditions at ground level, by regulating 
temperature and providing a shield against 
excessive ultraviolet radiation. Many organ- 
isms, from microscopic bacteria to bats and 
birds, spend part of their lives suspended in 
the atmosphere; however, no organism is 
known that passes its complete life cycle in 
the air, and living biomass per unit volume 
above the Earth's solid or liquid surface is 
extremely low. 

Photosynthesis and the biosphere 

Life on Earth is based essentially on the 
chemistry of water and carbon. Indeed, in 
biochemical terms, living organisms are 
simply elaborate systems of organic macro- 



molecules dispersed in an aqueous medium. 
The average cell is about 70 percent water by 
weight; the remainder consists very largely of 
carbon-containing (organic! compounds com- 
posed mainly of the four elements hydrogen, 
carbon, nitrogen and oxygen. These com- 

8 849 m 



Figure 1.1 
Hypsographic curve 

The horizontal baseline in 
this figure represents the 
Earth's total surface area 
of 510 million km'. 
The figure shows that 71% 
of this surface is covered by 
marine waters and 29% is 
dry land. It also shows the 
mean land elevation and 
mean ocean depth, and the 
amount of Earth's surface, 
in percentage terms, 
standing at any given 
elevation or depth. 



: 


average elevation 840 m 




^"■"■^^^ average depth 3 800 m 


29% 


71% 



6 ~ 



10 

11 035 m 



i WORLD ATLAS OF BIODIVERSITY 



Energy from the sun 
drives photosynthesis, 
responsible for the vast 
majority of organic 
production. 



pounds include four major types of large 
organic molecule - proteins, carbohydrates, 
lipids and nucleic acids - and about 100 
different small organic molecules. A number 
of other elements are required in much 
smaller, though still vital, quantities. These 
include phosphorus, sulfur, iron and 
magnesium. All these elements cycle through 
the biosphere in a variety of forms, both 
organic and inorganic, following complex and 
interlinked pathways many of which are yet to 
be fully elucidated. Except for some micro- 
organisms that use energy derived from 
inorganic chemicals, the engine that drives 
the organic part of this turnover is photo- 
synthesis - the capture by living tissues of 
energy from the sun. 

Photosynthesis essentially involves the use 
of energy from sunlight to reduce carbon 
dioxide ICO2I with a source of electrons 
(almost invariably hydrogen) to produce 
carbohydrates, water IH2OI and, generally, a 
by-product from the hydrogen donor. In some 
bacteria the hydrogen donor is hydrogen gas, 
in others it is hydrogen sulfide; but, in 
cyanobacteria, algae and plants, water is the 
hydrogen donor and gaseous elemental 
oxygen IO2I is the by-product. This is over- 
whelmingly the predominant and most impor- 
tant form of photosynthesis on the planet, and 
is described by the following equation: 

2nH20 + nC02 + light -^ nHjO + nCH20 + 2nO 

The initial products of photosynthesis in 
plants are simple sugars such as glucose 




(041-1,2041. Larger carbohydrate molecules 
made from glucose include cellulose, the 
main component of plant cell walls and 
woody tissues, and starch, a key storage 
carbohydrate found in roots and tubers. 
Energy is needed to make the chemical bonds 
within these organic molecules, and energy is 
released when the bonds are broken down 
again. The controlled breakdown of these 
molecules within cells is the mechanism 
by which all cells obtain energy to do 
useful work. 

All organisms can break down sugars very 
directly without the need for oxygen. Many 
bacteria that live in aerobic conditions use only 
this method. Virtually all eukaryotes (see 
Chapter 2) have evolved a more complex 
additional pathway that requires oxygen but 
yields much more energy. This latter pathway 
- aerobic respiration - essentially reverses the 
basic photosynthetic reaction shown above. 

The major cycling process of the biosphere, 
therefore, consists of the photosynthetic fixing 
of carbon dioxide with water to produce 
organic compounds, in which energy is stored, 
and oxygen; this is followed by respiration of 
these compounds, in which the stored energy 
is released and carbon dioxide and water are 
produced. Photosynthesis therefore is not only 
responsible for the vast majority of organic 
production, but also for the maintenance of 
free oxygen in the atmosphere, without which 
aerobic organisms (the great majority of 
eukaryotic organisms, including humans) 
could not survive. 

Although photosynthesis is the primary 
engine of the biosphere, in the sense that it 
injects energy into the system and creates 
basic organic molecules, production of the 
full range of organic molecules on which life 
depends requires additional elements. Of the 
four key elements, nitrogen is often the one 
in limited supply, but it is an essential com- 
ponent of nucleic acids and proteins. 
Although the atmosphere consists of 79 
percent nitrogen, this inert gaseous form of 
the element cannot be used by plants or most 
other organisms until combined (fixed) with 
other elements. In the biosphere, atmos- 
pheric nitrogen is fixed by a range of bacteria, 
including cyanobacteria, some free-living soil 



The biosphere 7 



bacteria, and most importantly by specialized 
bacteria that live symbiotically in the root 
nodules of leguminous plants Ipeas, beans, 
etc.) Some nitrogen is also fixed by lightning 
in electric storms and, in the modern world, 
industrially in the production of fertilizer. 
Fixed nitrogen is made available to plant roots 
through association with fungi ImychorrhizasI 
and as nitrogen-fixing organisms decay. 
From plant roots, it is transported to 
metabolizing plant cells. On death, these 
steps are reversed, and the fixed nitrogen 
may be immediately recycled or revert to 
elemental nitrogen. 

PRODUCTIVITY AND THE CARBON CYCLE 

About half of the solar energy reaching the 
upper atmosphere of the Earth is immediately 
reflected. Most of the remainder interacts with 
the atmosphere, ocean or land, where it 
evaporates water and heats air, so driving 
atmospheric and ocean circulation. Much less 
than 1 percent of the incoming energy is 
intercepted and absorbed by photosynthetic 
organisms. On land these photosynthesizers 
are overwhelmingly green plants, although 
cyanobacteria and algae are also present, the 
latter particularly in the symbiotic associations 
with fungi known as lichens. In aquatic 
habitats, particularly the sea, virtually all 
photosynthesis is carried out by cyanobacteria 
and algae, although green plants are also 
present in shallow coastal and inland waters. 

Photosynthesizers fix carbon and therefore 
accumulate organic mass or biomass loften 
measured in dry form - that is, the once-living 
tissues of an organism with the water 
extracted]. These organisms are the primary 
producers. The amount of carbon fixed is 
referred to as gross primary production and 
is typically measured in grams (gl of carbon 
ICl per unit of space larea or volume) per unit 
of time. 

The photosynthetic producers also respire 
to meet their own energetic needs. Under 
some circumstances, respiration of photo- 
synthesizers over a given period may balance 
their carbon fixation, so that there is no net 
accumulation of organic carbon. More nor- 
mally, however, there is a surplus of fixation 
over respiration, so that organic matter is 



accumulated over time. This accumulation is 
referred to as net primary production iNPPl. 
The accumulated matter is available to the vast 
suite of organisms of all sizes, including 
humans, that cannot synthesize their own 
organic compounds from an inorganic base or 
harness energy from inorganic sources. Such 
organisms are referred to as heterotrophs, 
while photosynthesizers and the few l<inds of 
microorganisms that use other energy sources 
to synthesize organic compounds are referred 
to as autotrophs. 



Organic products pass 
through the food chain, 
through processes such 
as predation. 




Food webs 

An organic product produced by a photo- 
synthesizer may pass through a number of 
heterotrophs before finally being brol<en 
down again to its inorganic constituents. 
Conventionally this can be viewed as a food 
chain. At macroscopic level, a green plant 
may be eaten by a herbivore - a grasshopper, 
say - which is eaten by a lizard, which is 
itself eaten by a hawk, which dies and is 
disassembled and partially consumed by 
animal scavengers, with the remainder 
decomposed by bacteria and fungi. 

In reality, this is an enormous over- 
simplification. The plant will almost certainly 
have a complex network of symbiotic fungi 
associated with its roots, which make 
use of some of the gross production of the 
plant but which also provide it with some 
essential nutrients. The plant itself may shed 
leaves which are directly broken down by 
other fungi, protoctists such as slime molds. 



8 WORLD ATLAS OF BIODIVERSITY 



Map 1.2 

Primary production In the 

biosphere 

Global spatial variation in 
annual net primary 
production INPPI, in g C 
per m" per year, calculated 
from an integrated model 
of production based on 
satellite indices of 
absorbed solar radiation. 



Source: Map created (rom data 
supplied by Chrts Field and George 
Merchant, Department of Global 
Ecology, Carnegie Institution of 
Washington. See 

http://jaspenstanford.edu/chrisweb/flab/ 
flab, html, and Field et 3l. , 



g C per m' per year 




1 782 


-3 859 


1 107 


- 1 781 


881 - 


1 106 


671 - 


880 


487 - 


670 


3A1 - 


486 


230- 


340 


U3- 


229 


61 - 


142 


0-60 



and many forms of bacteria. The grass- 
hopper is likely to be parasitized by a host of 
smaller organisms, some of which are 
themselves in turn parasitized. It will also 
support a host of benign microorganisms in 
its intestine that are themselves constantly 
growing and reproducing. The lizard may die 
and decompose and the hawk may eat the 
grasshopper directly. The overall pattern of 
feeding relationships thus forms a web of 
immense complexity in any but the simplest 
ecosystems. 

Each organism in the food web respires, 
releasing energy which is eventually dissi- 
pated in the form of heat, carbon dioxide and 
water At each stage, therefore, some carbon 
is returned to the inorganic part of the 
carbon cycle. In addition, all living organisms 
produce waste products, some of which are 



r. 




incompletely metabolized organic compounds. 
Heterotrophic organisms are also not 
completely efficient in their appropriation of 
the organic material they consume, so that 
some proportion of this is excreted as waste 
product. These organic wastes are theor- 
etically available to other organisms in the 
food web. The assimilation efficiency of 
heterotrophic organisms may be anything 
from 20 percent (in the case of some 
terrestrial herbivores! to 90 percent (in 
the case of some carnivores), with the 
remainder excreted. 

Of the amount assimilated, a high 
proportion is expended as respiration, with 
the remainder available to add biomass, i.e. to 
enable the organism to grow and reproduce. 
The proportion available to add biomass is 
dependent on the organisms involved as well 



The biosphere 9 



'jr 



./ " ^ * 



X, 





as a range of other factors. It can be as low as 
10 percent or less and as high as 50 percent 
or more. This proportion is a measure of the 
net growth efficiency of the organism. 

For purposes of ecological analysis, 
particularly involving productivity estimates, 
the gross growth efficiency is the most 
commonly used measure. This is simply the 
product of the assimilation efficiency and 
the net growth efficiency of a particular 
heterotroph and is a measure of the pro- 
portion of food consumed by that organism 
that, after excretion and respiration, is 
ultimately available for its growth. As a 
very coarse generalization, a value of 
10 percent is widely used, although it is 
acknowledged that in terrestrial herbivores 
the figure is likely to be lower and in 
planktonic communities and terrestrial 



V , 



carnivores it is likely to be higher. Using the 
figure of 10 percent in the example above, for 
every kilo of plant matter eaten by the 
grasshopper, the latter would add 10 grams 
to its body weight. When the grasshopper was 
eaten by the lizard, this would add 1 gram to 
the lizards body weight, and when the lizard 
was eaten by the hawk, this would add 
0.1 grams to the hawks weight. This explains 
why, at the species level, so-called higher 
predators are rarer than herbivores and in any 
given area have a lower biomass, while the 
biomass of primary producers exceeds that of 
all heterotrophs combined. 

Measures of local and global productivity 

Primary productivity varies enormously, both 
spatially and temporally, at all scales. Most ob- 
viously, under natural conditions productivity 



10 WORLD ATLAS OF BIODIVERSITY 



Table 1.1 

Global annual net primary 

production 

Note: This represents one 
among several attempts to 
estimate global production; 
see text for further details. 
Ocean data averaged 1978- 
83, land 1982-90, units in 
petagrams II Pg = 10"g). 

Source: Adapted Irom Field et al . 



Biosphere units 


NPP 




lx10"gCl 


Ocean 


48.5 


Terrestrial 


56.4 


Tropical rainforest 


17.8 


Deciduous broadleaf forest 


1.5 


Broadleaf and needleleaf forest 


3.1 


Evergreen needleleaf forest 


3.1 


Deciduous needleleaf forest 


1.4 


Savannah 


16.8 


Perennial grassland 


2.4 


Broadleaf shrubs with bare soil 


1.0 


Tundra 


0.8 


Desert 


0.5 


Cultivation 


8.0 



effectively ceases every night. Seasonal 
variations In most parts of the world are also 
marked. Productivity is, however, difficult to 
measure, so that estimates at all scales are 
subject to considerable uncertainty 

On land one major source of uncertainty is 
below ground productivity: In natural eco- 
systems, less than 20 percent of plant produc- 
tion Is typically consumed by herbivores The 
remainder enters the soil system, either 
through the plant roots or as leaf litter 
Measuring this portion of terrestrial produc- 
tivity - probably over 80 percent of the total - 
Is particularly problematic. In the past there 
has been a marked tendency to under- 
estimate it. Similarly, It had long been 
assumed that the nutrient-poor waters of the 
open ocean were extremely unproductive, but 
It Is now known that large populations of 
extremely small photosyntheslzing unicellular 
organisms - the so-called picoplankton - form 
the basis of a surprisingly productive eco- 
system in these regions. 

One approach to estimating global pro- 
ductivity Is based on measurements in 
particular ecosystems and extrapolation from 
these using estimates of the global extent of 
those ecosystems. This suggests that net 
primary production on land is of the order of 
^5-65 X 10'= g C per year (that is, 45-65 
petagrams or thousand million metric tons), 
and at sea Is around 51 x 10'^g C per year This 



gives a global estimate of annual net primary 
production in the order of 100 x 10'^ g C. Gross 
primary production is estimated to be about 
twice this. 

However, global measures using a 
somewhat different technique, involving 
assessments of relative concentration of 
oxygen Isotopes, have Indicated that annual 
gross primary production on land may be 
greater than 180 x 10'* g C, while that in the 
sea Is around 140 X 10'* g C. This would give a 
global figure of over 320 x 10'* g C, implying 
global net primary production of more than 
160 X 10'* g C. This global figure Is 60 percent 
higher than those global estimates based 
on summation of Individual ecosystem 
measurements. 

Another approach' has used a com- 
prehensive set of satellite indices of 
photosynthetic activity In the ocean and on 
land, combined with a model of primary 
production, to generate a more integrated 
global estimate of NPP. There is considerable 
spatial and temporal variability, but on 
average annual NPP on land amounts 
to around 56 x 10'* g C. while that in the sea is 
around 48 x 10'* g C Isee Table 1.11. 

The carbon cycle and global blomass 
estimates 

Carbon fixation by photosynthesis forms one 
crucial step in the carbon cycle*. Once fixed, 
the carbon will remain for a greater or lesser 
period within living tissues, that Is, form part 
of the planets biomass. As all cells and 
individual organisms have a limited lifespan, 
eventually the carbon will rejoin the non- 
living carbon pool (see Table 1.2). It may, 
however, remain as organic carbon com- 
pounds for a far greater period than It 
remained part of the blomass; e.g. the woody 
tissues of Paleozoic forests were formed 
several hundred million years ago but remain, 
fossilized, as a source of coal and oil. 
Eventually all carbon will recycle through the 
inorganic pool, as carbon dioxide in the 
atmosphere, in the soil or dissolved In the sea, 
or as inorganic carbon compounds (car- 
bonates) In rocks or dissolved in the sea. 

The great majority of carbon at any one 
time lies within the lithosphere. around 



The biosphere n 



80 percent as carbonate and the remainder as 
organic carbon compounds. A large propor- 
tion of this carbon is effectively inaccessible to 
the biosphere in the short term but itself 
participates in the overall carbon cycle, 
mainly through tectonic activity As seafloor 
crust is gradually consumed along subduction 
plate margins, carbon sediments are taken 
into the Earth's mantle and later released as 
carbon dioxide by volcanic and hydrothermal 
activity. Although the loss of carbon to the 
mantle is extremely slow, without volcanic 
activity tectonic processes would eventually 
exhaust the available carbon pool. 

Most of the carbon incorporated in living 
organisms is associated with green plants, 
and almost all of this is in the form of 
cellulose-rich woody tissues. The total 
terrestrial animal biomass appears to be 
Insignificant in comparison, probably more 
than two orders of magnitude less. The 
cyanobacteria and algae that are the primary 
producers In the ocean are estimated' to 
amount to only 0.2 percent of the biomass of 
all primary producers globally, although they 
generate in the region of half the global NPP 
Ithey cycle organic material much more 
rapidly than land plants, which also sequester 
large amounts In woody tissues). 

The role of diversity In the biosphere 

The biota play the pivotal rile In the major 
blogeochemlcal cycles, with different groups 
of organisms (e.g. nitrogen-fixing bacteria 
and photosyntheslzing plants] mediating 
different processes. Simplistically, therefore, 
at least some biological diversity Is necessary 
to maintain the biosphere as It currently 
operates, and microbial diversity is partic- 
ularly fundamental. However, just how much 
diversity Is needed, and how much redun- 
dancy. If any, is built Into the system, remains 
unclear. Indeed, the relationship between 
biological diversity and a whole suite of 
ecological measures including stability, 
resilience and productivity remains incom- 
pletely understood', although there Is an 
increasing volume of theoretical and ex- 
perimental work" indicating that diversity 
may play an Important role In long-term 
ecosystem functioning. 



Human Influence on the biosphere 

There are believed to be more than 6 billion 
humans on the planet at present. A significant 
proportion of global net primary production is 
diverted to support this population. Using a 
relatively conservative definition of appro- 
priation that takes account of the global 
agricultural and natural production used by 
humans', that proportion has recently been 
estimated at around one third of the 
terrestrial global total'". This result is very 
similar to that obtained in an earlier study", 
but because of uncertainties in almost alt the 
figures on which it and the earlier estimate 
were based, the margin for error is very high. 
Less conservative definitions also attempt to 
take into account the loss of overall net 
primary production that may result from 
human actions (e.g. through severe land 
degradation or accumulation of waste] but 
which IS not directly used by humans. This Is 
done by estimating the net primary pro- 
duction of what Is thought would have been 
the prevailing biomes in the absence of 
humans. A detailed analysis In one European 
country using this approach" " suggested that 
around 50 percent of NPP there was 
appropriated by humans. 

Human efforts to appropriate the pro- 
ducts of photosynthesis and other actions 

Total carbon content on Earth 

Amount buried in sedimentary rocl<S: organic 
carbonate 

Active carbon pool near surface; 
of which 

Dissolved inorganic carbon in sea 

Atmospheric CO2 

Organic carbon in soil 

Biomass on land 

Biomass in the sea 



associated with the development of complex 
societies have had enormous impacts on 
natural biomes and blogeochemlcal cycles. 
Over large areas of the Earth's surface, 
humans have replaced complex and species- 
rich natural habitats with simplified modified 
habitats specialized for agricultural production. 



1.6xlO"g 
6.5xl0"g 

40 000xlO'=g 

38 000xl0"g 

750xl0"g 

ISOOxlO'^g 

560x10''g 

5-10x10'*g 



Table 1.2 

Estimated global carbon 

budget and biomass totals 

Note: Biomass figure on land 
refers to plants. 

Source; Adapted from Schlesinger . 



12 WORLD ATLAS OF BIODIVERSITY 



Clearance by fire, burning of fuelwood and 
charcoal, soil cultivation, and fossil fuel use all 
increase movement of organic carbon Into the 
atmosphere. Global cycling of nitrogen, 
phosphorus and sulfur has also been per- 
turbed. Application of industrially produced 
fertilizer has doubled the rate at which nitro- 
gen in fixed form enters the terrestrial cycle, 
and industrial processes have doubled 
movement of sulfur from the lithosphere into 
the atmosphere. Increasing levels of nitrogen 
and phosphorus lead to shifts in nutrient 
availability which can cause radical change in 
natural communities, and sulfur is a major 
contributor to acidification phenomena. 

That human activities may have profound 
local impacts on natural biota Is Indisputable. 
What IS now becoming clear is that these 



REFERENCES 



activities may also have planet-wide impacts, 
particularly on climate. Analysis of atmos- 
pheric samples trapped in polar ice cores 
indicates that present-day concentrations of 
atmospheric carbon dioxide and methane 
(CH^l are unprecedented in the past 420 000 
years". Although their absolute concentration 
In the atmosphere Is low (CO2 around 360 
and CH^ at 1,7 parts per million by volume) 
these two gases play an extremely important 
role in determining atmospheric temperature. 
It IS indisputable that the rise In these gases 
is a result of human activities, so clearly 
these activities are having some Impact on 
global climate. The extent of this impact, 
particularly when compared with natural 
climatic fluctuations, remains a subject of 
great controversy. 



1 Hutchinson, G.E. 1970. The biosphere. Scientific American 233(31: 45-53. 

2 Lenton, T.M. 1998. Gala and natural selection. Nature 394: 439-447. 

3 Parkes, R.J. 1999. Oiling the wheels of controversy. Nafure 401: 644. 

4 Field, C.B. e( at. 1 998. Primary production of the biosphere: Integrating terrestrial and 
oceanic components. Science 282: 237-240. 

5 Schleslnger, W.H. 1997. Biogeoctiemistry: An analysis of global cfiange. 2nd edition. 
Academic Press, San Diego and London. 

6 Schulze, E.D. and Mooney, H.A. (edsl 1993. Biodiversity and ecosystem function. Springer, 
Berlin. 

7 Grime, J. P. 1997. Biodiversity and ecosystem function: The debate deepens. Science 277: 
1260-1261. 

8 t«1cGrady-Steed, J., Harris, P.M. and t«1orin, P.J. 1997. Biodiversity regulates ecosystem 
predictability Nature390: 162-165. 

9 Wright, D.H. 1990. Human impacts on energy flow through natural ecosystems, and 
Implications for species endangerment. Ambio 19(4): 189-194. 

10 Rojstaczer, S., Sterling, S.M. and Moore, N.J. 2001. Human appropriation of 
photosynthetic products. Science 294: 2549-2552. 

11 Vitousel<, P.M. et al. 1986. Human appropriation of the products of photosynthesis. 
BioScienceZb: 368-373. 

12 Haberl, H. et al. 1999. Colonizing landscapes: Human appropriation of net primary 
production and Its Influence on standing crop and biomass turnover In Austria. IFF-Social 
Ecology Papers No. 57. Institute for Interdisciplinary Research of Austrian Universities, 
Vienna. Also see: http://www.cloc.org/conference/presentations/ln4/npp-abstract.htm 

13 Schulz, N. 1999. Effects of human land-use on the amount of biologically available 
blomass-energy throughout the landscape. An empirical case study about Austria. 
Conference paper available online in pdf at: 

http://vwvw.univle.ac.at/lffsocec/conference99/htmlfiles/blue.html (accessed January 
20021. 

14 Petit, J.R. et al. 1999, Climate and atmospheric history of the past 420,000 years from the 
Vostok Ice core, Antarctica. Nature399: 429-436. 



The diversity of organisms 13 



2 T 



he diversity of organisms 



g-^ YSTEMATICS AIMS TO DEFINE SPECIES and sort them into a hierarchy of named groups 
^^^ congruent with the branching pattern of evolution. There is no single operational 
definition of what a species is, and taxonomy at all levels is subject to change as a result 
of new methods and data, but species diversity of better known organisms can often be 
assessed with useful accuracy. Globally, about 1.75 million species have been described and 
named, but the total including undescribed species might be up to ten times greater. 

All species known are assigned on the basis of shared patterns of form and function to one 
of about 100 major groups (phyla). There are marked differences between phyla in overall 
morphology, physiology and mode of life. These differences imply the existence of major 
genetic diversity, and contribute directly to structural, trophic and other dimensions of 
diversity within ecosystems. 

The phyla of living organisms fall into three primary lineages: the true bacteria, the 
archeans and other organisms. The first two are prokaryotes, the remainder (protoctists, 
animals, fungi and plants! are eukaryotes. 



EVOLUTION AND SYSTEMATICS 

A basic principle in evolution is that just as new/ 
individuals arise from ancestral individuals, so 
new populations arise from existing popu- 
lations, and ultimately new species arise from 
existing species. The chief mechanism by 
which this occurs is believed to be reproductive 
isolation. For example, physiographic or 
climatic change may divide an existing single 
population into two or more separate popu- 
lations, or individuals may colonize a new and 
geographically separate habitat. The genetic 
makeup of these isolated populations will 
diverge, mainly through natural selection 
acting on them, but probably also through 
other mechanisms. This genetic divergence 
may be manifested in various ways, physically, 
physiologically and behaviorally. If the period of 
isolation continues for long enough, the 
populations will diverge enough that they can 
be regarded as separate species. Each one of 
these species may itself in turn give rise to 
other species in due course, although some 
will die out without giving rise to any progeny. 
The surviving descendant species may 



themselves give rise to new species and so on 
through the long march of evolutionary time. 

The result of this process is a branching 
tree-like structure - a phylogenetic tree - 
rooted in the distant past. In archeans and 
bacteria, but probably rarely in other groups, 
elements of two separate branches may 
combine into one, giving a reticulate rather 
than exclusively tree-lii<e pattern. If it is 
assumed that all life on Earth had a common 
origin in the distant past (see Chapter 3), then 
all existing organisms form the topmost 
extremities of a vast and unimaginably 
complex single phylogenetic tree. 

Systematics has two roles' '. The first is 
to name the immense variety of different 
sorts of organisms that exist. The second is to 
try to elucidate the relationships between all 
these different organisms, that is to develop 
hypotheses of where they are positioned in 
the phylogenetic tree. Systematics provides 
the basic framework for the whole of biology, 
and IS a fundamental discipline for bio- 
diversity studies. Taxonomy is the subset of 
systematics that deals in particular with the 



u WORLD ATLAS OF BIODIVERSITY 




Relationships between 
organisms have to be 
inferred from their 
genetic, morphological, 
biochemical or behavioral 
characteristics. 



definition, naming and classification of 
species and. in some cases, subspecitic 
populations. The traditional output consists of 
species descriptions or revisions, or lists of 
species in a given group (possibly with 
hypotheses of their evolutionary relation- 
ships), or checklists of all the species in some 
higher taxon in a site or region. 

Because the actual evolutionary events that 
generated the overall phylogenetic tree are lost 
in history, the relationships between organisms 
have to be inferred from the evidence to hand. 
The most important forms of evidence are the 
characters of organisms, both living and fossil. 
These characters may be genetic, morpho- 
logical, biochemical or behavioral. Methods to 
reconstruct phytogeny generally use two 
working assumptions: that species sharing a 
large number of characters are likely to be 
related, and that species sharing some 
uniquely complex and specialized feature are 
likely to be more closely related than species 
not possessing this feature. 

Groups and names 

In the current system for naming species 
(nomenclature), each has a two-part sci- 
entific name (binomial], based on Latin or 
latinized Greek, comprised of the genus name 
(e.g. Vipera] and specific epithet (e.g. berus]. 
The author of the specific epithet may be 
given after the binomial. By convention, both 
parts of the binomial are italicized when 
printed, and the author name is shown in 
parentheses if the species was originally put 
into a different genus. 

Similar species (e.g. the European adder 
Vipera berus and asp viper Vipera aspis\ are 
grouped together in the same genus (Wpera), 
similar genera in families (Viperidae), families 
in orders (Serpentes), orders in classes 
(Reptilia) and classes in phyla (Craniata or 
Vertebrata) up to the highest level, the 
kingdom. An organism can only be assigned to 
a single species, genus, family, etc., and the 
taxonomic system forms a hierarchy with each 
lower taxonomic level being nested entirely 
within each increasingly inclusive higher level. 
Although the traditional Linnaean hierarchy 
includes only the seven obligatory categories 
above, intermediate categories are some- 



times used, and a further more inclusive 
category - the domain. 

Groups such as mammals or snakes, that 
because of shared unique characters are 
considered to contain all the living descen- 
dants of a common ancestor, are called 
monophyletic groups. Groups with no shared 
unique characters but only unspecialized or 
non-unique characters in common, are 
termed paraphyletic groups. These typically 
are groups of related species left over after 
one or more clearly monophyletic lineages 
that evolved within the group have been 
recognized and named. Examples are fishes 
(the craniate vertebrates without the unique 
features of tetrapods], reptiles (the amniote 
vertebrates without the unique features of 
birds or mammals), and the entire kingdom 
Protoctista. Groups defined on characters 
that appear to have evolved more than once 
are called polyphyletic groups. Although the 
goal of most systematists is to recognize only 
monophyletic groups in order to be able to 
retrieve evolutionary relationships from a 
classification, many paraphyletic groups, 
such as the three named above, continue to 
be very widely used in practice. 

Given a classification congruent with 
phytogeny, the taxonomic hierarchy becomes 
a device to store information on hypotheses 
about evolutionary history. Because hypoth- 
eses about relationships are always subject 
to revision as new information becomes 
available, or existing data are reinterpreted, 
the taxonomy of species is not fixed. The fact 
that species names are liable to change can 
cause confusion, for example when, as is 
commonly the case, conservation legislation 
uses a name no longer current. 

Species concepts and diversity 
assessment 

Despite the importance of 'the species', 
there is no unequivocal and operational 
definition of what a species is or how species 
can be recognized' I There are several 
definitions, differing in mainly theoretical 
and often subtle ways, but much of the 
existing body of systematic knowledge has 
been built up around elements of 'the 
biological species concept'. This defines a 



The diversity of organisms 15 



species as a population of organisms tliat 
actually or potentially interbreed in nature, 
and tliat are reproductively isolated by 
morphological, behavioral or genetic means 
from other such groups. It is, however, 
applicable only to organisms where sexual 
reproduction is the norm. 

In most real cases, especially where all 
the systematist has to hand is a collection 
of preserved specimens, whether criteria 
concerning reproductive isolation are met 
or not cannot in fact be tested, but an 
experienced worker will come to hold some 
particular level of morphological or other 
difference as deserving of species status. 
Where there is good evidence from fieldwork 
and geographic data attached to specimens 
that two somewhat similar populations 
occur in the same locality (i.e. are sympatric) 
but maintain their differences, they may be 
presumed not to interbreed, and will be 
treated and named as species. 

Different taxonomists will often use 
different criteria for the same group of 
organisms, so that one specialist may regard 
a group of fundamentally similar populations 
as a single species, whereas another will treat 
each smaller distinct population as a separate 
species. In the latter case, often associated 
with the 'phylogenetic species concept', the 
assumption is that each distinct lineage once 
established on its own evolutionary course is 
de facto a separate species. Many populations 
that were formerly described at subspecies 
level (i.e. somewhat distinct, but regarded as 
part of a single polytypic species complex! 
have subsequently been elevated to full 
species in this way, particularly if geo- 
graphically isolated. However, subspecies 
have often been named on the basis of just a 
few superficial features, not representative of 
the overall pattern of variation within species, 
and the formal subspecies category is now 
much less used than in the past (particularly 
among vertebrates). 

Different taxonomic characters and 
criteria are used to classify species in 
different groups of organisms. Those used, 
for example, to define species of fungi are 
very different from those used to define 
species of bird, and their application 




demands taxonomists with relevant specia- 
lized knowledge. Some organisms are diffi- 
cult or logically impossible to accommodate 
in any species concept involving criteria that 
assume all individuals reproduce by out- 
breeding, that is by sexual reproduction with 
another individual. Many higher plants are 
self-fertile or make use of various forms of 
asexual reproduction; in the latter case all 
individuals in a lineage are identical clones 
(assuming a very low or non-existent 
mutation ratel. Among the prokaryotes, 
bacteria can readily receive extraneous genes 
through direct entry of genetic material from 
the fluid environment, or from viruses or 
other bacteria', and this horizontal' transfer 
is independent of reproduction. Different 
kinds of bacteria have traditionally been 
defined by the cytological or biochemical 
properties of colonies in culture but, more 



Different taxonomic 
characters and criteria are 
used to classify species in 
different groups of 
organisms. 



16 WORLD ATLAS OF BIODIVERSITY 



recently, comparison of DNA and RNA 
sequences from sample collections has been 
used to distinguish one lineage from another 
In either case, the biological species concept, 
developed with reference to sexually 
reproducing, outbreeding animals and plants, 
is not appropriate. Such factors mean that 'the 
species' cannot provide a standard unit in 
which to evaluate all biodiversity because it 
does not define a single level in the hierarchy 
and its significance is not equivalent across all 
groups of organisms; it can, however, serve 
this purpose for most plants and animals. 

The use of different species concepts by 
different systematists can make a very 
large difference to the number of species 
recognized in a group and to complications 
in nomenclature (these will both affect the 
outcome of biodiversity inventory, an 



Box 2.1 New species discoveries 

The discovery of entirely new species of mammals and birds is rare, and often involves small 
and obscure forms. Remarkably, two large mammals previously unknown to science were 
discovered in one small area, the Vu Quang Nature Reserve in Truong Son, Viet Nam (along 
with many new species in other groups!. The Vu Quang ox or soala Pseudoryx nghetinensis 
was described in 1993, followed a couple of years later by a giant muntjac deer 
Megamuntiacus vuquangensis from the same area. The world's smallest muntjac deer, the 
Truong Son muntjac Munt/acus truongsonensis, was recently found in another part of the 
same region in Viet Nam. The soala is of particular interest because it does not appear to fit 
neatly in any of the main bovid groups as currently recognized. It is now known also to occur 
in adjacent parts of Laos. However, claims that another new bovid species existed in 
Southeast Asia, described as Pseudonovibos spiralis, were premature because the 
distinctive horns described as new were later shown to be domestic cattle horns, apparently 
reshaped artificially 



important application of systematicsl. 
Usually, however, only a small or very 
small number of taxonomists is working on 
any one group of organisms at any time, so 
that while the species level taxonomy of 
organisms is in a continual state of flux, it 
is generally not subject to radical and 
wholesale change. Exceptions arise where 
new techniques, chiefly molecular infor- 
mation and DNA data in particular, are 
applied to previously neglected groups or 
ones that have not been revised for many 
years. Nevertheless, the key point appears 



to be that units corresponding more or less 
closely to the biologists' model of the 
species' do indeed exist in nature and, in 
animals in particular, they define themselves 
to an extent through their reproductive 
behavior It has thus been possible to reach 
some measure of consensus on species-level 
classification of well-studied groups of larger 
organisms such as terrestrial vertebrates, 
and to estimate and compare the number and 
kinds of species in different sites, areas or 
countries. 

NUMBERS OF LIVING SPECIES 

From a practical point of view it is more 
important to know how many species, and 
which ones, occur in some spatially restricted 
area, such as a protected area or a country, 
than in the world overall. However, proper 
evaluation of each local situation requires 
some knowledge of the wider context and. 
where the goal is maintenance of global 
biodiversity in the face of increased risk, it is 
clearly important to have a sound appreci- 
ation of the full baseline range of diversity. 
This requires both an estimate of the number 
of known valid species, and an estimate of the 
number of unknown species, neither of which 
is readily available. 

The number of known species can be 
estimated by collating data from systematists 
and the taxonomic literature. Although many 
species names are synonyms (i.e. different 
names inadvertently applied to the same 
species), this can be done with reasonable 
precision for more familiar and well-reviewed 
groups of species. Recent calculations of this 
kind suggest that around 1.75 million of the 
probably far larger number that exist have 
been discovered, collected and later named 
by systematists". 

Any estimate of how many undiscovered 
and hence undescribed species are likely to 
exist in any given group, and in the biosphere 
overall, involves substantial uncertainty''. In 
taxonomic groups where individuals are 
readily visible, popular or economically 
important, and subject to sustained system- 
atic attention, e.g. mammals and birds, the 
number of known species is certainly very 
close to the total number of species in the 



The diversity of organisms 17 



group that exist. On average around 25 new 
species of mammals and and five of birds 
have been described annually in recent years'. 
Changing systematic opinion on which 
populations should be regarded as separate 
species and which should not, rather than 
completely new discoveries, is the major 
source of change in the number of named 
species in such groups. 

The converse applies to groups whose 
individuals are small, difficult to collect, 
obscure and of no popular interest, e.g. many 
groups of invertebrate animals. Frequently 
there are so few systematists actively working 
on a group that the number of named species 
appears to be limited mainly by the rate 
at which collected specimens waiting on 
museum shelves can be studied and 
described, and changing opinion on which 
populations are separate species is insig- 
nificant. In some cases, where new sampling 
and collection methods have been used, 
unexpectedly large numbers of new species 
have been found (e.g. tropical forest canopy 
insects and marine sediment nematodes). If 
findings from such local work are extrapolated 
to global level the total number of species 
calculated to exist is many orders of mag- 
nitude greater than the number actually 
known. Some estimates suggest that most 
undescribed terrestrial forms are likely to be 
tropical forest beetles, but new molecular 
techniques are revealing unsuspected divers- 
ity among microorganisms'. 

Although the goal of systematics is to 
recognize and name species, and to maintain 
an ordered body of information on names and 
associated biological data, there is no master 
catalog of all known species. Developing such 
a resource has only become feasible with 
advances in information technology during 
the past ten years. However, while many 
systematic data, in the form of checklists and 
museum catalogs, are now available in digital 
form over the Internet, and more will become 
so, a harmonized catalog in this format of all 
known species remains a distant prospect. 

Recent estimates of the numbers of known 
and possibly existing species in the world 
biota are given in Table 2.1. These are mostly 
large numbers, and the fossil record suggests 



that overall diversity has been increasing for 
some 600 million years up to the very recent 
past, but the numbers themselves are of little 
significance except in a wider context. 
Currently, much concern is focused on 
species numbers in relation to anthropogenic 
environmental change. 



Box 2.2 Improving taxonomic knowledge and capacity 




The Convention on Biological Diversity decided in 1998 to establish a Global Taxonomic 
Initiative IGTII", in recognition of a major 'taxonomic impediment' to effective biodiversity 
management. The objective is to improve decision-making for conservation, sustainable 
use and benefit-sharing by increasing taxonomic knowdedge and the number of trained 
taxonomists and curators. Similarly, the Global Biodiversity Information Facility IGBIFj" 
aims to develop an interoperable net«/ork of biodiversity databases and information 
technology tools that will enable users to access the viforld's stores of biodiversity 
information. In the same field. Species 2000'- (a global network based in the UK and 
Japan], and the Integrated Taxonomic Information System IITISI" in North America have 
joined forces to create a unified Catalogue of Life, planned to cover all known species of 
living organisms. Basic reference data on 250 000 species had been collated by mid-2001, 
and the plan is to reach 500 000 by 2003. 



DIVERSITY AT HIGHER LEVELS 

Until van Leeuwenhoek observed micro- 
organisms through a primitive microscope in 
the late 17th century, humans had been 
aware only of organisms visible to the naked 
eye (macroscopic! and regarded all living 
things as either plants or animals. At the 
end of the 19th century, with improved 
cytological techniques and new views on 
evolution, a third kingdom of organisms 
(Protista) was recognized for bacteria and 
other unicellular organisms. Around this 
time it became widely accepted that the cell 
was the fundamental unit of organization of 
all living organisms. Subsequent work, in the 
mid-20th century, recognized a basic dis- 
tinction between two kinds of cellular 
organization - prokaryotic and eukaryotic. In 
prokaryotic organisms, the genetic material 
is free within the cell. In eukaryotes, the 
genetic material is linked to proteins and 
organized into chromosomes that are packed 
within a membrane-bounded cell nucleus. 
There are several other profound differences. 
In eukaryotes the enzymes needed to extract 
energy from organic molecules are organized 



18 WORLD ATLAS OF BIODIVERSITY 



Table 2.1 

Estimated numbers of 
described species, and 
possible global total 



into discrete membrane-bounded organelles 
Imitochondna) within the cell, and in the 
eukaryotes that photosynthesize (plants, 
some protoctistsi the pigments and enzymes 
needed to fix solar energy are also in 
discrete organelles Ichloroplasts) within the 
cell. On the basis of this, it became accepted 
that the major taxonomic divide in organisms 
was between the prokaryotes, containing 
only the bacteria, and eukaryotes, which 
included all other organisms, including 
many unicellular forms included in the 
original kingdom Protista. 

Systematics has traditionally generated 
information on similarity and hypotheses of 
relationship on the basis of characters 
restricted to particular sectors of the 
phylogenetic tree, but molecular sequencing 
has broadened the range of useful evidence. 



with the potential for radically dissimilar 
groups to be compared and their phylogeny 
estimated. Analysis of small subunit ribo- 
somal RNA ISSU rRNAl - a molecule that is 
universal, functionally constant (central to 
protein manufacture in all cells! and very 
highly conserved over time - has proved 
especially informative. This work has revealed 
that there are two kinds of prokaryote: the true 
bacteria and the archeans. The archeans were 
first known only by extremophiles', i.e. forms 
living under exceptional conditions of high 
temperature or salt concentration, but repre- 
sentatives are now known to be widespread 
alongside bacteria in less extreme habitats. At 
biochemical level, the bacteria and archeans 
are as different from each other as from 
eukaryotes, leading to the conclusion that all 
organisms can be assigned to three basic 



Domain Eukaryote 
kingdoms 

1 



Arcliaea 

Bacteria 
Eukarya 



Animalia 

Craniata (vertebrates!, total 

Mammals" 

Birds" 

Reptiles'* 

Amphibians" 

Fishes" 
Mandibulata (insects and myriapods! 
Chelicerata (arachnids, etc.] 
Molluscs 
Crustacea 
Nematoda 



No. of 


Estimated 


described 


total 


species 





175 



10 000 





1 320 000 


52 500 


55 000 


4 630 




9 750 




8 002 




i950 




25 000 




963 000 


8 000 000 


75 000 


750 000 


70 000 


200 000 


40 000 


150 000 


25 000 


400 000 





Fungi 


72 000 


1 500 000 






Plantae 


270 000 


320 000 






Protoctista 


80 000 


600 000 


TOTAL 




1 750 000 


U 000 000 



Notes: This table presents recent estimates of the 
number of species of living organisms in the high- 
level groups recognized, and in some selected 
groups v^ithin them. Vertebrate classes are 
distinguished because of the general interest in 
these groups. The described species column refers 
to species named by taxonomists. Most groups lack 
a formal list of species. All estimates are 
approximations. They are inevitably inaccurate 
because new species wiill have been described since 
publication of any checklist and more are 
continually being described, and other names turn 
out to be redundant synonyms. In general, the 
diversity of microorganisms, small-sized species, 
and those from habitats difficult to access, are likely 
to be seriously underestimated. Among Archaea and 
Bacteria the figures of 175 and 10 000 are very 
rough estimates of 'species' defined on features 
shov^n in culture"; there appears to be no sound 
estimate of the total amount of prokaryote diversity. 
The estimated total column includes provisional 
vKorking estimates of the number of described 
species plus the number of unknown species; the 
total figure is highly imprecise. Only a small 
selection of animal phyla is shown, but the figure for 
Animalia applies to all. Figures in the total row are 
for all species in all domains. 

Source: Data mainly from United Nations Environment 
Programme^ and Hammond^; vertebrates from individual 
sources indicated. 



The diversity of organisms 19 



BACTERIA 



ARCHAEA 



EUKARYA 



Animals 




forms of life, or domains: Archaea, Bacteria 
and Eukarya, rattier than two lprol<aryotes and 
eukaryotesl. The distance between lineages. 
In terms of amount of change in rRNA 
sequence, and their branching sequence In 
evolution, have been represented as a 
'universal phylogenetic tree'" ". 

Where the root of the tree Is located, and 
Its basal branching pattern, i.e. which two 
of the three domains are more closely 
related than either Is to the third, remains 
open to discussion"". It has been argued'" 
that this deep branching took place in very 
early evolution, possibly before modern cell 
types had been established, when genes 
were widely spread by horizontal transfer 
las persists to a lesser extent among 
extant prokaryotesl and in a sense, shared 
communally. 

Reticulate evolution would have been 
common, with distinct lineages of organisms 
emerging only when the evolving cell became 
sufficiently integrated that horizontal gene 
transfer was reduced and vertical transfer 
down closed ancestor-descendent series 
became the norm. It has also been per- 
suasively argued that the Eukarya arose 
through the permanent symbiotic fusion of a 
number of different prokaryotic organisms In 
one cell, with organelles such as mitoc- 
hondria and chloroplasts representing the 
vestiges of different lineages of formerly 
independent prokaryotes' '". 



PROTOCTISTS 



Within the Eukarya, the fungi, animals and 
plants form Insignificant clusters over- 
shadowed by the morphological and physio- 
logical diversity of the protoctists Iprotistsl. 
Protoctista is a name of convenience for the 
enormous and very diverse collection of all 
the small-to-mlcroscopic eukaryotes that 
lack the distinguishing features of fungi, 
animals or plants. Further changes of pers- 
pective are expected as research on protoctlst 
systematics continues. For example, a recent 
tendency^' Is to recognize the Chromista as 
another major group distinct from remaining 
eukaryotes. The chromists are aquatic 
species distinguished by structural and 
biochemical characters, and include organ- 
Isms with the largest linear dimensions 
known Ikelpl, as well as microscopic but 
ecologically important organisms (e.g. 
diatoms, downy mildews). 

The only known organisms that are not 
cells, or assemblages of cells, are viruses. They 
exist on the very boundary of most definitions of 
life. Consisting only of nucleic acids and protein, 
they are much smaller than the smallest 
bacteria, they can only replicate Inside other 
living cells, and they are totally inert outside 
other cells, when they can survive for years in a 
crystallized state. Each type of virus may be 
more closely related to the organism in which It 
grows than to other viruses'. They are not 
discussed elsewhere In this book. 

Some characters of the high level group- 



Figure 2.1 

The phylogenetic tree 

Note: This diagram 
represents in highly 
simplified form the distance 
between the three domains 
of organisms and the 
general branching pattern 
within them; because of 
conflicting interpretations 
the root and branching 
sequence of the three 
domains are not 
represented. 

Source: Adapted from Woese . 



WORLD ATLAS OF BIODIVERSITY 



Table 2.2 

Key features of the major 

groups of living organisms 

Note: The four groups within 
the Eukarya are commonly 
regarded as kingdoms, the 
highest formal category of 
the Linnaean system of 
taxonomy The Archaea and 
Bacteria have been treated as 
a single group, or as separate 
kingdoms, but a more recent 
tendency is to treat Archaea, 
Bacteria and Eukarya as 
three separate domains in 
recognition of the deep 
phyletic divergence between 
them, and fundamental 
differences in RNA 
composition. 

Source: Principally Margulis and 
Schwartz . also University of California 
and University of Arizona 



Arcfiaea 

Prokaryotic. Composition of the cell wall and of lipids in cell membranes differ from those in 
Bacteria. Distinctive SSU rRNA, more similar in some respects to Eukarya than to Bacteria. 
Reproduce asexually by cell splitting, or produce genetic recombinants without any fusion of cells by 
accepting genes from other bacteria, or from the fluid medium, or through viruses, independent of 
cell reproduction. Flourish in habitats with radical extremes of temperature or salinity that are 
unavailable to other organisms lapart from bacteria in some casesi, but also occur in other 
environments. 

Bacteria 

Prokaryotic. Reproduce asexually by cell splitting, or produce genetic recombinants without any fusion 
of cells by accepting genes from other bacteria, or from the fluid medium, or through viruses, 
independent of cell reproduction. Metabolically uniquely versatile; key mediators of major 
biogeochemical cycles. Permeate the entire biosphere, including other organisms, although dominant 
only in exceptional habitats. 



Eukarya 



Animalia 



Fungi 



Multicellular, mainly macroscopic, eukaryotes. Reproduce through fertilization cf an egg by a 
sperm, the fertilized egg Inow diploid, i.e. a duplicate set of chromosomes] is called a zygote 
and (except spongesl this forms a characteristic hollow multicelled blastula from which the 
embryo develops. All heterotrophic. 



Mainly multicellular, micro- to macroscopic eukaryotes. Fungi develop directly without an 
embryo stage from resistant non-motile haploid (one set of chromosomes] spores that can be 
produced by a single parent. Sexual reproduction also results in haploid spores. Most consist 
of a network of threadlike hyphae. Heterotrophs, vital to decomposition processes; form 
mycorrhizal symbioses with plants, facilitating exchange of soil nutrients. 

Plantae 

Multicellular macroscopic eukaryotes. The fertilized egg develops into a multicelled embryo 
different from blastula of animals. Alternate spore-producing generations and egg or sperm- 
producing generations. Virtually all are terrestrial photosynthetic autotrophs. 

Protoctista 

Mainly microorganisms. Possess the features of eukaryotes, but lack the characteristics 
of fungi, animals or plants. Extraordinary variation in life cycle and morphology Early 
evolution probably based on symbiotic relationships between different kinds of bacteria 
forming lineages of composite organisms resulting in the protoctist grade of 
organization. Include photosynthetic algae (formerly classed as plants] and heterotrophs 
(formerly called 'protozoa']. 

ings of organisms (domains and kingdoms] is presented in Appendix 1. The objective of 

are outlined in Table 2.2. A synopsis of this material is to provide a convenient 

information on eacfi of the 96 phyla recognized overview of global organismal diversity, in 

in the most recent comprehensive synthesis' terms of higher taxon diversity. 



The diversity of organisms 21 



REFERENCES 

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2 Vane-Wright, R.I. 1992. Systennatics and diversity. In: World Conservation Monitoring 
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pp. 7-12. Chapman and Hall, London. 

3 Mayr, E. and Ashlock, P.D. 1991 . Principles of systematic zoology. McGraw-Hill Inc., New York. 
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Science Ltd, Oxford. 

5 Vane-Wright, R.I. 1992. Species concepts. In: World Conservation Monitoring Centre. 
Groombridge, B. led.) Global biodiversity: Status of the Earth's living resources, pp. 13-16. 
Chapman and Hall, London. 

6 Margulis, L. and Schwartz, K.V. 1998. Five kingdoms. An illustrated guide to the phyla of 
life on earth. 3rd edition. W.H. Freeman and Company, New York. 

7 United Nations Environment Programme 1995, Heywood, V. (ed.l Global biodiversity 
assessment. Cambridge University Press, Cambridge. 

8 Hammond, P. 1992. Species inventory. In: World Conservation Monitoring Centre. 
Groombridge, 8. (ed.l Global biodiversity: Status of the Earth's living resources, pp. 17-39. 
Chapman and Hall, London. 

9 Williams, D.M. and Embley, T.M. 1996. Microbial diversity: Domains and kingdoms. 
Annu. Rev Ecol. Syst. 27: 569-595. 

10 GTI. http://www.biodiv.org/programmes/cross-cutting/taxonomy/ (accessed January 20021. 

1 1 GBIF. http://wviw.gbif.org/ (accessed January 20021. 

12 Species 2000. http://vww.sp2000.org/ (accessed January 2002). 

13 ITIS. http://www.itis.usda.gov/ (accessed January 2GG2I. 

14 Woese, C.R., Kandler, 0. and Wheelis, M.L. 1990. Towards a natural system of organisms: 
Proposal for the domains Archaea, Bacteria and Eukarya. Proceedings of the National 
Academy of Sciences 87: 4576-4579. 

15 Woese, C.R. 2000. Interpreting the universal phylogenetic tree. Proceedings of the 
National Academy of Sciences 97: 8392-8396. 

16 University of California, Berkeley, Museum of paleontology. 
http://www.ucmp.berkeley.edu/alllife/threedomains.html (accessed January 20021. 

17 University of Arizona, Tree of Life project. 
http://phylogeny.arizona.edu/tree/phylogeny.html (accessed January 2002]. 

18 Margulis, L. 1998. The symfa/of/c p(anef; /\ new (oofc af ei/o/uf/on. Weidenfeld and 
Nicolson, London. 

19 Philippe, H. and Forterre, P. 1999. The rooting of the universal tree of life is not reliable. 
Journal of Molecular Evolution 49: 509-523. 

20 Woese, C.R. 1998. The universal ancestor Proceedings of the National Academy of 
Sciences 9^3: 6854-6859. 

21 http://www.ucmp.berkeley.edu/chromista/chromi5ta.html (accessed January 20021. 

22 Woese, C.R. 1998. Default taxonomy. Proceedings of the National Academy of Sciences 
95: 11043-11046. 

23 Wilson, D.E. and Reeder, D.M. ledsl 1993. Mammal species of the world: A taxonomic and 
geographic reference. 2nd edition. Smithsonian Institution Press, Washington DC and 
London. Available (accessed January 2002] in searchable format at 
http://nmnhwww.si.edu/msw/ 

Covers existing or recently extinct species, as of book publication year. 

24 Clements, J.F. 2000. Birds of the world: A checklist. Pica Press, Sussex. Other lists are 
available online, e.g. a list based on Sibley, C.G. and Monroe, Jr, B.L. 1990. Distribution 
and taxonomy of birds of the world. Yale University Press, New Haven and London is 
available (accessed January 20021 at http://www.itc.nl/~deby/SM/SMorg/sm.html 



22 WORLD ATLAS OF BIODIVERSITY 



25 http;//www.embl-heidelberg.de/-uetz/LivingReptiles.html (accessed January 2002). 

26 Estimate is sum of species numbers for Anura 14 701 1, Caudata (505) and Gymnophiona 
(caeciliansl (1591 retrieved from ontine resource Amphibia Web at 
http://elib.cs.berkeley.edu/aw/lists/index.stitml (accessed January 20021. Also see 
Duellman, W.E. 1993. Amphibian species of the world: Additions and corrections. Special 
Publication No. 21, University of Kansas Museum of Natural History, Lawrence, Kansas, 
and database with later revisions available online in searchable format at 
http://research.amnh.org/herpetology/amphibia/ (accessed January 20021. 

27 Eschmeyer, W.N. et ai 1998. A catalog of the species of fe/ies. Vo[s 1-3. California 
Academy of Sciences, San Francisco. Available online in searchable format at 
http://www.calacademy.org/research/ichthyology/catalog/fishcatsearch.html (accessed 
January 20021. 



Biodiversity through time 23 



3 



Biodiversity through time 



Two FUNDAMENTAL PATTERNS CAN BE DISTINGUISHED in the fossil record. On one 
hand, new groups of organisms appear, diversify and generally persist for very long 
periods of time; on the other, most such groups and their Included species eventually 
cease to exist. Most analyses of the fossil record show an erratic rise In overall biodiversity, 
increasing through the Mesozolc and Cenozolc and reaching a peak around the end of the 
Tertiary. However, diversity has been greatly reduced during several periods of radical 
environmental change during each of which more than half the multicellular species then 
living became extinct. Such mass extinction phases have provided Important new 
opportunities for diversification in remaining lineages, and the spread of new communities. 
Evidently, the large number of species existing now on Earth Is the result of a modest net 
excess of originations over extinctions during the 3 800 million years of evolution of life. 



THE FOSSIL RECORD 

Knowledge of the history of diversity through 
geological time is based on analysis of the 
fossil record. When fossils, as the inert 
mineralized parts or casts or imprints of dead 
organisms, are interpreted in a biological 
context they provide the only direct evidence 
of the history of life on the planet. 

The fossils discovered and described by 
paleontologists represent more than a quarter 
of a million species, virtually all of them now 
extinct', but these are believed to make up only 
a very small fraction of all the species that have 
ever existed. For example, the fossil record of 
marine animals is far more comprehensive 
than that of terrestrial forms, but the marine 
sample is estimated to represent only about 
2 percent of all the marine animals that have 
lived'. The fossil record overall may represent 
as little as 1 percent, or less, of all the species 
that have existed'. Clearly, statements about 
broad patterns in the evolution of life, and the 
ascendancy or extinction of groups of organ- 
isms, thus rest on a very narrow base of 
tangible evidence. 

Macroscopic animals with hard skeletons 
that lived in shallow marine environments, 
where their remains could be buried by 



sediment, petrified, and later exposed in 
uplifted rock strata, are by far the most likely to 
be both preserved and found. The fossil record 
from the past 600 million years is thus 
dominated by mollusks, brachiopods and 
corals'. For many other kinds of organism, 
exceptional circumstances are required if a 
dead individual is to become preserved and 
found. With some exceptions, microscopic 
or soft-bodied organisms rarely leave discern- 
able traces of their existence, and larger 
organisms are usually decomposed, dis- 
assembled and never discovered. Terrestrial 
vertebrate fossils are often of individuals that 
must have been preserved by the smallest of 
chances, perhaps sudden burial during a 
natural disaster, or as a result of the body falling 
into a rock crevice out of reach of scavengers. 

All else being equal, the probability of 
preservation will rise greatly the more 
widespread and abundant the species is, and 
the longer it persists through time. Conversely, 
there is a very low probability that any individual 
of a numerically rare or restricted-range 
species with a short persistence time will die in 
circumstances conducive to fossilization and 
subsequently be found. Factors such as these 
mean that even the relatively better known 



24 WORLD ATLAS OF BIODIVERSITY 



Figure 3.1 

The four eons of the 

geological timescale 

Arrows indicate 
approximate age of oldest 
confirmed fossils of the 
groups named. 

Source: Adapted from Mdrgulis and 
Schwartz . 



groups are certain to be very incompletely 
l<nown. The plant and animal species now living 
include a substantial number of rare or local 
species and it is difficult to imagine many of 
them being represented in the fossil record of 
our time as recovered in the future^ 

Because the fossil record gives an 
incomplete and biased view of the past history 
of life, the reconstruction of that history has 
been the subject of great debate. It has been 
generally accepted that the record can give a 
reasonable insight into past diversity in terms of 
taxonomic richness", particularly at higher 
taxonomic levels. On the other hand, recent 
analysis of a new database of fossil occurrence 
has indicated that sampling effects (reflecting 
variation in the nature of the fossil record and 
the way it is reported! can make a very 
significant contribution to the shape of global 
diversity curves. Preliminary results of this 
analysis have suggested, for example, that the 
widely accepted post-Paleozoic increase in 
marine diversity Isee belowl may to some extent 
be an artifact of the analytic methods used^ 

PATTERNS OF DIVERSIFICATION 
The early history of life 
In its earliest history, there was no life on the 
Earth. Now, there are about 1.75 million 
different species of all kinds known, with 
perhaps many times that number still 
unknown. Self-evidently, life has both arisen 
and diversified. 

The planet Earth is between 4 500 million 
and 5 000 million years old. The oldest known 
rocks are about 4 000 million years IMyl in 
age, and mineral grains IzirconI of apparently 
greater age have been reported. Carbon 



chemofossils' suggest that life had evolved 
by 3 800 My ago' but periodic intense ocean 
heating caused by extraterrestrial impacts is 
suspected to have restricted early organisms 
to thermophilic non-photosynthetic forms, 
possibly associated with hydrothermal 
vents". Most of the principal biochemical 
pathways key to the modern biosphere were 
probably in place by 3 500 My ago'. The first 
tangible evidence of cellular organisms 
themselves in the fossil record consists of 
filaments and spheroids, believed to be the 
remains of prokaryote microorganisms, in 
rocks of about this age, including traces from 
an apparent Precambrian submarine thermal 
spring system of about 3 235 My in age'. 

Stromatolites [rock domes formed in 
shallow waters from multiple layers of sedi- 
ment and bacterial also first appear at around 
3 500 My ago and are the most abundant fossils 
so far known during the 3 000 My up to the start 
of the Phanerozoic eon, marked by the start of 
the Cambrian period around 5^5 My ago. For 
much of this time, bacteria-like organisms 
were the only known life forms; gross 
morphological diversity was very low and many 
kinds apparently persisted for hundreds of 
millions of years, some outwardly indis- 
tinguishable from existing forms. Probable 
microbial mats, similar to stromatolites but 
developed on an exposed soil surface, have 
been interpreted as the first evidence of 
terrestrial ecosystems around 2 600 My ago'". 

The next major step in the evolution of life 
was the development of eukaryotic organisms. 
Biochemicals characteristic of eukaryotes 
have been found in shales 2 700 My old", far 
earlier than the first fossils, reported from 



Eon 



Millions of 
years ago 



t 



HADEAN 



4 500 



origin 

of Earth 




4 000 3 900 



3 500 



3 000 



t t t 



t 



oldest 


'chemofossils' 


oldest 


Earth 


possible 


microfossils 


rocks 


evidence 
of life 


of bacteria 



evidence of 
eukaryotes 



Biodiversity through time 25 



shales of around 1 500 My age". These earliest 
eukaryote microtossils Include a number of 
'acritarchs' (widely interpreted as the cysts or 
resting stages of marine algael some of which 
show cytological features characteristic of 
protoctists (see previous chapter]' . Diversity in 
acritarchs, and the rate at which different 
forms replaced one another in the record, 
were low until around 1 000 My ago when 
both species number and species turnover 
increased markedly". 

Radiations around the early 
Phanerozoic boundary 

For many years it was assumed that animals 
originated in the Cambrian period at the base 
of the Phanerozoic Ithe Phanerozoic is the eon 
of time characterized by presence of animal 
fossils; it includes the Paleozoic, Mesozoic and 
Cenozoic). This is now known not to be the 
case, as a wide range of fossil animals, 
including recognizable arthropods and possibly 
echinoderms, is now known from about 1 00 My 
before the Cambrian. Most fossils from this 
time, however, appear completely unrelated to 
extant forms, and consist mainly of enigmatic 
frond- and disc-shaped soft-bodied animals: 
the so-called Ediacaran fauna. 

The lower Cambrian marks a dramatic 
change from this early fauna, with the sudden 
appearance in the fossil record (e.g. the 
Chengjiang and Burgess Shale faunas! of a 
wide range of animals, many with calcareous 
skeletons. It is generally accepted that this 
represents a genuine explosion of diversity that 
took place over only a few million years, and is 
not an artifact of the fossil record. The lower 
Cambrian thus represents the most important 



period of high-level diversification In the his- 
tory of animal life on Earth. These archaic 
Invertebrates had by the end of the Cambrian 
period, around 500 My ago, established all the 
basic body plans seen in extant animals, and 
many others besides. Each such basic lineage 
Is recognized taxonomically at phylum level, 
and the range of morphological diversity was 
higher at 500 My ago than at any time before 
or since. As many as 100 different animal 
phyla may have existed during the Cambrian" 
including every well-skeletalized animal phy- 
lum living today (except perhaps the Bryozoa). 
whereas in the latest synthesis all extant 
animals are placed in 37 phyla'^ 

Plants and animals began to extend into 
terrestrial habitats during the first half of the 
Paleozoic, with the first fossil material known 
from the late Silurian, around iOO My ago. At 
this point approximately 90 percent of the 
history of life to the present had already 
passed. Fossils suggest low diversity tor the 
next 100 My until the later Devonian period. 
No new animal phyla appeared with the 
colonization of land, millions of years after 
the initial Cambrian radiation of animal phyla. 

Diversity of marine animals in the Phanerozoic 

The overall pattern of diversity [assessed as 
numbers of families! appears to show a 
possible early peak around the start of 
Phanerozoic time, followed by a plateau of 
somewhat higher diversity extending through 
most of the Paleozoic era, and then, after the 
end-Permian mass extinction (see below] a 
steady increase in diversity over remaining 
geological time'. Some recent methods of 
analysis' giving special attention to sampling 




2 000 



PROTEROZOIC 



1 500 



t 



oldest 

protcctist 

fossils 



PHANEROZOIC 



4 



1 000 



545 500 

t ft 

oldest oldest oldest 

animal fungus plant 

fossils fossils fossils 



24 WORLD ATLAS OF BIODIVERSITY 



Figure 3.2 

Periods and eras of the 

Phanerozoic 

This IS an expanded version 
of the most recent segment 
of the geological timescale 
shown in Figure 3.1 . 
The Phanerozoic is the eon 
of time extending from the 
base of the Cambrian, 
some 545 million years ago, 
to the present, and to which 
the entire fossil record was 
formerly thought to be 
restricted. 



Source: Adapted from Margulis and 
Schwartz , Cambrian base date from 
International Subcommission on 
Cambrian Stratigrapfiy website . 



procedures suggest this broad view may be 
subject to significant revision. 

Although the number of phyla has 
decreased markedly since the Cambrian, 
diversity at all lower taxonomic levels appears 
to have either increased overall or, in a few 
cases, remained more or less level. The 
number of orders of marine animals present 
in the fossil record climbed steadily through 
the Cambrian and Ordovician, slowing 
towards the end of the Ordovician to a figure 
of between 125 and 1A0, which has been 
maintained throughout the Phanerozoic. 

The diversity of marine families repre- 
sented in the fossil record shows a similar 
pattern of increase through the Cambrian 
(possibly falling during the latter half of the 
period! and Ordovician, leveling off at around 
500, a figure which was maintained until the 
late Permian mass extinction. This extinction 
event resulted in the loss of around 200 
families, but diversity increased subsequently 
to the modern level of around 1 100 families, 
with a number of temporary reversals during 
minor extinction events. The trend in number 
of species in the fossil record is even more 
extreme. From the early Cambrian until the 
mid-Cretaceous, the number of marine 
species remained low; since then, that is 
during the past 100 million years, it appears to 
have increased dramatically, perhaps by a 
factor of 10. (See Figure 3.3.) 

Diversity of terrestrial organisms 
in the Phanerozoic 

Although low to moderate peaks and troughs 
are evident in the record, the overall pattern 
of family diversity in terrestrial organisms 
shows a continuing rise from the Silurian to 
the present, thus differing somewhat from 



the pattern shown by many analyses of 
marine animals. 

It is generally accepted that vascular 
terrestrial plants first arose in the Silurian, 
although some paleobotanists argue for a late 
Ordovician origin, a time when microfossils 
suggest that bryophyte-like (non-vascularl 
plants already existed". Diversity increased 
during the Silurian, and then more rapidly 
during the Devonian, owing to the first appear- 
ance of seed-bearing plants, leading to a peak 
of more than AO genera during the late 
Devonian. Diversity then declined slightly, but 
started to increase markedly during the 
Carboniferous, with 20 families and more than 
250 species in the mid-Carboniferous record of 
the northern hemisphere. Following this, 
diversity increased only slowly until the end of 
the Permian. There was a marked decrease in 
diversity at the end of the Permian, coinciding 
with or preceding the mass extinction of 
animal species. Subsequent increase in diver- 
sity was slow, reaching around 400 species in 
the early Cretaceous, but apparently more 
rapid from the mid-Cretaceous. 

This overall pattern masks important 
changes with time in the composition of the 
flora, most notably in the relative importance of 
the three main groups of vascular plant: the 
pteridophytes, gymnosperms and angio- 
sperms (see Figure 3.4|. The Silurian and early 
Devonian are marked by a radiation of primitive 
pteridophytes. During the Carboniferous, more 
advanced pteridophytes and gymnosperms 
developed and underwent extensive diversifi- 
cation. Following the late Permian extinction 
event, pteridophytes were largely replaced by 
gymnosperms (although ferns remained 
abundant! and these became the dominant 
group until the mid-Cretaceous. The dramatic 



Period Cambrian 



Era 



Ordovician 



Silurian 



PALEOZOIC 



Devonian 



Carboniferous 



Millions of 
545 years ago 490 



438 



408 



360 



Biodiversity through time 




600 



300 200 

Geological time I lO' years! 



K 

100 



increase in plant diversity since then is entirely 
due to the radiation of the angiosperms, fossils 
of which first appear in the lower Cretaceous, 
although an early to mid-Jurassic origin, 
around 1 70 My ago, has been argued". 

Colonization of land by animals has 
occurred many times; although the oldest 
body fossils of terrestrial animals date from 
the early Devonian, it is generally accepted 
that the primary period of land invasion by 
animals was the Silurian. 

The overwhelming number of described 
extant species of terrestrial animals are 
insects. Their fossil record is more extensive 
than might be expected, but had been little 
studied until recently. Data on insect diversity 
at family level have been collated, based on 
nearly 1 300 families'". This analysis shows a 
very slow increase in families from the first 
appearance of insects in the Devonian, a rise 



in numbers during the Carboniferous, and a 
steeply increasing rise throughout the 
Mesozoic to the Tertiary. The apparent 
explosion in insect diversity had previously 
been attributed to ecological opportunities 
provided by the expansion of flowering plants, 
but the insects are now known to have begun 
their diversification some 150 fvly before the 
flowering plants". 

The fossil record of vertebrates includes 
around 1 iOO families, with tetrapods (amph- 
ibians, reptiles, birds, mammals) somewhat 
outnumbering fishes. The bird record is much 
less substantial than that for other groups, 
probably because their light skeletons have 
been less frequently preserved. 

Terrestrial vertebrates first appear in the 
fossil record in the late Devonian. Diversity 
remained relatively low during the Paleozoic, 
with around 50 families, and may have 



Figure 3.3 

Animal family diversity 

through time 

The lines plotted represent 
the number of families in 
the fossil record. 

Notes: The blue line 
essentially represents marine 
invertebrate animals. 
Although a small number of 
vertebrate groups, notably 
fishes and a few tetrapod 
species, are included, these 
make up a tiny proportion of 
the total marine family 
diversity shown. The curve for 
fishes includes an increasing 
proportion of freshwater 
forms through the Cenozoic. 
Tetrapods are amphibians, 
reptiles, birds and mammals. 
The extent to which such 
results, based on sampling 
and interpretation of the 
fossil record, represent 
actual diversity has been 
subject to discussion. 

Source Marine animals, adapted Irom 
Sepkoski ; fishes and tetrapods, 
adapted after Benton : insects, adapted 
from Labandeira and Sepkoski . 



rboniferous Pernnlan Triassic 



Jurassic 



Cretaceous 



Tertiary Quaternary - 



^V PALE( 



PALEOZOIC 



MESOZOIC 



KI-IH,-U«.1INU 



286 



248 



213 



U4 



CENOZOIC 



65 



WORLD ATLAS OF BIODIVERSITY 



800 



700 



ui 600 



500 



400 



300 



200 



100 



i..„ Pteridophytes 
Gymnosperms 
^H Angiosperms 




Tr 



400 



300 



200 



K 

100 



Geological time 110° yearsi 



Figure 3.4 

Plant diversity through time 

Notes- Pteridophytes are 
ferns Filicinophyta and allies, 
gymnospernns are conifers 
Coniferophyta and allies, 
angiosperms are flowering 
plants Anthophyta. Note 
changing numerical 
dominance of each group 
over time. 

This generalized diagram is 
to illustrate changing 
abundance of major groups 
over time; it does not 
represent short-term 
extinction events. 

Source: Adapted from Kemp . after 
Niklas. 



declined during the early Mesozoic. From the 
mid-Cretaceous the number of families 
started to increase rapidly, reaching a recent 
peak of around 340. Diversity of genera follows 
this overall pattern in a more exaggerated 
form. It appears that periodic increase in the 
number of tetrapod families is mainly a result 
of lineages becoming adapted to modes of life 
not already followed by other organisms, i.e. 
by adopting new diets or new habitats". 

PATTERNS OF EXTINCTION 

If living species represent between 2 and 4 
percent of all species that have ever lived™, 
almost all species that have lived are extinct, 
and extinction can be presumed to be the 
ultimate fate of all species. 

Numerous estimates have been made of the 
lifespan of species in the fossil record; these 
range from 0.5 My to 13 My for groups as varied 
as mammals and microscopic protoctists. 
Analysis of 17 500 genera of extinct marine 
microorganisms, invertebrates and verte- 
brates, suggests an average lifespan of 4 My in 
these groups'. Given this average lifespan, at a 
very gross estimate, the mean extinction rate 
would be 2.5 species per year if there were 
around 10 million species in total. However, 
because of bias inherent in the fossil record, 
such lifespan estimates are likely to relate to 



widespread, abundant and geologically longer- 
lived species, in effect, the extinction-resistant 
forms; and so do not represent the biota as a 
whole-'. If most species survived for less than 
4 My, the overall extinction rate at any given 
time would have been correspondingly higher 

Major extinctions in animals 
In general the Precambrian fossil record is too 
incomplete to allow detailed analysis of extinc- 
tion rates. However, there is good evidence of a 
major loss of diversity during the Vendian 
period in latest Precambrian times, around 
550 My ago, when the entire Ediacaran fauna 
disappeared lalong with many acritarchsl. 
Another wave of extinction affected archaeo- 
cyathid sponges, mollusks and trilobites during 
the lower Cambrian, some 530 My ago. 

By far the most severe marine invertebrate 
mass extinction was in the late Permian 
1250 My ago). At that time, the number of 
families of marine animals recorded in the 
fossil record declined by 54 percent and 
the number of genera by 78-84 percent. 
Extrapolation from these figures indicates 
that species diversity may have dropped by as 
much as 95 percent. Other major extinctions 
in marine invertebrates occurred at the end of 
the Ordovician 1440 My ago) (see Figure 3.31, 
when around 22 percent of families were lost, 
and during the late Devonian and late Triassic 
121 percent and 20 percent, respectively!. 
Around 15 percent of marine families 
disappeared at the end of the Cretaceous. 

The vertebrate fossil record, especially for 
terrestrial tetrapods, is much less amenable to 
analysis of extinction rates than the inverte- 
brate record, chiefly because it is less complete 
and less diverse. However, studies indicate 
that fishes have been subject to at least eight 
important extinction events since their recorded 
origin in the Silurian, while tetrapods have 
experienced at least six such events since their 
appearance in the late Devonian. Some of these 
events coincide with each other and with those 
recorded for marine invertebrates; in particular, 
the five major mass extinction events outlined 
above are paralleled by losses in vertebrate 
diversity. The most significant is the late 
Permian event, which, in terms of percentage 
loss, IS the largest recorded extinction both for 



iiodiversity through time 29 



fishes (44 percent of families disappearing from 
the fossil record) and tetrapods (58 percent of 
families disappearing). The late Cretaceous 
event was more significant for tetrapods than 
for fishes, with at least 30 of the 80-90 
families then in the fossil record disappearing 
at this time. These families were, however, 
virtually confined to three major groups which 
suffered complete extirpation - the dinosaurs, 
plesiosaurs and pterosaurs. Most other 
vertebrates were almost completely unaffected. 

Major extinctions in vascular plants 

Fewer major extinction events have to date 
been distinguished in the plant fossil record 
than in the animal record. Periods of elevated 
plant extinction appear in some cases more 
protracted than animal extinction events and 
not usually coincident with them". However, 
there is good evidence for extensive reduction 
in woody vegetation at the end of the Permian, 
with widespread loss of peat forests in humid 
areas and of conifers in some semi-arid 
regions. A fourfold increase in atmospheric 
carbon dioxide around the Triassic-Jurassic 
boundary is correlated with a more than 
95 percent turnover in the megaflora (i.e. leaf 
fossils, etc., as opposed to pollen or spores]". 
The end-Cretaceous catastrophe appears to 
have had a major influence on the structure and 
composition of terrestrial vegetation and on the 
survival of species. Data from fossil leaves 
suggest that perhaps 75 percent of late 



0.20 



0.15 



0.05 




Figure 3.5 

Frequency of percent 
extinction per million year 
period 

See text for explanation. 
Source- Adapted from Raup', 



40 60 

% extinction 



Cretaceous species became extinct, although 
data from fossil pollens indicate a lower though 
still significant level of extinction. During the 
Tertiary there are two other periods of 
widespread heightened extinction rates, during 
the late Eocene and from the late Miocene to 
the Quaternary, although in the latter extinction 
of taxa at generic level and above appears to 
have been mainly regional rather than global. 

Mass extinctions 

The very many species extinctions represented 
in the fossil record are not distributed evenly 
through time, nor do they occur randomly In 
paleontology much attention has been devoted 
to mass extinction periods, during which some 
75-95 percent of species then living became 



Figure 3.6 
Number of family 
extinctions per geological 
interval through the 
Phanerozoic 

Note: 76 geological intervals 
represented, average 
duration around 7 million 
years. 

Source: Benton . 



350 



■2 300 



S 250 



^ 200 



i 150 

E 

z 100 

50 





kill 



500 



Paleozoic . 

400 



iiiil.liilaiLi..liljLi 



Tr 



300 200 

Geological time (1 0' years! 



Mesozoic 



dilluk 



Cenozoic 



100 



30 WORLD ATLAS OF BIODIVERSITY 



Table 3.1 

The principal mass 
extinctions in the 
Phanerozoic fossil record 

Source: Summarized from Kemp , 
Hallam and Wignalt , 



extinct during geologically very short periods of 
time, in some cases possibly as little as a few 
hundred thousand years or even less. Five such 
phases (based chiefly on extinction of marine 
species] are recognized during Phanerozoic 
time, late in each of the Ordovician, Devonian, 
Permian, Triassic and Cretaceous periods 
(Table 3.1). 

Although each of the 'Big Five' mass 
extinctions had a very profound effect on then 
contemporary life, they are not isolated peaks 



standing out from a constant, very low, 
background rate. Rather, the extinction rate 
has varied continuously throughout the 
Phanerozoic with periods of more or less 
elevated rates of extinction, of which the most 
extreme can be characterized as mass extinc- 
tion events. A frequency plot (Figure 3.5| of the 
percentage of species becoming extinct in each 
1 -My interval of the 600-My record of animal life 
provides an indication of this variation in 
extinction intensity'. The plot takes the form of 



D3TB species 

(l^lyagol loss (%1 



Late Ordovician 4A0 



85 




□loiic cnange 



The last but largest of several extinction events during the Ordovician. More than 
25% of marine invertebrate families lost. Entire class Graptolithina reduced to a 
few species; acritarchs, brachiopods, conodonts, corals, echinoderms, trilobites, 
all much reduced. 



Cooling; Warming 
Marine regression 
Marine transgression 
and anoxia 



Late Devonian 365 



End Permian 250 V5 



End Triassic 205 



End Cretaceous 66 75 



Mass extinctions came at the end of prolonged period of diversity reduction. Marine transgression 

Rugose corals lost >95% of shallow water species; stromatoporoid corals and anoxia 

reduced by half and reefs disappeared; brachiopods lost 33 families; ammonoids 

and trilobites severely affected. Fishes suffer only major mass extinction; all 

early jawed fishes (placodermsl disappear and most agnatha. First major crisis 

in plants; diversity greatly reduced, but spread of first tree, the gymnosperm 

Archaeopteris. 

The most severe extinction cnsis: metazoan life came within a few percent of total Volcanism 

extinction.Tabulate and rugose corals terminated, complex reefs disappear (return Warming 

after 8-My gap); echinoderms almost wiped out; worst crisis in history of Marine transgression 

foraminifera; severe extinction in ammonites, brachiopods, bryozoa, mollusks. 

Some losses in early ray-fin fishes. Major loss of terrestrial vertebrates (75% of 

families! and insects (8 of 27 insect orders extinctl. Mass extinction in plants: large 

plants including peat-forming trees lost (spread of small conifers, lycopods and 

guillwortsi; sudden unprecedented abundance of fungal spores at end of period. 

Mass extinction in marine invertebrates, especially brachiopods, cephalopods Marine regression 

and mollusks; also mass disappearance of scleractinian corals and sponges. 
Several seed fern families lost; some land vertebrates lost, but evidence for 
mass extinction questionable. 

Radical change in planktonic foraminifera; 85% of calcareous nanoplankton lost. Impact of large meteor 
also all ammonites, belemnites and many bivalves; losses in echinoids and Volcanism 

corals. Many marine reptiles extinct (ichthyosaurs, plesiosaurs, mosasaurs); Cooling 

significant losses in freshwater and terrestrial vertebrates, including last Marine regression 

dinosaurs (high turnover throughout dinosaur history - end Cretaceous unusual 
in that no replacements emerged]. Mass extinctions in plants: highest (possibly 
60% species loss! in angiosperms, lowest in ferns. 



m 



Biodiversity through time 31 



an even [eft-sl<ewed curve, with no substantial 
discontinuity between rare periods of high 
extinction rate Imore than 60 percent of species 
extinct per one million years) and the most 
frequent rate (10-15 percent); the mean 
intensity is 25 percent of species extinct per 
1-f^y interval. This suggests that mass extinc- 
tions may arise from causes not qualitatively 
different from those associated with extinctions 
at other times. It should also be remembered 
that because of the extremely long duration of 
the Phanerozoic during which species have 
been constantly becoming extinct, and the very 
short duration of the mass extinction events 
themselves, the latter account for only a small 
percentage (estimated at around 4 percent) of 
all extinctions. 

The precise causes and timespans of each 
of the mass extinctions have been the subject of 
much debate and study"". It is now widely 
accepted that the late Permian mass extinction 
was a long-term event, lasting for 5-8 million 
years. It appears to have been associated with 
geologically rapid global physical changes 
(including the formation of the supercontinent 
Pangaea], climate change, and extensive, 
tectonicatly-induced marine transgression and 
increased volcanic activity. The late Cretaceous 
extinction is probably the best known, but in 
terms of overall loss of diversity is also the least 
important. There is strong evidence that this 
extinction event was associated with climate 
change following an extra-terrestrial impact, 
although this remains somewhat controversial. 
The late Ordovician event appears to be 
correlated with the global Hirnantian glaciation, 

REFERENCES 

1 Raup, D.M. 1994. The role of extinction in evolution. Proceedings of the National 
/Academy o^Sc/ences 91:6758-6763. 

2 Sepkoski, J.J. 1992. Phylogenetic and ecologic patterns in the Phanerozoic history of 
marine biodiversity. In: Eldredge, N. (ed.) Systematics, ecology and the biodiversity 
crisis, pp. 77-100. Columbia University Press, New York. 

3 Jablonski, D. 1991. Extinctions: A paleontological perspective. Science 253: 754-757. 

4 Benton, M.J. 1995. Diversification and extinction in the history of life. Science 268: 52-58. 

5 Alroy, J. et al. 2001. Effects of sampling standardization on estimates of Phanerozoic marine 
diversification. Proceedings of the National Academy of Sciences 98: 6261-6266. 

6 Mojzsis, S.J. et ai 1996. Evidence tor life on Earth before 3,800 million years ago. Nature 
384: 55-59. 

7 Nisbet, E.G. and Sleep, N.H. 2001. The habitat and nature of early life. Nature A09: 1083-1091. 

8 Farmer, J.D. 2000. Hydrothermal systems: Doorways to early biosphere evolution. GSA 



with three separate episodes of extinction 
spread over only 500 000 years. In all cases, 
however, the ability to determine accurately the 
timing and periodicity of extinction is heavily 
dependent on the completeness of the fossil 
record and the reliability and precision of 
stratigraphic analysis. 

A mass extinction period is typically 
followed by a phase of 5-10 My of very low 
diversity, with a handful of species dominant in 
fossil faunas and floras. When diversity again 
increases the biota may be very different in 
composition from those preceding. In several 
instances, groups previously showing low 
diversity have radiated and spread widely 
following the demise of groups previously 
dominant; e.g. ray-finned fishes diversified 
following loss of placoderms; quillworts and 
seed ferns diversified for a time after loss of 
glossopterids; and the mammals radiated 
after loss of many terrestrial reptile groups. 
Reef organisms provide a classic example: 
reef communities have been lost several 
times during the Phanerozoic but with each 
reappearance the main reef-building forms 
have been different'^ Important new evo- 
lutionary opportunities arise for the lineages 
that survive mass extinction events, which can 
radically redirect the course of evolution'"'. 

The present-day diversity of living 
organisms is the result of a net excess of 
originations over extinctions through geological 
time. Figure 3.6 shows the number of family 
extinctions in each interval of geological time, 
and the cumulative diversity of families, both 
marine and terrestrial. 



32 WORLD ATLAS OF BIODIVERSITY 



Today 10. Available online (accessed January 2002] at http://www.geosoclety.org 

9 Rasmussen, B. 2000. Filamentous microfossils in a 3,235-mi[licn-year-old 
vulcanogenic massive sulphide deposit. Nature AOb: 676-678. 

10 Watanabe, Y., Martin, J.E.J, and Ohmoto, H. 2000. Geochemical evidence for terrestrial 
ecosystems 2.6 billion years ago. Nature ^08: 574-578. 

1 1 Brocks. J.J. ef al. 1999. Archean molecular fossils and the early rise of eukaryotes. 
Sc/ence 285: 1033-1036. 

12 Javaux, E.J., Knoll. A.H. and Walter. M.R. 2001. Morphological and ecological 
complexity In early eukaryotic ecosystems. Nature k\l: bb-k'^ . 

13 Knoll. A.H. 1995. Proterozolc and early Cambrian protlsts: Evidence for accelerating 
evolutionary tempo. In: Fitch. W.M. and Ayala. F.J. ledsl. Tempo and mode in 
evolution: Genetics and paleontology 50 years after Simpson, pp. 63-68. National 
Academy Press, Washington DC. 

U McMenamin. M.A.S. 1989. The origins and radiation of the early metazoa. In: Allen. K.C. 
and Bnggs. D.E.G. ledsl. Evolution and the fossil record, pp. 73-98. Belhaven Press, a 
division of Pinter Publishers Ltd, London. 

15 Margulls, L. and Schwartz, K.V. 1998. Five kingdoms. An illustrated guide to the phyla 
of life on earth. 3rd edition. W.H. Freeman and Company, New York. 

16 Wellman, C.H. and Gray, J. 2000. The microfossil record of early land plants. 
Philosophical Transactions of the Royal Society of London. Series B. Biological 
Sciences 255: 717-731. 

17 Wikstrom, N., Savolalnen, V. and Chase. M.W. 2001. Evolution of the angiosperms: 
Calibrating the family tree. Proceedings of the Royal Society of London. Series B. 
Biological Sciences 268: 221 1 -2220. 

18 Labandeira. C.C. and Sepkoski. J.J. 1993. Insect diversity in the fossil record. Science 
261: 310-315. 

19 Benton, M.J. 1999, The history of life: Large databases In palaeontology. In: Harper, 
DAT. (ed.l Numerical palaeontology: Computer-based modelling and analysis of 
fossils and their distributions. John Wiley and Sons, Chichester and New York. 

20 May, R.M., Lawton, J.H. and Stork, N.E. 1995. Assessing extinction rates. In: Lawton, 
J.H. and May, R.M. ledsl. Extinction rates, pp. 1-24. Oxford University Press, Oxford. 

21 Jablonski, D. 1995. Extinctions In the fossil record. In: Lawton, J.H. and May, R.M. 
ledsl. Extinction rates, pp. 25-44. Oxford University Press, Oxford. 

22 Crane, PR. 1989. Patterns of evolution and extinction in vascular plants. In: Allen, K.C. 
and Bnggs, D.E.G. ledsl. Evolution and the fossil record, pp. 153-187. Belhaven Press, 
a division of Pinter Publishers Ltd, London. 

23 McElwain, J.C, Beerling, D.J. and Woodward, Rl. 1999. Fossil plants and global 
warming at the Triassic-Jurassic boundary. Science 285: 1386-1390. 

24 Kemp, T.5. 1999. Fossils and evolution. Oxford University Press, Oxford. 

25 Hallam, A. and Wignall, PB. 1997. Mass extinctions and their aftermath. Oxford 
University Press, Oxford. 

26 Erwin, D.H. 2001. Lessons from the past: Biotic recoveries from mass extinctions. 
Proceedings of the National Academy of Sciences 98: 5399-5403. 

27 International Subcommlssion on Cambrian Stratigraphy, http://www.unl- 
wuerzburg.de/palaeontologie/Stuff/casu6.htm laccessed January 20021. 

28 Benton, M.J. 1989. Patterns of evolution and extinction In vertebrates. In: Allen, K.C. 
and Briggs, D.E.G. ledsl. Evolution and the fossil record, pp. 218-241. Belhaven Press, 
a division of Pinter Publishers Ltd, London. 

29 Benton, M.J. led. I 1993. The fossil record 2. Chapman and Hall, London. Data available 
online In interactive format at http://lbs.uel.ac.uk/ibs/palaeo/benton/ laccessed 
January 20021; also see http://palaeo.glybrls.ac.uk/frwhole/fr2.familles 



Humans, food and biodiversity 33 



k 



HumanSp food 

and biodiversity 



THE LINEAGE LEADING TO THE HUMAN SPECIES EMERGED from an ancestry among the 
apes, almost certainly in Africa around 5 million years ago. Fossils usually attributed 
to the genus Homo itself date from the late Pliocene, perhaps 2 million years ago, and 
anatomically modern humans appeared some 100 000 years or more before the present. 

Agriculture developed independently in several regions around 10 500 years ago, in 
association with increased population density. The global human population continued to grow 
only slowly until the 19th century, when revolutionary developments in agriculture, industry 
and public health triggered an exponential rise that has continued to the present day. 

Agriculture is a means to channel the Earth's resources into production of human bodies. 
Humans have converted large areas of terrestrial habitat and use more than one third of net 
primary production on land. They are strongly implicated in the extinction of many large 
terrestrial mammal and bird species in prehistory, and are responsible for habitat change 
and exploitation that have caused the decline and extinction of many species in recent times. 



HUMAN ORIGINS 

The human species evolved as a natural 
element of diversity in the living world, and it 
is a simple ecological imperative that humans 
depend on other species and communities to 
supply the basic requirements of existence 
and to maintain biosphere function. 

The creation of organic compounds by 
photosynthesizing organisms is the point at 
which the sun's energy enters the biosphere 
(Chapter II; humans and other animals are 
unable to capture energy in this way and must 
consume and digest either primary producers 
or other organisms that are themselves 
dependent on primary producers, in order to 
obtain these energy-rich organic compounds 
for their own activities. 

While humans are doing nothing funda- 
mentally different from other animals, with 
the benefits of society and technology, which 
serve to increase the rates of resource 
extraction, they are uniquely successful at it. 
Self-evidently, humans have not arrived at 
their extraordinarily dominant position on 
the planet overnight. The growth of their 
influence can be traced back several million 



years, to well before the Pleistocene when a 
stone-tool-wielding hominid first emerged 
somewhere in eastern Africa. 

Climate during the past 2 million years 
The Earth's climate appears always to have 
been in a state of flux. Generally the degree 
of accuracy in our understanding of global 
climate, and certainly the degree of resolution 
in the timescale of climate change, decreases 
the further back in time we go. It is therefore 
difficult to compare periods that in geological 
terms are recent with more distant times. 
However, it does seem that during the past 
2 million years there have been numerous, 
intense climate changes that were at least as 
severe, or perhaps more severe, than any 
recorded earlier in the Earth's history'. It is 
during this period that hominids very similar 
to modern humans first appear in the fossil 
record. By the early Holocene, some 10 000 
years ago, technologically sophisticated 
humans had spread to all the major land 
masses except Antarctica, and had evidently 
started to exert a major, and ever-growing, 
impact on the biosphere. Indeed there is 



34 WORLD ATLAS OF BIODIVERSITY 



Map 4.1 

Early human dispersal 

A highly generalized view 
of the colonization of the 
world by advanced humans 
from the early Pleistocene 
onwards. Coastline is shown 
150 m lower than at present 
and. with northern 
hemisphere ice cover, 
represents an approximation 
of that at periods of glacial 
maximum. Dates represent 
earliest well-established 
time of arrival; arrows 
indicate general net 
direction of dispersal, not 
actual routes. Presence and 
dispersal of early humans 
are not represented: the 
lineage is believed to have 
arisen in Africa and later 
fossil material dated to 
between 1.8 and 1.4 million 
years before the present is 
known from several sites in 
Eurasia. 



Ice 

Human dispersal 



^< 



Hawaiil 




ilCuba,"' 
HIspani&la). 



x^<^ 



-y^ 



000 Years before the present 



evidence that such impacts - for example in 
the extinction of large mammal and bird 
species - were already being felt con- 
siderably earlier than this. However, because 
the rise of humans coincides with a period of 
major climatic and ecological fluctuations, it 
is often difficult to disentangle the effects 
of the former from the latter, and so the 
precise nature of these impacts remains 
controversial. 

The Tertiary period, which began some 
65 million years ago and ended with the start 
of the Quaternary (the Pleistocene and 
Holocenel 1.8 million years ago, is charac- 
terized overall by a gradual decrease in 
global temperature and increase in aridity 
Superimposed on this general pattern were 
many oscillations, occurring on a timescale 
of thousands of years'. These oscillations are 




\ 


1 500 




% 








" ■ Vv , ■ 


-1 500 



-^^^,^.J6.. ::\ I 






believed to be linked to cyclic variations in the 
Earth's position in its orbit around the sun, 
known as Milankovitch cycles. These cycles 
notwithstanding, the climate during virtually 
the whole of the Tertiary was notably warmer 
than at present. 

Around 1.8 million years ago, at the 
very start of the Pleistocene^ there was 
apparently rapid global cooling leading to the 
start of a period dominated by marked 
climate cycles of around 100 000 years' 
duration. For long periods of each cycle 
global temperatures were significantly lower 
than they are today, and extensive areas of 
the northern hemisphere land masses were 
covered in ice sheets. Detailed analysis of 
climate changes over the past AOO 000 years 
or so, particularly through examination of 
Antarctic ice cores', indicate that each cycle 



/ 



Humans, food and biodiversity 35 








I 



V-.\ ■ • ^<^ t ■ -"i' ■! ' ^''''^-^' 



:.>V. 



■Jl^' 




^'•■\^^J^- 



56 000 










}i' (M, 



' 3 500 



(New 
Zealand! 



ii-..^. 



over this period Inas been broadly similar, 
vi/ith a gradual decline in average temp- 
erature, although w/ith many minor and some 
large oscillations (relatively colder periods 
being referred to as stadials and warmer 
ones as interstadialsl, followed by a short 
period of intense warming in which temp- 
eratures rose from those of fully glacial 
conditions to those characterizing a warm 
interglacial state, perhaps sometimes over 
only a few decades. 

Mean global temperature during glacial 
periods was around 6''C cooler than during 
interglacials, with cooling more pronounced at 
the poles than the equator. The mid-latitudes 
and equatorial regions were probably some- 
what more arid than at present. The large 
amount of water locked up in the greatly 
expanded polar ice caps meant that mean sea 



level during glacial periods was probably 
around 100-150 meters lower than at present. 
Until around 11000 years ago, each 
temperature peal< was apparently followed 
by an almost immediate decline as the 
next glacial cycle began'. Overall, it appears 
that during the past half million years the 
Earth's climate has been as warm as, or 
warmer than, today's for only around 
2 percent of the time', it seems lil<ely 
that this also holds true for the early 
Pleistocene. The end of the last glacial cycle, 
around 1 1 000-12 000 years ago, marked the 
start of the Holocene. Temperatures similar 
to today's have prevailed throughout the 
(Holocene, making it by far the longest true 
interglacial period for at least the past half 
million years and probably for the last 
1.8 million years. 



36 WORLD ATLAS OF BIODIVERSITY 



Human origins and dispersal 

The origins and early history of humans are 
among the most controversial subjects in 
paleontology. Remains are generally scarce 
and often open to varying interpretation. 
Nevertheless, consensus has emerged over 
the broad outlines. It is likely that the direct 
ancestors of humans - the homlnid line - 
diverged from the apes In Africa during a cool, 
dry phase of the late Miocene, around 
6 million to 5.5 million years ago. The primary 
evidence for this divergence is in the form of 
'molecular clocks' calculated from comparison 
of human and ape genetic materiall Relevant 
fossil material from this time horizon Is 
sparse and often of uncertain status, but 
material from Ethiopia dated to 5.8 million to 
5.2 million years ago' is believed to represent 
Ardipethecus, generally accepted as the 
earliest known member of the homlnid 
branch. Other species of this genus are 
known from before A million years ago, with 
various species of Australopithecus and 
Paranthropus, all from eastern Africa, some- 
what younger in age. 

Sometime during the middle to late 
Pliocene, 2.5 million to 1.8 million years ago, 
the genus Homo is thought to have evolved 
from Australopithecus stock^'. Until com- 
paratively recently it had been assumed that 
early man then remained confined to Africa 
until less than 1 million years ago. However, 
two well-preserved skulls recently found In 
the Caucasus (southern Georgia) have been 
dated to about 1.8 million years' (and variously 
attributed to l-lomo ergaster, otherwise 
known from Africa, or H. erectus]. Elsewhere, 
the earliest stone tools from Turkey and 
southwest Asia have been dated to about 1.5 
million years ago, and from northeast Asia to 
nearly 1.A million years', while tools and 
fossils from East and Southeast Asia might be 
as old as 1.9 million years. These finds 
suggest that populations of early forms of 
l-iomo spread through the southeastern 
fringes of Europe as well as into Asia within at 
most a few hundred thousand years of the 
genus originating in Africa^ 

The earliest hominid remains elsewhere in 
Europe date from 780 000 years ago in Spain'", 
followed by somewhat different material 



known from several sites widely distributed in 
northern Europe of about 500 000 years in age 
or less". Fossil material from these two 
periods may be attributed to H. antecessor 
and H. heidelbergensis. respectively. The 
diversity of homlnid material and the range 
of dates suggest that the spread of early 
hominids was complex and multidirectional, 
with possibly three lineages moving into 
Eurasia from Africa. 

Ivlodern humans, i-iomo sapiens, are 
believed to have appeared sometime after 
200 000 years before the present. Anatomically 
modern human remains are known from 
around 100 000 in Africa and the Middle East, 
with more recent dates In East and Southeast 
Asia. The first appearance in Europe dates 
from about AQ 000 years ago"", with 
expansion apparently from the east toward 
the west, where in places premodern 
(Neanderthal) humans remained until about 
28 000 years ago. One interpretation of the 
several lines of genetic evidence suggests, on 
the basis of mathematical models, that the 
modern human population Is derived from a 
small founding population of perhaps 10 000 
breeding Individuals that existed sometime 
between 130 000 and 30 000 years before 
the present'-' ". 

The earliest evidence of hominids outside 
Africa, Europe and Asia Is much more recent. 
The age of the human skeleton found In 197i 
at Lake Mungo in Australia, evidently the 
earliest human remains known on the 
continent, has been estimated at 62 000 years 
before the present'', but a date close to 56 000 
years Is now widely accepted for human 
arrival on the continent". 

in the Americas, the oldest good evidence of 
human presence Is that of a coastal settlement 
in Chile dated around U 000-15 000 years 
ago'"', although even this Is far from universally 
accepted amongst archeologlsts. There are 
very controversial and now widely questioned 
claims for evidence of human settlement much 
earlier, most notably that dating from 32 000 
years before the present from Pedra Furada in 
northeast Brazil". The earliest unequivocal 
evidence of widespread occupation in the 
Americas comes from the so-called Clovis 
hunting culture whose oldest remains are 



Humans, food and biodiversity 37 



generally dated at around 12 000 years before 
the present". Settlement of the Caribbean 
islands including Cuba and Hispaniola appears 
to have taken place considerably later, around 
6 000 years ago'", 

Archeological evidence indicates that 
colonization of the Pacific Islands east of New 
Guinea began around it 000 years ago, when 
much of Melanesia and Micronesia was 
settled. Fiji and Samoa were probably colon- 
ized around 3 500 years ago. and the outliers 
of Hawaii, Easter Island and New Zealand 
within the last 1 500 years-'. In the Indian 
Ocean. Madagascar was probably first settled 
around 1 500 years ago'*. 

Technology 

The earliest evidence of tool manufacture by 
hominids is from the Gona River drainage in 
northern Ethiopia where stone tools have 
been dated to at least 2.5 million years ago. 
These tools are small (generally less than 10 
centimeters long) and simple, but are alteady 
of relatively sophisticated manufacture, 
suggesting that even older artifacts will 
eventually be found". They are of essentially 
the same design as those dated from 2.3 
million to 1.5 million years ago from other 
sites in East Africa (e.g. Lokalalei in the Lake 
Turkana basin in northern Kenya and the 
Olduvai gorge in Tanzania) indicating effective 
technological stasis tor at least 1 million 
years (from 2.5 million to roughly 1.5 million 
years agol". Collectively these tools are 
referred to as products of the Oldowan stone 
tool industry". 

Around 1.5 million years ago much 
more sophisticated and often larger tools 
including hand axes and cleavers suddenly 
appear in the archeological record in East 
Africa. These are referred to as Acheulian 
tools. The Acheulian tool industry spread into 
Europe, the Near East and India, and remained 
apparently relatively unchanged until around 
200 000 years ago, showing similar temporal 
persistence to the Oldowan industry. 

Evidence of tools and artifacts made from 
organic materials from the Paleolithic is 
understandably extremely scarce. In 1995, 
however, three large wooden implements 
very similar in design to modern-day javelins 




and dated to around AOO 000 years ago were 
discovered at Schdnmgen in Germany. These 
were found in association with a smaller 
wooden implement (probably a throwing 
stick), stone tools and the butchered remains 
of more than ten horses, and can be per- 
suasively interpreted as throwing spears used 
in systematic, organized hunting". 

There is also intriguing although indirect 
evidence from Southeast Asia dating to 
around 900 000 years ago that hominids at 
this time, at least in this region, were capable 
of repeated water crossings using watercraft. 
The evidence is in the form of stone tools 
dated to this age from the island of Flores in 
eastern Indonesia. Even at the time of the last 
glacial maximum (when global sea levels 
would have been at their lowest), reaching 
Flores from the Asian mainland would have 
required crossing three deep-water straits 
with a total distance of at least 19 kilometers. 
The impoverished - and typically island - 
nature of the Paleolithic fauna of Flores would 
appear to preclude the existence of any now 
submerged land bridge". The next oldest, 
again indirect, evidence for the use of 
watercraft is the colonization of Sahul, 
probably sometime around ^0 000 to 60 000 
years ago. Even at times of lowest sea level, 
this would have necessitated crossing some 
100 kilometers of open sea. 

Fire 

At some point in their evolutionary history, 
hominids clearly learnt to control, manipu- 
late and, presumably later, to start fires. 
Determining even very approximately when 
this may have happened is difficult, and as 



■4 



m 



Watercraft may have played 
an important role in human 
dispersal for millennia. 



WORLD ATLAS OF BIODIVERSITY 




Large-scale burning of 
vegetation is one of the 
major fiuman impacts on 
the biosphere. 



with all else to do with human evolution, 
controversial. This is chiefly because the 
existence of natural fires caused by lightning 
strikes and volcanic activity greatly compli- 
cates the interpretation of the archeological 
and geological record - an association 
between artifacts and evidence of burning 
does not necessarily indicate a direct link 
between the two'°. 

The earliest dated associations between 
artifacts and burning that could be construed 
as deliberate use of fire are from Africa. Stone 
tools and splintered bones, which can be 
interpreted as evidence of butchery, have 
been found associated with clay baked at 
several hundred degrees for several hours at 
Chesowanja in Kenya, in deposits around 
1.4 million years old''". The characteristics of 
the clay are consistent with formation 
beneath a campfire, but are also consistent 
with formation around a slow-burning tree 
stump that could be associated with a natural 
bushfire". Charred animal bones and other 
evidence of human occupation from just over 
1 million years ago have been found at 
Swartkrans cave in South Africa, although 
similar problems of interpretation apply. 

Numerous sites in Europe and Asia provide 
evidence of human occupation and associated 
fire from the mid-Pleistocene, some 400 000 
years ago. The best known of these is at 
Zhoukoudien in China, although even here 
the evidence for deliberate use of fire is 
widely considered equivocal"™. Others are 
at Torralba-Ambrona (Spain), Terra Amata 
IFrancel, Westbury-sub-Mendip (England! 
and Vertesszollos (Hungary]''. 



Large-scale burning of terrestrial vege- 
tation is undoubtedly one of the major 
present-day impacts by humans on the 
biosphere, and is believed to constitute 
around one third of current annual anthropo- 
genic carbon dioxide emissions. Evidence of 
earlier impact is invariably circumstantial. It 
is difficult to demonstrate widespread 
biomass burning in the fossil or subfossil 
record and even harder to demonstrate a link 
with human activities - natural fires may be 
caused by storms or volcanic activity and may 
be expected to vary in extent, frequency and 
intensity according to prevailing climatic 
and ecological conditions. 

However, the abundance of elemental 
carbon in marine sediments off Sierra Leone 
in Africa can be persuasively interpreted as a 
measure of intensity of biomass burning in 
sub-Saharan Africa. Analysis of a core from 
these sediments covering the past million 
years or so indicates that inferred fire 
incidence in the region was low until around 
400 000 years ago. Since then, five episodes of 
intense burning of vegetation can be inferred, 
all except the most recent coinciding with 
periods when the global climate was changing 
from interglacial to glacial. The current peak 
is unique in that it is occurring during an 
interglacial period. The change from inter- 
glacial to glacial climate is generally 
associated with increased aridity, so that 
vegetation may be expected to be more 
vulnerable to fire. It would also be expected 
that a substantial fuel base in the form of 
woody biomass would have accumulated 
during the warmer, wetter, interglacial. It may 
well be merely coincidental, but it is intriguing 
that the period of increased fire incidence in 
the Sierra Leone core - around 400 000 years 
ago - coincides with the timing of the first 
widespread evidence for hominid use of fire. 

Food resources 

The early australopithecine members of the 
hominid lineage appear, on craniodental 
evidence, to have been well adapted to 
consume hard but brittle items, such as nuts 
and seeds, and soft items, such as fruits'^ 
Use and development of stone tools by later 
hominids is usually thought of as associated 



Humans, food and biodiversity 39 



with meat consumption, and cut and hammer 
marks on large mammal bones from hominid 
sites are evidence of this, but microscopic 
wear patterns found on some tools indicate 
they were used to scrape wood and cut coarse 
plants such as reeds or grass". Oldowan 
and Acheulian stone tools are rarely found 
more than 10-20 kilometers from their rock 
sources, suggesting that their owners ranged 
over a relatively small area", so if hominid 
populations at the time were small in size, the 
environmental impacts of targeted resource 
extraction would have been minimal. 

The ratio of different forms of carbon and 
nitrogen incorporated in human bone colla- 
gen can provide information on the relative 
importance of food from terrestrial, coastal or 
freshwater habitats. Such 'stable isotope 
analysis' of collagen extracted from archeo- 
logical remains, coupled with other material 
evidence, suggests that pre-modern humans 
(Neanderthals] in Europe, although omni- 
vorous and opportunistic, behaved as preda- 
tory carnivores^'. Similar evidence for slightly 
later human remains suggests a significant 
broadening of the resource base, owing 
particularly to increased use of freshwater 
resources". Remains found at Ivlediterranean 
sites suggest further diversification of human 
diet and the means of food gathering". 



associated with an apparent rise in human 
numbers during the later Paleolithic and 
approaching the start of the Neolithic. Here, 
there was increasing use of agile small game, 
such as partridges and hares, and apparently 
decreasing reliance on slow-maturing, easily 
collected forms such as tortoises and shell- 
fish, the average size of which decreased over 
time at a rate consistent with the effects of 
excess harvesting, possibly indicative of 
increasing human density^'. 

Origins of agriculture 

Until the end of the Pleistocene, humans 
evidently depended on hunting and gathering 
of wild resources for their sustenance. 
Around the end of this period a radical 
change was initiated - the emergence of 
crop-based agriculture and domestication of 
livestock - phenomena which appear to have 
arisen independently in Africa, Eurasia and 
the Americas. 

The earliest direct evidence for animal 
domestication, of the dog Canis. dates from 
around 14 000 years ago in Oberkassel in 
Germany". However, analysis of molecular 
clocks' appears to show that the dog diverged 
from its wild ancestor, the wolf Canis lupus, 
far earlier than this - perhaps more than 
100 000 years ago". It is quite possible that 




The dog diverged from 
its wild ancestor, the 
wolf, more than 100 000 
years ago. 



40 WORLD ATLAS OF BIODIVERSITY 



domestication of the dog as a guard animal 
and aid in hunting predates domestication of 
other animats and plants as food sources 
by many tens of thousands of years, but In 
the absence of archeological evidence this 
remains speculative. 

The study of plant and animal domestication 
for agricultural purposes is a rapidly expand- 



Box iA Loss of diversity in agricultural genetic resources 



Cfilna 


Wheat varieties 


Korea 


Garden tandraces 


IRep.l 




Mexico 


Maize varieties 



With the rise of Industrial-scale agriculture and commercial breeding, many local 
agricultural genetic resources, both crops and livestock, have been lost and replaced by 
modern, genetically uniform types specialized for superior production in higher Input 
systems. There is much evidence that many local varieties possessed features of adaptive 
value in a particular environment and cultural context, and the precautionary principle 
argues that the diversity of domesticated forms and their wild relatives should be conserved 
where possible in order to maintain options for future breeding Improvements. A complete 
picture of the global reduction In local genetic resources is not available, but there is 
abundant evidence at national level of the enormous scale of genetic erosion in crop plants. 

About 1 000 110 percent) of 10 000 vaneties used 
in 1949 remained in 1970s 

About 25 percent of landraces of U crops grown 
in home gardens In 1985 remained in 1993 

Only 20 percent of maize varieties planted in 
1930s remain 

USA Varieties of apple, cabbage. Only 15-20 percent of varieties grown in 1804- 

field maize, pea, tomato 1904 available at present 

A growing number of countries have documented and evaluated livestock resources, see 
Appendix 3 for summary of mammal breed status. 



ing field, based on study of fossil pollen 
records, archeological remains, the genetics 
of present-day crops and their close relatives, 
and remaining indigenous agricultural 
systems". The development of agriculture 
began independently in different continents 
and proceeded at different rates, while early 
cultivators undoubtedly continued to rely 
heavily on hunting and gathering from the wild. 
The first development of agriculture as an 
integrated system for food production was 
based in the Fertile Crescent, composed of 
the uplands of Anatolia and western Iran and 
the arid lowlands to the south, with the first 
tangible evidence approaching 1 1 000 years in 



age. Wheat Triticum. barley Hordeum, rye 
Secale, pea Pisum and lentil Lens, cattle 80s, 
sheep Ovis. goat Capra and pig Sus were all 
domesticated in this region, and formed the 
basis of the Neolithic peasant economy that 
spread steadily through much of western 
Eurasia into surrounding areas after about 
10 000 years before the present. The system 
integrated the use of food plants, cereals 
especially, and domesticated animals for 
fertilizer and power as well as food. 

Although early plant domesticates are 
known elsewhere in the world. Integrated 
agricultural systems appear to have taken 
longer to develop, perhaps in part because of 
the absence of domesticated animals. For 
example, in Middle America seeds of domes- 
ticated squash Cucurbita are known from 
about 10 000 years ago, maize from 6 300 years 
and beans from 2 300, with village-based 
farming economies evidently taking several 
thousand years to develop'". Farming systems 
based on rice cultivation In Asia appear to date 
from 7 500 years ago. 

Analysis of mitochondrial DNA has 
indicated that a multiple maternal origin is 
general among domestic livestock species, 
i.e. female animals from more than one wild 
stock have contributed, at different times and 
places, to the present genetic diversity among 
each breed". For each of the four major 
breeds (cattle, sheep, goats, pigs), this 
evidence is consistent with archeological 
findings in suggesting a primary center of 
origin around the Fertile Crescent region, but 
also suggests additional centers in Asia. 

Ivlost of the major crops on which humans 
presently depend have been grown con- 
tinuously since the early or middle Holocene. 
They have been constantly selected over this 
period and have developed large amounts of 
useful genetic variation. Indeed the success 
of Individual crop species over wide geo- 
graphical areas is partly determined by 
their flexibility in evolving and sustaining 
genotypes suitable for local environments. 
Conventional breeding Involves the selection 
and crossing of desirable phenotypes within 
a crop in order to create more productive 
genotypes. The process of harvest, storage 
and sowing alone may have assisted In the 



Humans, food and biodiversity 41 



selection of traits sucfi as non-shattering seed 
fieads, uniform ripening of seeds, uniform 
germination, targe fruits and seeds, and easy 
storage. Breeding methods have increased the 
rate of introgression of desired genetic traits 
into new cuttivars; genetic modification is now 
possible at the level of incorporating individual 
genes directly into genomes. 

Domesticated crops and livestock have 
been transported around the world probably 
since full-scale agriculture began, e.g. 
wheats are recorded in areas outside their 
presumed center of origin at least 8 000 
years before the present. Some crops became 
increasingly widely distributed after the 1 500s 
when European colonists moved out of their 
home continent. 

CURRENT FOOD AND NUTRITION 

Perhaps as many as 7 000 of the 270 000 
described plant species have been collected 
or cultivated for consumption'"', but very few, 
some 200 or so, have been domesticated, and 
only a handful are crops of major economic 
importance at global level". The variety of 
species used is limited more by production 
and cultural factors, such as tradition and 
palatability, than by nutritional value. Twelve 
crop plants together provide about 75 percent 
of the world's calorie intake. These comprise 
lin alphabetical order]: bananas/plantains, 
beans, cassava, maize, millet, potatoes, rice, 
sorghum, soybean, sugar cane, sweet potatoes 
and wheat''. More than 40 mammal and bird 
species have been domesticated, around 12 
of which are important to global agricultural 
production (see Appendix 3). Some, such as 
cattle, pigs, goats, sheep and chickens, are 
fundamental to many non-industrial agri- 
cultural systems, and provide a wide range of 
products in addition to foodstuffs. 

Although the global consumption of wild 
terrestrial species cannot be assessed acc- 
urately, the amount is likely to be insig- 
nificant in comparison to products from just 
three domestic forms: pigs, cattle and 
chickens. On the other hand, fishes from wild 
sources are an important nutrient source, 
and the amount harvested and consumed 
is known to be greatly under-reported in 
official statistics. 



At world level, cereals are the most 
important single class of food commodities, 
providing around 50 percent of daily calorie 
intake and 45 percent of protein. In contrast, 
meat provides around 15 percent of the 
protein, and fishery products only some 
6 percent, or 15 percent of all animal protein. 
In each case, when comparing sources of 
calories, protein and fat in the global human 
diet, just two or three commodities stand out 
from a large number of commodities of lesser 
importance. Rice and wheat together provide 
around 40 percent of the world supply of both 
calories and protein, while milk and pigmeat 
are key sources of fat. calories and protein 
(see Table 4.11. 

At country level, a much wider variety of 
plant species are important in that they make 
a significant contribution to human nutrition". 
Around 22 species and groups of species 
provide more than 5 percent of the per capita 
supply of either calories, protein or fat in at 
least ten countries of the world. Notes on 
the origin, uses and genetic resources in 
these 22 crops are tabulated in Appendix 2, 
together with briefer information on the 
remaining 50 or so crops that are also 
nutritionally important but to a smaller 
number of countries. 

Although the vast bulk of human food 
supply on a global level is derived from 



Table 4.1 

Top ten food commodities, 
ranked by percentage 
contribution to global 
food supply 

Note: This information is 
partly determined by the way 
food commodities are 
aggregated for reporting 
purposes; for example, 
fishery products collectively 
provide about 6% of the 
hypothetical global protein 
supply, but no individual 
fishery commodity is 
important enough to appear 
in the table. 



Source: FAO food balance sheets 
1997 data. 



^^^HJ^^^^HJH^^HHH 


Rice (milled equiv.l 


21 


Wheat 


22 


Pigmeat 


14 


Wheal 


20 


Rice (milled equiv.] 


15 


Soybean oil 


10 


Sugar (raw equiv.l 


7 


Milk (excl. butter] 


9 


Milk (excl. butter] 


9 


Maize 


5 


Pigmeat 


5 


Rape and mustard oi 


6 


Milk (excl. butter] 


4 


Bovine meat 


5 


Sunflowerseed oil 


6 


Pigmeat 


4 


Poultry meat 


5 


Palm oil 


5 


Soybean oil 


2 


Maize 


5 


Fats, animal, raw 


5 


Potatoes 


2 


Vegetables (other than 




Bovine meat 


4 


Vegetables (other than 




tomatoes, onions] 


3 


Poultry meat 


4 


tomatoes, onions] 


2 


Eggs 


3 


Butter, ghee 


4 


Cassava 


2 


Pulses (other than 
peas, beans] 


2 






Other 


31 


Other 


25 


Other 


34 


Global mean: 2 745 




Global mean: 73.4 




Global mean: 70.1 




calories/person/day 




grams/person/day 




grams/person/day 
J 


^B 



42 WORLD ATLAS OF BIODIVERSITY 



Map 4.2 

Livestock breeds: 
numbers and status 

Color represents the 
number of mammal breeds 
in each country. Recording 
of breeds is incomplete 
globally. Pie charts 
represent the proportion of 
all mammal breeds 
associated with each FAO 
region assessed as 
threatened Igrayl or extinct 
Iblackl. 



Source; Charts calculated from FAO 
World Watch List for Domestic Animal 
Diversity \3rti edition. 2000); country 
data derived from FAO Domestic Animal 
Diversity Information System IDAD-ISI 
database, available online at 
tittp://www fao org/dad-is/ laccessed 
February 20021- 





Status 



% threatened 
% extinct 



farming, a large number of people worldwide, 
including many who are principally agricul- 
turalists or pastoratists, make extensive use of 
wild resources. In parts of Africa, bushmeat 
(meat from wild animalsl supplies most of the 
protein intake; similarly some 80 percent of 
people in sub-Saharan Africa are believed to 
rely largely or wholly on traditional medicines 
derived almost exclusively from wild sources. 
Indeed, traditional medicine continues to be 
the source of health care for the majority of 
people living in developing countries, and is 
widely incorporated in primary health care 
systems. Wild plants in many areas are 
extremely important as famine foods when 
crops fail, or may provide important dietary 
supplements, and use of fuelwood and 
charcoal from wild sources is almost universal 
in the developing world. 




V 



A small number of people continue to 
derive most of their requirements from wild 
sources by hunting, fishing and gathering. In a 
large sample 12201 of such societies, nearly AO 
percent have a high dependence on fishing, 
around one third are highly dependent on 
gathering and 28 percent on hunting of 
terrestrial resources". Fishing and gathering 
tend to be alternative activities, both of them 
complementary to terrestrial hunting. In 
contemporary conditions, fishing tends to be 
more important where temperature is lower 
(especially in high northern latitudes! while 
the converse applies to gathering. People in 
such communities also derive shelter, medi- 
cine, fuelwood and esthetic or spiritual fulfill- 
ment from wild species, and the immense 
variety of plant and animal species used has 
been well documented. 



Humans, food and biodiversity 43 




Global food supply 

National data on reported food commodity 
supplies are collated by the Food and 
Agriculture Organization of the United 
Nations (FAO)". Given information on the food 
value of these commodities, and the size of 
the human population, it is possible to 
estimate the average national food supply per 
person. Dietary food value can be broadly 
assessed in terms of energy or materials. The 
former is conventionally measured in calories 
Icalories/person/dayl, the latter in terms of 
weight of protein or fat Igrams/person/dayl. 
These standard measures take no account of 
the vitamins and minerals (micronutrientsi 
that are required for maintenance of full 
health. The nutritional value of the human diet 
varies geographically and over time, and to 
a great extent according to the state of 



development and purchasing power of the 
people concerned. 

The human population of the world 
doubled between 1950 and 1990, reached 
around 6 billion in the late 1990s, and will 
continue to grow for decades, albeit more 
slowly because of decreased reproduction 
rates. At global level, there has been enough 
food available in recent years to supply the 
entire human population with a very basic 
diet, largely vegetarian, providing 2 350 kcal 
per day. In 1992 the average food supply was 
estimated at 2 718 kcal daily lafter losses 
during storage and cooking], made up of 2 290 
kcal from plants and ^28 kcal from livestock 
products". Thus the global food supply was 
nominally sufficient for the calorific needs of 
a population 15 percent larger than the 
estimated population; a similar small annual 



w WORLD ATLAS OF BIODIVERSITY 



Diet class Class 1 


Class 2 


Class 3 


Class i 


Class 5 


Class 6 


Rice 


Maize 


Wlieat 


Milk, meat. 


Millet, 


Cassava, plantain 








wheat 


sorghum 


taro, yams 


iHain Asia 


Central 


N.Africa, 


Europe, 


Sahel, 


Central Africa, ' 


geographic 


and 


West Asia, 


N.America, 


Namibia 


Madagascar 


distribution 


S. America 

InorthI 


S. America 
(south! 


Australia 




^^^ 


Human 












population 












Ithousandsl 2 920 923 


5U911 


664 507 


9i2 924 


61867 


357 361 


% of world 












population 54 


9 


12 


17 


1 


7 


Increase in 












supply needed 












by 2050 X 2.37 


xL96 


X2.84 


xl.13 


X4.82 


x7.17 



Table U.l 

World diet classes 

Notes: Population calculation 
based on 1997 data, from 
FAO website. Excludes Japan 
and Malaysia, in anomalous 
position using six-part 
classification. Total 
population in 1 1 7 countries in 
sample: 5 462 ^93 000. Tiie 
penultimate row in fact 
shows the percent eacti class 
forms of ttiis total population 
In the sample countries; this ■ 
is assumed to be an 
acceptable surrogate for the 
total world population. 



surplus has existed since the 1970s". Talking 
these aggregated global data at face value, It 
appears that a more than sufficient amount of 
food has been produced annually during the 
past two decades to maintain the world's 
human population. Global food supply has 
doubled since the 19i0s; the Increases in 
supply over the last two decades are attri- 
buted mainly to Increased productivity (69 
percent] and secondarily to an increase in 
production area 131 percent). 

Each year during the past two decades, 
between 850 million and 900 million people 
have been undernourished". Given that there 
has been sufficient food available in the world 
overall, undernourishment must be an effect 
of unequal access to the appropriate amount 
and type of food. Unequal distribution is 
evident at macro and micro scales: some 
countries are more productive and richer in 
resources than others and, whether at nation- 
al or village level, high-status social groups 
secure better diets than others, compounded 
in some cases by gender differences. Poverty 
Is a key cause of undernutrition, and often 
also an effect of it, forming a self-reinforcing 
cycle from which It is difficult to escape 
without appropriate outside Intervention. Put 
in different terms 'food insecurity is a problem 
of lack of access resulting from either 
inadequate purchasing power or an Inade- 



quate endowment with the productive 
resources that are needed for subsistence'". 
Although the 1996 World Food Summit in 
Rome called for a 50 percent reduction In the 
global number of chronically malnourished 
people by 2015, the absolute number of 
people affected remains high, and if trends 
continue this target will not be met". 

Regional variation 

The global average diet' Is a simplifying 
abstraction that ignores an enormous amount 
of regional, national and local variation in food 
sources and in supply. Nor does it take account 
of micronutrlent availability (vitamins, miner- 
als, trace elements). I.e. substances that do 
not directly contribute to energy or protein 
Intake but which are nevertheless essential 
elements of a healthy diet. 

The human diet can be assessed in 
several ways. The FAO devised a simple 
classification to serve as the basis for an 
analysis of food requirements In relation to 
population growth". In this scheme, diet is 
assessed In terms of calorie sources as 
recorded at national level In the FAO food 
balance database, and countries with similar 
diet structures are clustered together In one 
of six diet classes (Table 4.2, Map 4.3). Each 
class is named after the food product that 
best characterizes the diet (although this is 
not necessarily the principal calorie source). 

The countries In each food class tend to 
share broad demographic features. Rice 
countries (class 1) have high population 
densities, higher than average mortality rates 
and little diet diversification. Maize countries 
(class 2) generally have population densities 
near the world average and low mortality, 
especially Infant mortality. Wheat countries 
on average have low population density, but 
this masks serious land and water shortages 
in many. Class U countries, with a diet 
described as mixed 'mllk-meat-wheaf, 
include the worlds most highly developed 
nations, with fertility, mortality and popu- 
lation growth rates well below the global 
average. Millet countries (class 5) In general 
tend to have high population growth, high 
fertility and low life expectancy; the diet 
provides only a marginal surplus of energy 



Humans, food and biodiversity 45 



supplies over requirements. Diet class 6 
countries contain the hunnan populations 
most at risk from food Insecurity. The diet is 
characterized by roots and tubers, and on 
average does not provide basic energy 
requirements. These populations show high 
fertility, high mortality and a rapid grovirth in 
numbers. Poverty is vi^ldespread and the 
infrastructure weak, but there are significant 
reserves of under-exploited arable land. 

These major diet patterns have been used 
to help characterize the improvements to 
production and food supply that may be 
needed in order to meet the per capita energy 
needs of the population projected to exist 
In 2050. In terms of diet class, availability In 
class k could remain little changed, supply 
in classes 1, 2 and 3 would need to approxi- 
mately double, and classes 5 and 6 would 
need at least a fourfold increase. 

In some cases, food security may be 
better ensured by diversification rather than 
Increasing yields. The limitations of agri- 
culture based on uniform varieties and a high 
Input of fertilizers and pesticides become 
more acute for farmers who rely on poor 
resources or marginal environments. By 
growing diverse and locally adapted crops 
farmers can bring about greater security in 
food production and more efficient use of 
limiting resources. Many traditional agri- 
cultural systems manage to varying degrees 
a high diversity of both cultivated and wild 
food species (Table A. 31, and in a sense reduce 
the distinction between wild plants and 
domesticated crops. Strong selection pres- 
sures exerted by natural processes and 
humans over several millennia and wide geo- 
graphical areas have resulted In thousands of 
varieties within most crop species. 

High agricultural diversity can not only 
provide insurance against crop failure in 
difficult agricultural environments, but tends 
to have nutritional benefits. Transition to 
more uniform diets, with high Intake of fats 
and sugar, has resulted In declining nutri- 
tional status among numerous indigenous 
groups*"'^ Low dietary diversity Is associated 
with micronutrlent deficiency, a problem far 
more common than general protein-energy 
malnutrition, and particularly prevalent In 



children, pregnant women and breastfeeding 
mothers. Diverse cropping systems In back- 
yard and home gardens, whether rural or 
urban, can lead to direct Improvement in 
family nutrition and in some cases provide 
cash Income. Even a small mixed vegetable 
garden is capable" of providing 10-20 percent 
of the recommended dally allowance of 



Table 4.3 

Examples of diversity in 

agricultural systems 

Source; Multiple sources 





Brazil 


Kayapo 


Over 45 tree species planted for food or to attract game; 
86 varieties of food plants grown 


Ecuador 


Slona-Secoya 


Major staples: 15 varieties of manioc, 15 of plantain, 
9 of maize; pre-1978 traditional gardens yield 12 300 kilos of 
food or 8.8 million calories; 72% calories and U.8% protein, 
22.2% fats, 90.9% carbohydrates; po5t-1978 horticulture 
provides 67.8% calories, 10.2%, purchased 


Indonesia 


Java 


500 species in home gardens In a single village; i^^m 
ability to support 1 000 people per km' ^^H 


Indonesia 


West Sumatra 


6 main tree crops, many vegetables and fruit; 

53 species of wild plants also protected and harvested < 


Kenya 


Bungoma 


100 species of fruit and vegetables; 47% households ' 
collect wild plants, 49% maintain wild plants in gardens i 


Kenya 


Chagga 


Over 100 species produced in gardens 


l«lexlco 


Huastec 


Over 300 useful species found In managed forest plots \ 
called te'lom, 81 food species ' 


Mexico 


Migrant rural 
community 
In southeast 


338 species of plants and animals In home gardens, ' 
including 62 species of wild plants 


Papua New 
Guinea 


Gidra 


Approximately 54% calories and 82% of protein come 
from non-purchased sources (wild, sago, coconut gardens! 


Peru 


Bora 


22 varieties of manioc and 37 tree species are planted; 
118 useful species found in fallow fields 


Peru 


Santa Rosa 


168 species Identified in 21 home gardens 


Philippines 


Hanunoo 


System of intercropping with 40 crops in a single field; 
over 1 500 plants considered useful, of which 430 may be 
grown in swiddens 


Sierra 
Leone 


Gola forest 


Of food items 14% are hunted, 25% are from fallow land, 
8% are from plantations, 19% are from farm, swamp or 
garden, 21% are from streams and rivers, 13% is bought 
or given 


Ttialiand 


Lua 


1 1 varieties of food plants and 27 wild food plants are 
found in swidden fallows 



46 WORLD ATLAS OF BIODIVERSITY 



^^^^BNd. of 


No. of individuals 

1 


BlomaHlkUosT^^H 

Per km' Globally J 
of habitat -I^^H 


Per km' 
of habitat 


Globally 


Bacteria" 400 000 


7.84x10-' 
-1.18x10" 


4x10" 
-6x10" 


1.39x10' 
-2.14x10' 


7.06x10" 
-1.09x10= 


Coliembola" b 500 


2.4x10' 


2x10'= 


64 


5x10' 


Termites"" 2 760 


2.3x10' 


2.4x10" 


1400 


1.44x10" 


Antarctic krill" 1 


1.43x10' 


5x10" 


4.29x10= 


1.5x10" 


Birds" 9 946 


1 600 - 3 200 


2x10" 
-4x10" 






Eleptiants"" 2 


0.07-0.1 


4.26x10= 
-6.31x10= 


85-126 


5.34x10' 
-7.29x10' 


Great whales"" 10 


<0.01 


2.83x10' 
-3.6x10* 


52-65 


1.89x10" 
-2.33x10" 


Domestic livestock 
(excluding pets)" ca 1 5 








7.3x10" 


Humans 1 




6x10' 




3.9x10" 


Humans plus 
livestock 








11.2x10" 



Table 4.4 

Number of individuals and 

biomass, selected organisms 

Notes; Calculations based on 
data in sources cited after 
group names. Biomass is 
estimated dry mass 
standardized on 30% wet 
mass. Number of birds per 
km- estimated by dividing 
global bird population 
(2-4 X 10") by land area minus 
area of extreme desert, rock, 
sand and ice lapprox. 
125 million km'l. Estimates of 
abundance for Coliembola 
represent minimum global 
totals. Whale estimates 
presume a sex ratio of 1:1. 
Mean Asian elephant mass 
estimated at 3 500 kilos; range 
estimated at 500 000 km'. 
African elephant mass 
estimated at 4 250 kilos. 



protein, 20 percent of iron, 20 percent of 
calcium, 80 percent of vitamin A and 100 
percent of vitamin C. 

HUMAN NUMBERS AND IMPACTS 
Human population size 

Information on early human population 
numbers is based heavily on inference from 
circumstantial evidence, and remains on an 
uncertain footing even when written historical 
material becomes available in some abun- 
dance for the past few hundred years. 

Highly speculative estimates based on 
extrapolations of population densities of great 
apes=' and on studies of contemporary human 



hunter-gatherers== indicate that the global 
late Pleistocene human population may have 
been between 5 million and 10 million. It 
seems likely that any increase in human 
population up to then had been a result of 
increasing the total area occupied by the 
species, rather than by any major increase in 
population density in already occupied areas. 
Information on human population size in 
historic times is fragmentary, and populations 
lacking written records are likely to be 
inadequately represented. 

One analysis'"' distinguishes three main 
phases of population change. First is a cycle 
of primary increase in Europe, Asia and the 
Mediterranean brought about by the spread 
and further development of Neolithic 
agriculture, which appears to have allowed a 
great increase in population density. At the 
start of the Iron Age in Europe and the Near 
East, some 3 000 years ago, the world 
population may have been doubling every 500 
years, and the total probably reached 100 
million around this time or soon after. Growth 
appears to have slowed to reach near zero by 
around year 400, possibly because the limits 
of then current technology had been reached. 
After the Dark Ages in Europe, a second 
growth cycle began around the 10th century in 
Europe and Asia, with numbers rising to a 
peak of around 360 million during the 13th 
century, followed by a slight fall. The global 
population then increased slowly until the 
19th century, when an increasingly rapid 
rise began as a result of revolutionary 
developments in agriculture, industry and 
public health [Figure 4.1|. 

Crucially, the rate of global population 
growth peaked during the late 1960s: it was 
then at just over 2 percent per annum, but 
is now about 1.7 percent. The absolute 
increase per annum has also peaked; it was 
around 85 million more people per annum in 
the late 1980s and is now about 80 million. 



12 000 



11 000 



10 000 



9 000 



iOOO 



7 000 Years ago 



Humans, food and biodiversity 47 



Such trends suggest that the present global 
total of some 6 billion may not Itself double, 
as all previous totals have done. The medium 
variant of the current UN long-range forecast 
suggests the total in 2050 may be 9.3 billion". 
Although several countries in Africa have 
yet to shift to lower reproduction rates, thus 
making a further doubling quite possible, it 
may be that 'children born today may be 
thinking about their retirement at a time 
when the global population count will have 
stabilized - or even begun to decline". 

The exponential rise in abundance of a 
single species, to a position of global eco- 
logical dominance, in the sense of using a 
disproportionate share of natural resources, 
is without known precedent in the history of 
the biosphere. This was not achieved without 
significant, often adverse, effects on the 
environment, many of which stem from the 
agricultural activities required to maintain 
human numbers. There is no single species in 
which so many individuals, of such large body 
size, are distributed so widely over the planet. 
There appear to be few macroscopic species, 
the Antarctic krill Euphausia suberba being 
one of these few, in which the number of 
individuals approaches the size of the present 
human population. No animal of comparable 
size has a population remotely similar in 
number to that of humans. Biomass provides 
a measure of the way in which global net 
primary production or NPP Isee Chapter 11 is 
partitioned. If the standing crop of domestic 
livestock is added to the human biomass, 
amounting to around 1 1.2 x 10" kilos in total, 
then only the global biomass of bacteria as a 
whole Is higher (Table A.A]. 

The current human population Is unevenly 
spread across the land surface of the Earth. 
While some areas, such as most of Antarctica, 
the interior of Greenland, and hyperarid hot 
deserts, have no permanent human presence, 
In others human densities may locally reach 



an extreme of 1 000 Inhabitants per hectare 
le.g. In Calcutta and Shanghai!. Map i.A 
shows the current density of human popu- 
lations over the Earth, based on census 
counts within administrative units of varying 
size. Among the most striking features of 
this map are the large areas of very high 
population density in parts of China and the 
Ganges-Brahmaputra lowlands, also on Java, 
and the large area of high population density 
extending over most of Europe. 

Human activities have now made them- 
selves felt throughout the biosphere, but it 
might be expected that the degree of trans- 
formation of terrestrial landscapes would be 
related to ease of access and proximity to 
population centers. Settlements, ranging in 
size from villages to cities of many million 
Inhabitants, are connected by networks of 
paths, railways and waterways that allow the 
influence of humans to diffuse far beyond 
these settlements. 

Map 4.5 shows the results of a GiS-based 
analysis of the relative distance of points on 
the Earths surface from all such human 
constructions. Those points most remote can 
be given a high naturalness' value II. e. 
Ignoring other possible Impacts, they are likely 
to be least disturbed] and, conversely, points 
surrounded by a high density of human 
structures can be given a low value. Although 
human population density was not part of the 
analysis, there are, unsurprisingly, strong 
similarities between Maps A.A and 4.5. 



t6 



- 5 




5 000 



4 000 



3 000 



2 000 



1 000 



Figure 4.1 
Human population 

This graph shows the long 
period of many thousands 
of years during which the 
world human population 
remained small, followed by 
an exceptionally rapid rise 
to a total of 6 billion at 
present. 

Source: Data from McEvedy and Jones ; 
FAOSTAT database". 



48 WORLD ATLAS OF BIODIVERSITY 



Map 4.3 

FAO world diet classes 

A classification of country 
dietary patterns, based 
mainly on calorie sources. 
Eacti class is named after 
the foodstuff that best 
characterizes the diet. The 
classification of a few 
countries (e.g. Japan, 
Ivlalaysial appears 
anomalous, and some 
countries are not included 
in the classification. 

Source: Analysis by FAO . 



Diet class 




Class 1 Rice \ 

Class 2 Maize 

Class 3 Wtieat 

Class A fvliLk, meat, wheat 

Class 5 Millet, sorghium 

Class 6 Cassava, plantain, tare, yams 

Not assigned to a class 



The human share of global resources 

Agriculture is a set of activities designed to 
secure a greater and more reliable share of 
the energy and materials in the biosphere for 
the benefit of the human species, i.e. to divert 
an increased amount of available energy 
toward production of human bodies. Naturally 
occurring plants and animals are replaced by 
specially cultivated or bred varieties that can 
produce nutrients efficiently from available 
resources, and in a form that humans can 
conveniently use. The growth and persistence 
of these selected species is subsidized by 
humans. In less-developed agricultural sys- 
tems, this subsidy may be very small, perhaps 
just the removal of competition for light or 
grazing, i.e. plots are cleared or weeded and 
wild herbivores discouraged. Efficiency can be 
high but output is usually low. In western 




J!5>. 




industrialized agriculture the subsidy is 
enormous. Competitors, pests and predators 
are removed from vast areas (through the use 
of herbicides and pesticides!; fossil fuel is 
consumed to process, transport and apply any 
nutrients that limit production (nitrogenous 
fertilizer) and to store produce. Efficiency is 
high in some respects, e.g. use of space and 
labor, but much lower if all hidden costs of 
fossil fuel use and waste impact are 
considered. Output can be very high. 

The area of land devoted to agricultural 
production is now a significant proportion of 
the global land surface. Five thousand years 
ago, the amount of agricultural land in the 
world was negligible. There is no direct 
evidence from the greater part of this period 
on the rate of expansion of agricultural land. 
Useful historical data relate to the past few 



Humans, food and biodiversity 49 




hundred years, and this evidence suggests 
that about 250 to 300 million hectares (ha) of 
land globally were devoted to crops in 1700. At 
present, arable and permanent cropland 
covers approximately 1 500 million ha of land, 
with some 3 400 million ha of additional land 
classed as permanent pasture (this figure 
includes rangeland and vi^ooded land used 
for grazingl. This represents a nearly sixfold 
increase in cropped land over the past three 
centuries (Table i.5|. The current extent of 
cropland is represented in Map 5.2. All the 
cropland is used to produce domestic plant 
material, and much of the land classed as 
pasture, together with large parts of the 
grassland and open shrubland landcover 
classes in Map 5.2 (of which 'permanent 
pasture' is a subset] is used to produce 
domestic herbivore biomass. 



Most domestic herbivores are destined to 
become human biomass or to meet other 
human requirements, so can be counted as 
surrogate humans. The rise in livestock 
numbers has been accompanied by a 
decline in wild herbivores. For example, the 
extinct wild ox Bos primigenius of Eurasia 
and North Africa has been replaced in its 
former range by domestic cattle, of which 
approximately 1 360 million head exist in the 
world". Similarly, the American bison Bison 
bison was reduced from perhaps 50 million 
before European arrival on the continent to 



Cropland area 
(million hal 



Table 4.5 

Land converted to 

cropland 



Source: 1700-1950 estimates from 
Richards^'; 1980 and 1999 data from 
FAOSTAT database Icomplete time 
series from 1961 available from tfiis 
sourcel. 



265 



537 913 

m il. — Tft-ifc' -oc- , ■■ 



1 170 



1431 



1501 



50 WORLD ATLAS OF BIODIVERSITY 



Table 4.6 

Estimated large herbivore 

numbers and biomass in 

Mesolithic and modern 

Britain 

Source: After Yalden . 



under 1 000 at the end of the 19th century; 
although there has been significant recovery 
under management, it has been replaced 
over much of its former range by domestic 
cattle, of which 100 million head are present 
in the United States. 

Data on the fauna of the Bialowieza forest in 
Poland la forest remnant with populations of 
large wild herbivores and predators) have been 
used'' to estimate the possible number and 
biomass of wild herbivores in heavily forested 
Mesolithic Britain. These estimates were 
compared with current herbivore populations 
in Britain. Indications are that there are now ^40 
times more domestic cattle than there were 
wild ox and 20 times more domestic sheep 
than there were wild deer; the overall large 
herbivore biomass has increased by a factor of 
ten Isee Table A.6]. Similar values are likely 
to apply to other heavily populated industria- 
lized countries. Not only are domesticated 
mammals far more abundant than their wild 



mSi«kS4)^aWi86rii£^i»tti&^fiUrWi^ir.V/.7\ . 


><«;;is"-^*ij<v.. 


.'v..:~'">, '4H^^^H 


Mesolithic Britain 






Modem Britain ^^^^| 


ft 


No. 


Kilos 




No. Kilos S 








Sheep 


20 364 600 916 407 000 


Wild boar 










Sus scrota 


1 357 740 


108 619 200 


Pigs 


853 000 127 950 000 


Wild ox 










Bos primigenius 


99 250 


39 700 000 


Cattle 


3 908 900 2149 895 000 


Red deer 










Cervus elepbus 


1 472 870 


147 287 000 


Red deer 


360 000 54 000 000 


Roe deer 










Capreolus capreolus 


1083 810 


21 676 200 


Roe deer 


500 000 10 500 000 


Moose 










Akes alces 


67 490 


13 498 000 


Introduced deer 111500 5 017 500 








Feral sheep 










and goats 


5 700 256 500 


^^^H 






Domestic herbivore 


^^^M 






total 


25126 500 3194 252 000 


^^^* 






Wild herbivore 


'^^ 




^^^■a 


total 


977 200 69 774 000 




4 as; 1M^^ 




^^00 3 264 02400^^ 



relatives ever were, the latter are in many 
cases extinct or near extinction. 

As mentioned in Chapter 1, calculations at 
global level and in a study of one country 
[Austria! estimate the proportion of terrestrial 
global production appropriated or diverted for 
human use at 30-50 percent (depending on 
how much change beyond direct harvesting is 
incorporated). One approach to assessing the 
effects of energy appropriation has used the 
empirical relationship between energy and 
species number". There is much evidence 
suggesting that at a range of spatial scales 
and within different taxonomic groups, the 
diversity of species present in an ecosystem 
tends to be positively correlated with the 
amount of energy available [see Chapter 5). 
Accepting the empirical evidence for this 
relationship, it has been argued that the 
number of species present will decline if the 
amount of energy available for use declines. 
At global level, a conservative estimate using 
this relationship predicted that 3-9 percent of 
terrestrial species would be extinct or 
endangered by the year 2000". The evidence 
from the Austrian study'" is consistent with 
the species-energy relationship: the curve 
developed by Wright" predicts that with 41 
percent of potential NPP at country level now 
being appropriated by humans, 5-13 percent 
of species should have been extirpated from 
the country; in fact, 8 percent of birds and 7- 
14 percent of reptiles have been lost. 

The significance human resource appro- 
priation has for other species, and for 
biosphere function, in part depends on the 
extent to which the resources are limited in 
availability. Space clearly is limited, and it may 
be that the main effects of appropriation will 
be exerted through direct changes to land- 
cover, and diminished availability of material 
resources for wild species, rather than 
through a diversion in energy flow. 

SPECIES EXTINCTION AND HUMANS 
Tracking extinction 

Change in biological diversity has principally 
been assessed in terms of declining popu- 
lations, species, communities and habitats. 
There has always been special concern about 
extinction, because this is a threshold from 



Humans, food and biodiversity 51 



whicfi tfiere is no turning back. As discussed 
in Chapter 3, ttie extinction of species is 
natural and expected, and self-evidently ttiere 
fiave always been species at risk of extinction. 
It seems very likely, however, that recent and 
current extinction rates are considerably 
higher than would be expected without the 
influence of humans. 

For several reasons it is difficult to record 
contemporary extinction events with precision. 
The species involved may well be unknown. 
Even if they have been discovered and named, 
they may be too small to be noticed without 
special sampling procedures. The entire 
process of decline and eventual extinction 
may take place over many years or even 
centuries in the case of particularly long-lived 
organisms such as many trees. The near- 
terminal stages in the process of species 
extinction are unlikely to be observed. Where 
such observations have been possible, it is 
because the species has been destroyed in 
part by unusually extreme hunting pressure 
(e.g. the passenger pigeon Ectopistes 
migratorius] or extreme ecological events 
le.g. extinction of many native land snails in 
French Polynesia and Hawaii following 
introduction of the predatory snail Eugiandina 
rosea], and has been the subject of sufficient 
interest to be closely monitored. 

In other cases, positive evidence of 
extinction is lacking. Typically, many years 
elapse before sightings of a species become 
sparse enough to generate concern, and 
many more years are likely to pass before 
negative evidence li.e. failure to find the 
species despite repeated searches] accumu- 
lates to the point where extinction is the most 
probable explanation. In other words, unless 
circumstances are exceptional, monitoring of 
recent extinction events has a resolution limit 
measured in years or decades. This is why it is 
not possible to state with precision how many 
species became extinct in a given month, year 
or even decade, nor to predict exactly how 
many species, let alone which ones, are going 
to become extinct this year or decade or 
century. The search effort and chance both 
play a part in determining whether a species 
not seen for decades is rediscovered, as 
shown by the occasional new encounter with 




Standing out as some good news against a background of widespread species depletion, a 
small but significant number of species have been considered extinct but rediscovered after 
a gap of several decades. In some cases, prolonged directed searches have been carried out, 
but others have emerged by chance. Sometimes a 'rediscovery' is in part a consequence of 
new systematic work confirming the species status of long-neglected populations, e.g. 
differentiation of the pygmy mouse lemur Microcebus myoxinus from other mouse lemurs 
on Madagascar The coelacanth Latimeria is an extreme 'Lazarus taxon', being the living 
representative of an entire order (CoelacanthiformesI of early fishes thought to have become 
extinct some 80 million years ago. There are at least two population groups, one off 
southeast Africa [L. chalumnae] first discovered in 1938, and one in Indonesia (named 
L menadoensis] discovered in 1998. 

• Bavarian pine vole Microtus bavaricus: an alpine species believed lost when a hotel was 
built near its single known locality, but recently rediscovered nearby in the Austrian Alps. 

• Fiji petrel Pterodroma macgiltivrayi: known by one specimen collected in 1855, but 
regarded as extinct until a bird flew into a researcher's headlight one night in 1984. 

• Jerdon's courser Rbinoptilus bitorquatus: known by just two museum skins and last 
recorded in 1900, until rediscovered in 1986 in a patch of scrub forest in southern India. 

• Jamaican iguana Cyclura collei: believed to have gone extinct during the 19'lOs but 
rediscovered in 1990 in the remote Hellshire Hills. 

• The Cranbrook pea Gastroiobium lehmannii: endemic to Western Australia where last 
collected from the wild in 1918 and since listed as extinct; rediscovered in 1995. 



species once feared extinct (see Box 4.21. 
Hidden survivors are even more likely in plants, 
which may have propagules that can remain 
viable but unseen for extremely long periods. 
Extinctions in the recent past are likely to be 
recorded with significant accuracy either where 
circumstances favor preservation of hard 
remains in good number (e.g. in caves, potholes 
or kitchen middensi or where early naturalists 
recorded the fauna or flora with sufficient care 
that they set a firm baseline against which the 
composition of the modern biota may be 
assessed. The detailed record of bird extinc- 
tion in Hawaii is a result of the former 
circumstance, and the record of mollusk 
extinction on many islands a result of the latter. 



Table i.7 

Late Pleistocene extinct 

and living genera of large 

animals 

Source: Adapted irom Martin . 



^H[ Extinct wRhin Still 


Total 




^^^; last 100 000 extant 


genera 


of maximum 1 


Africa 7 42 
North Amenca 33 12 


49 
45 


14 No peak 

73 11 000-13 000 years ago 


South America 46 12 


58 


79 11 000-13 000 years ago 


Australia 19 3 


22 


86 ca 50 000 years ago 



52 WORLD ATLAS OF BIODIVERSITY 



Map U.U 

Human population density 

The relative density of 
human population based on 
census data relating to 
administrative units of 
various sizes. 

Source- CIESIN, gndded population of 
the world, version 2. data available 
online at 

http://sedac.cie5in, columbia.edu/ 
plue/gpw/ [accessed April 20021 



Population density 



High 



Low 



Early human impacts on biodiversity 

A wide range of factors affects the frequency 
of occurrence of particular species in the 
fossil record, of which the abundance of that 
species in life is only one. There is not 
necessarily therefore any direct relationship 
between the former and the latter, so that 
deducing past changes in abundance of 
species from the fossil record is a problematic 
exercise. Cataloging, though not dating, 
extinctions is rather less contentious, al- 
though even this may be problematic as 
evinced by the existence of so-called 'Lazarus' 
taxa Ithose presumed extinct that are 
rediscovered alive. 

One of the unusual features of the 
Quaternary period (the Pleistocene and 
Holocenel has been the disproportionately 
high extinction rates in the largest ter- 



J^ 




'-W' 



f^ 




'^^■J^- 



V, 




restrial species, particularly mammals and 
birds (Table 4.71. These species are gen- 
erally referred to as the 'megafauna', often 
defined as those with an adult mass of UU 
kilos or more, although the term has not 
been used consistently. 

The extinct American fauna include such 
well-known genera as the sabretooth cats 
Smilodon. giant ground sloths Eremotherium, 
glyptodonts Glyptotherium and mammoths 
Mammuthus, as well as a number of 
scavenging and raptorial birds, including the 
giant Teratornis and Cathartornis. Those in 
Australia include the marsupial equivalents 
of rhinoceroses (family Diprotodontidael 
and lions \Thylacoleo\, giant wombats 
(Phascolonus, Ramsayia and Phascolomis], 
the large emu-like Genyornis and the giant 
monitor lizard Megatania. In all, some 40 



Humans, food and biodiversity 53 



•'^^^ 




■.yv 



~^- 




species of tfie larger Australian land mam- 
mals, reptiles and birds became extinct 
across the entire continent about 46 000 years 
ago", approximately 10 000 years after the 
first known human colonists. 

These extinctions have been followed by a 
similar series on islands during the Holocene. 
On New Zealand, the moas (giant flightless 
ratite birds in the family Anomalopterygidael 
became extinct after the first humans came 
to the islands. On tvladagascar, two endemic 
hippopotamus species Hippopotamus lemerlei 
and H. madagascariensis, the elephantbird 
Aepyornis maximus, and a number of large to 
very large lemur species all appear to have 
died out 500 to 900 years ago. Similarly on 
the Caribbean islands, a number of large 
mammals, including several ground sloths 
(order Xenarthra, family Megalonychidael 



appeared to have survived until human 
occupation but to have died out at some point 
since then. 

The precise causes of all these extinctions 
have been the subject of endless debate, 
which centers chiefly on the role of humans. 
At one extreme lies the 'blitzkrieg' hypo- 
thesis, applied particularly to the apparently 
sudden collapse (i.e. over a few hundred 
years! of the North American megafauna at 
the hand of humans. At the other are those 
who maintain that in most, if not all, cases 
the impact of early humans was negligible 
and climate change, particularly increasing 
aridity, was the cause. 

Several 'eatures of the phenomenon seem 
to point persuasively to humans having played 
a pivotal role in most, if not all, of these 
extinctions. The most compelling is their 



54 WORLD ATLAS OF BIODIVERSITY 



Map 4.5 

Terrestrial wilderness 

The wilderness value of any 
given point is essentially a 
measure of remoteness 
from human influence, 
assessed on the basis of 
distance from settlement, 
access routes and 
permanent manmade 
structures. 

Source: GIS analysis by R, Lesslie lANUI. 
method developed for the Australian 
Heritage Commission. 



Wilderness level 



High 





Low 



timing. In each case the arrival of humans 
seems to have preceded the major spate of 
extinctions (in as much as these can be 
dated], with no or very few such extinctions 
having been recorded prior to human arrival 
(compare fvjap 4.1 and Table 4.7). The 
cumulative weight of this coincidence is 
difficult to counter, and is supported by 
population modeling. For example, new 
archeological evidence suggests that the first 
Polynesian settlements in New Zealand date 
from the late 13th century, and moas were 
becoming scarce by the end of the Hth. This 
evidence, coupled with a mathematical model 
of population and predation, indicates that all 
11 moa species were driven to extinction 
within 100 years of human arrival". Similarly, 
a computer simulation" of population 
dynamics of humans and large herbivores in 




North America accurately models mega- 
faunal extinction in accord with archeological 
evidence for significant human arrival around 
13 400 years ago and the first wave of 
extinction starting some 1 000 years later 

The situation is somewhat different in Africa 
and Eurasia, where megataunal extinctions 
were relatively few, and were spread out over 
the whole of the Pleistocene. In Africa the peal< 
was in the lower Pleistocene |21 genera 
extirpated from the region between 1.8 million 
and 700 000 years ago compared with nine 
between 700 000 and 130 000 years ago, and 
seven later than 130 000 years ago]. It is 
noteworthy that this more gradual, earlier and 
less extreme pattern of extinctions has typified 
the region where humans evolved. 

It is difficult to formulate an entirely 
climate-based model of extinction that can 



Humans, food and biodiversity 55 




account for tfiis asynctironicity - outside Africa 
and Eurasia, ttiese species survived a series of 
climatic cfianges at least as extreme as those 
they faced at the start of the Holocene. The 
recent studies of Genyornis newtoni and other 
megafauna in Australia"" seem to indicate a 
widespread and largely synchronous dis- 
appearance from a wide range of habitat types 
during a period of relative climatic stability, 
strongly implying that some other agent was 
responsible. It is also noteworthy that the 
fossil and archeological evidence indicates 
that, on continents at least, these extinctions 
were not matched by parallel extinctions of 
smaller species (for example, as far as Is 
known no Insect species at all became extinct 
in Europe In the entire Pleistocene'!. If climate 
change were the cause, it would be expected 
that these species would be at least as affected 



as larger ones as In general their opportunities 
for long-range dispersal and migration are 
much more limited. 

Even if humans are accepted as the major 
agents in these extinctions, evidence for the 
mechanisms Involved in most cases remains 
elusive. It seems likely that extensive use of 
fire, and direct hunting of a fauna that had 
evolved In the absence of humans and was 
therefore unlikely to recognize them as 
potential predators, may have been sufficient 
cause to exterminate the large herbivores. 
The large carnivores and scavengers may 
then have suffered population collapses 
owing to the disappearance of their prey base. 

Extinction in modern times 

From the relatively sparse evidence that is 
available, it appears that amongst animals 



56 WORLD ATLAS OF BIODIVERSITY 



Map 4.6 

Vertebrate extinctions 

since AD1500 

An indication of the number 
and former occurrence of 
recently extinct mammal, 
bird and freshwater fish 
species. Each of these three 
vertebrate classes is 
represented by a differently 
colored symbol, sized 
according to number of 
extinct species. Only 
extinctions that are fully 
resolved according to firm 
Committee on Recently Extinct 
Organisms ICREO) criteria 
are covered. The exception is 
Lake Victoria, indicated by a 
blue circle: some reports 
have suggested that up to a 
third of the 500 or so 
endemic fishes are extinct, 
but others note that 
available evidence is 
inconclusive, in many cases, 
including most islands and 
lakes, the position of the 
symbol indicates last record 
or core of former range. 
Where several species 
ranged more widely over a 
country, the symbol is 
positioned at the centre of 
that country. 

Source; Based on several sources; see 
Appendix k. 




P^^^'^' 



Y 



Vertebrates 


• 


Birds 


• 


Fishes 


• 


Mammals 


o 


Possibly-extinct 




Lake Victoria fishes 


Number 


of species 


m 


10-25 


• 


6-9 


• 


3-5 


• 


1-2 



more than 300 vertebrates, including at least 60 
and possibly more than 80 mammals, more 
than 120 birds and around 375 invertebrates, 
have become extinct during the past AOO years 
(see numerical summary in Table UZ and list of 
extinct vertebrates in Appendix 41. Data for 
plants are much more equivocal, owing in part 
to the uncertain taxonomic status of many 
extinct plant populations. One source" lists 
some 380 extinct plant taxa and a further 370 or 
so classified as extinct or endangered (these 
include a number of infraspecific taxa and 
a number that although believed extinct in 
the wild are extant, and sometimes abundant, 
in cultivation]. 

Because mammals and birds tend to be 
relatively well recorded, and because they leave 
recognizable macroscopic skeletal remains, it 
is principally among these groups that known 




ij^N) 







extinctions may be reasonably representative of 
actual extinctions. In these groups the known 
extinction rate over the past 400 years, based 
on data in Appendix 4, averages out at around 
20-25 species per 100 years. 

A crucial question then, is how this 
observed extinction rate compares with some 
hypothetical or expected background extinc- 
tion rate. It is of course impossible to derive 
such a rate from observation of the modern 
world, as this has already been highly 
modified by human activity The only reason- 
able comparison is thus with historical 
records, for which we must turn to the fossil 
record, discussed in Chapter 3. Although 
extinction rates have evidently been highly 
variable during the history of life on Earth, it 
seems that the average persistence time of 
species in the fossil record is around 4 million 



Humans, food and biodiversity 57 



'n 




^- 



'Ml 



•• 




years. If 10 million species exist at any one 
time, on tliis basis the extinction rate would 
amount to around 2.5 species annually. 
However, it is unclear fiov*/ representative the 
fossil record is of species as a whole. It is 
likely that many rare species or those with 
very restricted distribution never appear in it 
at all. These rare species may almost by 
definition be expected to be inherently more 
prone to extinction than the species that are 
recorded and may therefore be expected to 
have a lower persistence time. This would 
mean that average species duration was less 
- perhaps much less - than A million years 
and the actual extinction rate in geological 
time considerably higher than the rate 
observed in the fossil record. 

Applying a mean persistence time of 
A million years to birds and mammals land 



assuming some 10 000 species of the former 
and around 5 000 of the latter], the back- 
ground extinction rate would be around one 
species every 500 years and 1 000 years, 
respectively, so that current rates would be 
some 100 or 200 higher than background. 
Even if background rates in these groups were 



Mammals 

Birds 

Reptiles 

Amphibians 

Fisfies 

Insects 

Mollusks 

Other invertebrates 



83 
128 

21 
5 

81 

72 
291 

12 



I 



Table 4.8 

Numbers of extinct animal 

species according to lUCN 

Note: Alternative criteria used 
by the Committee on Recently 
Extinct Organisms ICREOI 
result in a lower number of 
mammals and fishes being 
regarded as certainly extinct 
than IS given in this table (see 
full list in Appendix 4). 

Source; Hilton-Taylor . 



58 WORLD ATLAS OF BIODIVERSITY 



Map 4.7 

Threatened mammal 
species 

Color represents number 
of globally threatened 
mammal species in each 
country in 2000. Pie charts 
represent the proportion 
of the mammal fauna 
assessed as threatened at 
the national level in a small 
sample of countries. This 
IS a highly generalized 
comparison because of 
differences in status 
assessment methods. 

Note: To reduce ambiguity 
Alaska (United States] has for 
the purposes of this map 
been assigned to the same 
class as adjacent Canada 
rather than the conterminous 
United States. 



Source: Global data from Hilton- 
Taylor , country data (rom selection ol 
national Red Data books. 





National status 

( A — % threatened 



ten times this, the currently observed 
extinction rates would still be 10-20 times 
those expected. It seems therefore that, even 
if a high background extinction rate is 
postulated, recent extinctions are still much 
higher than might be expected (the alternative 
explanation is that the background rate was 
higher still). This elevated rate is particularly 
noteworthy as the Holocene appears to have 
been a period of relative climatic and 
geological stability, in which extinction rates 
might have been expected to be low. 

Most known extinctions have occurred on 
islands, including 42 of the 61 resolved 
mammal extinctions 168 percent], and 105 of 
the 128 bird extinctions (82 percent]. Reasons 
for the former are probably twofold. First, 
island species do appear to be particularly 
extinction-prone, by virtue of their limited 




ranges and usually small population sizes, 
and also because they have often evolved in 
the absence of certain pressures (e.g. ter- 
restrial predators, grazing ungulates!; if faced 
with these pressures (usually through human 
intervention] their populations may collapse 
completely. Second, it is much easier to arrive 
at some certainty that a given species is no 
longer present on an island of limited extent 
than that it has disappeared completely from 
a continental range, where the limits of its 
range were probably uncertain to start with. 

Most known or probable continental 
extinctions have been among freshwater 
organisms, particularly fishes and moUusks. 
Ivlany freshwater biota appear to have the 
characteristics of island organisms, in that 
they have limited and highly circumscribed 
ranges and they are similarly often sensitive to 



Humans, food and biodiversity 59 




ao Tome 
d Principe 



■^f^m-f 



rk 



Soutln Africa 



Australia 





external pressures (e.g. introduction of preda- 
tory fisti species, complete fiabitat destruction 
througfi drainage or dam construction). 

Figure 4.2 represents the number of 
accepted extinction events among mammals, 
birds and fisfies in eacfi third of the five 
centuries since AD1500 (see also Map 4.6|. 
The information in Appendix 4 allows some 
elements of this broad global picture to be 
disentangled: 

• early extinctions among the remaining 
archaic fauna of large islands, notably mam- 
mals on Hispaniola, Cuba and Madagascar; 

• somewhat later extinctions, especially 
among ground birds, on many isolated 
small islands, such as St Helena, the 
Mascarenes and others, during European 
colonial consolidation; 



50 



AO 



30 



I 20- 



Fishes 

Birds 

Mammals 



20 - -m 

i 



Figure 4.2 

Vertebrate extinctions by 

period since AD1500 

This graphic is derived from 
information in the 'Period' 
column of the list of extinct 
vertebrates given in 
Appendix 4. 'Early' refers to 
the first four decades of a 
century, 'mid' to the next 
three decades, 'late' to the 
final three decades. Only 
the extinction events 
regarded by Committee on 
Recently Extinct Organisms 
(CREOI criteria as 'resolved' 
are shown, and only the 
three groups recently 
assessed are represented. 

Source: Derived from Appendix 4. 




early mid late early mid late 
16th century I 17th century 



early mid late early mid late early mid late 
18th century I 19th century I 20th century 



60 WORLD ATLAS OF BIODIVERSITY 



^^^■l^^^pl 


f No. of 


nSH 


|P^% breeding 


^ 


breeding 


globally 


species 


1^^^^^^^ 


bird species 


threatened 


threatened ^^ 


Continental countries 








Brazil 


U92 


113 


8 


China 


1100 


73 


7 


India 


923 


68 


7 


Colombia 


1695 


77 


5 


■ Peru 

Small islands 


1538 


73 


5 








French Polynesia 


60 


23 


38 


Solomon Islands 


163 


23 


U 


Mauritius (and Rodrigues) 


27 


9 


33 


Sao Tome and Principe 


63 


9 


U 


L Saint Helena and dependencies 


53 


13 


25 



Table A.9 

Island diversity at risk: 

birds 

Note: The first five rows are 
the continental countries 
(Indonesia and Philippines 
excluded! with most 
threatened bird species; the 
other five rows are the small 
island groups with most 
threatened bird species. 

Source: BirdLife International . 



• increasing evidence of 20th century 
extinctions among fisfies in many countries, 
and among marsupial mammals on the 
Australian mainland, with continuing losses 
of island species everywhere. 

The apparent reduction in extinction rate from 
the late 19th century onwards may be in part 
attributed to management action designed to 
maintain highly threatened species, and there 
is indeed good evidence that populations of 
a small number of target species have 
recovered significantly, but it is sure also to 
reflect the difficulties of observing and 
documenting extinction events. As mentioned 
above, many years may pass before a 
possibly extinct species can be treated with 
certainty as extinct, and the declining 
extinction rate toward the end of the 20th 
century is most probably in part an artifact 
of this monitoring process. 

Clearly, in view of our very incomplete 
knowledge of the worlds species, and the fact 
that only a minute proportion of living species 
are being actively monitored at any one time, 
it is extremely likely that more extinctions are 
occurring than are currently known. Indeed, 
most predictions of present and near-future 
extinctions suggest extremely high rates. 
Most are based on combining estimates of 
species richness in tropical forest with 
estimates of rate of loss of these forests, and 



predict species extinction on the basis of the 
general species-area relationship (which 
predicts a decline in species richness as area 
declines, see figure 5.1). It is widely believed 
that the great majority of all terrestrial 
species occur in tropical forests, and most of 
these species will be undescribed arthropods 
(notably beetles). At present rates of forest 
loss, it has been predicted that between 2 and 
8 percent of forest species will become 
extinct, or committed to extinction, between 
1990 and 2015'"'. Depending on whether 
higher or lower estimates of tropical species 
richness are used, extinction at this rate could 
entail loss of up to 100 000 species annually. 
As a cautionary note, it should be observed 
that few extinctions to date have actually been 
recorded in continental tropical moist forests, 
although monitoring species in these habitats 
presents great difficulty. 

Threatened species assessment 

Except for species lost as a result of random 
environmental factors, extinct species must 
largely be drawn from a pool of species that 
could be assessed as in decline or at risk, all 
of which face eventual extinction if negative 
trends or threats to their populations are not 
reversed. Various national and other programs 
have developed methods to assess the relative 
severity of risks faced by species, and to label 
species with an indicative category name. 
Conservation activities can then be prioritized 
on the basis of relative risk, taking account of 
other relevant factors, such as feasibility, cost 
and benefits, as appropriate. 

The system" developed by the Species 
Survival Commission of lUCN-the World 
Conservation Union (lUCN/SSC) and collabo- 
rators in conjunction with its Red Data Book" 
and Red List Programme" has been designed 
to provide an explicit and objective frame- 
work for assessment of extinction risk, and to 
be applicable to any taxonomic unit at or 
below the species level, and within any 
specified geographical or political area. To be 
categorized as threatened, any species has to 
meet one of five sets of criteria formulated to 
permit evaluation of all kinds of species, with 
a wide range of biological characteristics. 
The criteria are defined on population 



Humans, food and biodiversity 61 



"W/j: 



reduction, population size, geographic area 
and pattern of occurrence, and quantitative 
population analysis. 

The size and connectedness of different 
populations of a species influence the like- 
lihood of its survival. In general, small Isolated 
populations will be more sensitive than larger 
connected ones to demographic factors (e.g. 
random events affecting the survival and 
reproduction of individuals! or environmental 
factors (e.g. hurricanes, spread of disease, 
changes in the availability of foodl. Islands tend 
to have a much higher proportion of their biota 
at risk than continental countries because they 
start with many fewer species, all or many of 
which face the risks associated with a small 
range size (see Table 4.9 for an example using 
bird data). Biogeographic theory based around 
the species-area relationship, supported by 
much empirical evidence, predicts that each 
'habitat island' created by fragmentation 
of a continuous habitat area will come at 
equilibrium to contain fewer species than 
previously. Human activities everywhere tend 
to promote fragmentation of natural and often 
species-rich habitats (e.g. primary tropical 
forest or temperate meadow grassland] and 
the spread of highly managed species-poor 
habitats (e.g. eucalypt plantations or cereal 
croplands!. As a result, many species occur 
in just the kind of fragmented pattern that 
Increases the risk of extinction. 

Recent declines 

Reduction in population numbers, or com- 
plete loss of a species from a site or an 
Individual country (often termed extirpation! 
are far easier to observe than global species 
extinction, and appear liable to occur 
wherever humankind has modified the en- 
vironment for its own ends. The conservation 
status of most species is not known in detail, 
and this certainly applies to the many million 
as yet undescrlbed species, but two large 
animal groups - the mammals and birds - 
have been comprehensively assessed and 
may be representative of the status of 
biodiversity in general. Approximately lU 
percent (1 130] of the world's mammals and 
12 percent (1 183] of the worlds bird species 
are regarded on the basis of lUCN/SSC 



criteria as threatened (see Table 4.10]. 
Proportions are a great deal lower in other 
vertebrates, but none of these has been 
assessed fully. Empirical observations such 
as these give sufficient grounds for serious 
concern for biodiversity maintenance, regard- 
less of any hypotheses that have been pro- 
posed regarding the future rate of extinction. 
Interestingly, the ratio of threatened mam- 
mals to threatened birds is near 1:1, as Is 
the ratio of recorded, recently extinct mam- 
mals to birds, giving some indication that 
threatened species categories may be a 
reasonably reliable indicator of proneness to 
extinction, and also that mammals may as a 
group be somewhat more susceptible than 
birds to extinction. 

Countless other species, although not yet 
globally threatened, now exist in reduced num- 
bers and fragmented populations, and many of 
these are threatened with extinction at national 
level. The significance of loss of diversity at 
gene level Implied by loss of local populations 
of species Is not clear, although It has been 
argued that loss of resilience in response to 
environmental change is Inevitable. 



Table 4.10 
Threatened species 

Notes: Includes species 
assessed as globally 
threatened and assigned to 
categories 'critically 
endangered', 'endangered' 
or vulnerable' under lUCN 
criteria. Ivlosses not included 
here; most plant taxa listed 
were assessed for the World 
list of threatened trees'" in 
1998. 



Source: Hilton-Taylor ; BirdLife 
Internalional . 



^^^r 


No. of 


Approx. 


Threatened 


%oftotafl|| 


^^^^^K 


species In 


%of 


species 


In group ' 1 


^^ 


group 


group 
assessed 




threatened | 


Vertebrates 










Ivlammals 


U30 


100 


1 130 


24 


Birds 


9 750 


100 


1186 


12 


Reptiles 


8 002 


<15 


296 


4 


Amphibians 


4 950 


<15 


U6 


3 


Fisties 


25 000 


<10 


752 


3 


Invertebrates 










Insects 


950 000 


<0.01 


555 


0.06 


Mollusl<s 


70 000 


<5 


938 


1 


Crustaceans 


ilOOOO 


<5 


iOB 


1 


Others 






27 




Plants 










Gymnosperms 










(Conlferoptiyta, Cycadopfiy 


a. 








Ginkgophytal 


876 


72 


141 


16 


Angiospernis 










(Anthopfiyta, flowenng 










plants] 


250 000 


<5 


5 390 


2 



62 WORLD ATLAS OF BIODIVERSITY 



Map 4.8 

Critically endangered 

mannmals and birds 

The general distribution of 
almost all the 362 mammal 
and bird species categorized 
as 'critically endangered', 
the highest risk category, in 
2000. Each circle represents 
one distribution record; 
some species are known 
from a single point locality, 
others are represented by a 
cluster of localities. On this 
map, a high density of 
symbols can represent many 
records of a single species, 
or single records of many 
separate species. Red and 
blue symbols represent 
mammals and birds, 
respectively 

Source: Mammals; species selection 
based on Hilton-Taylor* . tjtstrtbution 
researcti and mapping by UNEP-WCf^lC; 
birds: spatial data provided by BirdLife 
International (February 20021, further 
information in BirdLife International . 



^^!> 



Critically endangered 
o Mammals 

o Birds 



At global level, most of the species 
assessed as threatened are terrestrial forms 
(Table 4.111. The preponderance of terrestrial 
species is because the great majority of 
mammals and birds are terrestrial, and 
little or nothing is known of the population 
status of most aquatic species in most 
groups. Where significant numbers of aquatic 
species have been assessed, e.g. among 
crustaceans and mollusks, the proportion of 
threatened aquatic species rises markedly. 
Among fishes, the high number of freshwater 
species doubtless in part reflects the general 
lack of data on marine species, but to some 
extent indicates relative risk - many fresh- 
water species being restricted to small and 
isolated habitat patches. 

Evidence of the vulnerable nature of 
freshwater habitats and the risk faced by 



t' \ 



■■» 



\ 



*1 



'^ -^ I, 






many aquatic groups is accumulating. For 
example, in the United States, freshwater 
groups are considerably more threatened 
than terrestrial groups (specifically, nearly 70 
percent of the mussels, 50 percent of the 
crayfish and 37 percent of the fishesi". 

Forest is an important habitat for a high pro- 
portion of the threatened terrestrial verte- 
brates. Among birds, for example, about 70 
percent of the 1 186 species assessed as 
threatened in the year 2000 occur in forest, and 
25 percent in grassland, savannah and scrub 
habitats'". Of the threatened forest birds, 41 
percent occur in lowland moist forest and about 
35 percent in montane moist forest™. Among 
the 515 mammals regarded as threatened in 
2000 that were assigned to a habitat category, 
33 percent occur in lowland moist forest and 22 
percent in montane formations". 



Humans, food and biodiversity 63 







if 

:- %. 



Numbers of globally threatened species 
can be mapped at country level le.g. Map 4.7, 
mammals). Maps of tfiis kind provide at best a 
broad overview of ttie occurrence of sucti 
species, shown as direct numbers or as a 
proportion of the total number in that group in 
the country. As with other country-level 
biodiversity analyses, this information could 
be used to help focus efforts to slow or 
reverse biodiversity decline. Data with 
improved spatial resolution (although plotted 
in a highly simplified way! are shown in Map 
i.8, representing the individual ranges of all 
the mammals and birds assessed as 'critically 
endangered'. BirdLife International has gen- 
erated individual distribution maps of all 
threatened birds. Among other applications, 
these are being used to develop the first-ever 
world map of all the threatened species in 



% 



an entire major group of organisms. A 
preliminary version of this, as a global density 
surface, is shown in Map 4.9. Being geo- 
referenced rather than country based, and 
at relatively fine spatial resolution, this has 
considerable potential to focus bird con- 
servation efforts effectively. 



Table 4.11 

Number of threatened 

animal species in major 

biomes 

Notes: Counts include globally 
threatened species tabulated 
in each biome (lUCN 
categories 'critically 
endangered', 'endangered' or 
vulnerable'!. Only mammals 
and birds have been 
comprehensively assessed. 
Some species, e.g 
amphibians and migratory 
fishes, are counted in more 
than one biome row. Plants 
not tabulated; almost all 
assessed as globally 
threatened are terrestrial. 
Inland water includes saline 
wetlands, cave waters, etc., as 
well as freshwaters. 

Source: Hilton-Taylor . 



■ 








Animal gn 


■ 


11 




m 




J3 






in 

e 




3 






■■ 


n 


E 
E 


-0 


'1. 


:5 

a. 


lA 
01 




tn 

Qt 








m 








in 




<n 




^»..me.ype 


.2 


I 


OQ 


Q£ 


< 


LU 





1 


m 


Marine 


315 


25 


105 


9 





163 








13 


Inland water 


1932 


31 


78 


111 


131 


627 


409 


125 


420 


Terrestrial 


3 627 


1 111 


1 U4 


283 


U3 








438 


508 

■'-■ 1 



44 WORLD ATLAS OF BIODIVERSITY 



Map 4.9 

Threatened bird species 

density 

BirdLite International tias 
developed digital range 
maps of all threatened bird 
species. This map is based 
on distribution data for the 
1 186 species assessed as 
threatened in 2000. The 
data are plotted as a global 
density surface, representing 
the number of species 
potentially present in each 
location. The map is the 
first such treatment of any 
large group of threatened 
species. 

Source; Density surface provided by 
BirdLife Inlernational, further 
information on Iflreatened birds in 
BirdLife International . 



Threatened bird species 
density 

High 



Low 



Proximate causes of recent declines 

The continuing conversion of natural habitats 
to cropland'' and other uses typically entails 
the replacement of systems rich in bio- 
diversity with monocultures or systems poor 
in biodiversity. Habitat modification, from 
agricultural conversion and a variety of other 
causes, is in general the most important 
factor acting to increase species' risk of 
extinction. Among species assessed as 
globally threatened in 2000", habitat 
modification is the principal threat affecting 
more than 80 percent of the mammals, birds 
and plants; it is similarly predominant in 
several other major groups, notably in 95 
percent of threatened bivalve mollusks, and is 
the main cause of loss in 75 percent of extinct 
freshwater fishes". 

A second major source of biodiversity loss 




"'^^ 






( 






IS the widespread introduction of species 
outside their natural range where they 
typically induce change at the community and 
ecosystem level. The effects of alien species 
are especially pronounced in closed systems 
such as lakes and islands. Introduced 
species, such as rats and cats, are cited as a 
cause of extinction in nearly 40 percent of the 
approximately 200 species where cause could 
be attributed; the majority were island 
forms". Seven endemic snails in French 
Polynesia have been extirpated following the 
late 1970s introduction of a carnivorous snail 
species lEuglandina rosea] intended to 
control another introduced species Ithe giant 
African snail Achatina fulica], itself an 
agricultural pest. Accidental introduction of 
the brown tree snake Boiga irregularis to 
Guam in 1968 led to decline of the entire 



Humans, food and biodiversity 65 




m>^''^^^ 






f 




.r*7 A. 



^7 



avifauna, of wfiicfi one species is now tfiougfit 
extinct and one is extinct in the wild'". 
Introduction of tfie Nite perch [Latss 
nitoticus] to Lal<e Victoria has contributed to 
the decline or extinction of nearly 200 native 
and endemic cichlid fishes. Overall, Intro- 
duction events probably number in the 
thousands at the global level"and, although 
cases involving animals have been cited to 
illustrate the scope of the problem, intro- 
duced plants are in many places as pervasive 
and damaging. 

High trade demand for certain species and 
products, whether for international markets 
consuming hardwoods, sea fish, live animals, 
and plants and derivatives, or local markets 
consuming commodities such as bushmeat 
or turtle eggs, can readily push exploitation 
beyond the production capacity of the 



resource. The direct impact of hunting, 
collecting and trade Is the second most 
important threat category among globally 
threatened mammals and birds, with around 
35 percent In each group affected". 

Rapid environmental change, such as that 
associated with El Nino Southern Oscillation 
(ENSOl events, can have significant impacts 
on natural habitats. For example, in 1997-98 
climate fluctuation associated with El Nlfio 
was implicated In the persistence and spread 
of fires in Brazil, Indonesia and elsewhere: 
an estimated 1 million hectares of savannah 
woodland burned in Brazil and a similar 
area of forests in Indonesia were affected 
by fire. The effect of events of this type will 
be multiplied many times wherever habitats 
are already fragmented and species are 
depleted. 



66 WORLD ATLAS OF BIODIVERSITY 



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7A Juma, C. 1989. Biological diversity and innovation: Conserving and utilizing genetic 

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75 Alvarez-Buylla Roces, M.A., Lazos Chavero, E. and Garcia-Burrios, J.R. 1989. 
Homegardens of a humid tropical region in South East Mexico: An example of an 
agroforestry cropping system in a recently established community Agroforestry Systems 
8: 133-156. 

76 Alcorn, J.B. 1984. Development policy, forests and peasant farms: Reflections on Haustec- 
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77 Davies, A.G. and Richards, P. 1991. Ram forest in Mende life: Resources and subsistence 
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report to the Economic and Social Committee on Overseas Research (ESCORl, UK 
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78 Michon, G., Mary, F. and Bompard, J. 1986. Multistoned agroforestry garden system in 
West Sumatra, Indonesia. /Igroforesfry Sysfems 4[4|: 315-338. 

79 Kunstadter, P., Chapman, E.C. and Sabhasri, S. 1978. Farmers in the forest: Economic 
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80 Padoch, C. and de Jong, W. 1991. The house gardens of Santa Rosa: Diversity and 
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81 Denevan, W.M, and Treacy, J.M. 1987. Young managed fallows at Brillo Neuvo. In: 
Denevan, W.M. and Padoch, C. (edsl. Swidden-fallow agroforestry in the Peruvian Amazon. 
Advances in Economic Botany 5, pp. 8-46. New York Botanical Garden, New York. 

82 Ohtsuka, R. 1993. Changing food and nutrition of the Gidra in lowland Papua New Guinea. 
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83 Conklin, H.C. 1954. An ethnoecological approach to shifting agriculture. Transactions of 
the New York Academy of Sciences 17: 133-142. 

84 Posey, D.A. 1 984. A preliminary report on diversified management of tropical forest by the 
Kayapo Indians of the Brazilian Amazon. Advances in Economic Botany 1: 1 12-126. 

85 Vickers, W.T. 1993. Changing tropical forest resource management strategies among the 
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Biocultural interactions and applications to development. Man and the Biosphere Series. 
Volume 13. UNESCO and Parthenon Publishing, Paris and Carnforth. 

86 Michon, G. 1983. Village forest gardens in West Java. In: Huxley, P. led.l Plant research 
and agroforestry, pp. 13-24. International Centre for Research in Agroforestry, Nairobi. 

87 Whitman, W.B., Coleman, D.C. and Wiebe, W.J. 1998. Prokaryotes: The unseen majority 
Proceedings of the National Academy of Sciences 95: 6578-6583. 

88 Rusek, J. 1998. Biodiversity of Collembola and their functional role in the ecosystem. 
Biodiversity and Conservation 7: 1207-1219. 

89 Zimmerman, P.R. et al. 1982. Termites: A potentially large source of atmospheric 
methane, carbon dioxide, and molecular hydrogen. Science 218: 563-565. 

90 Nicol, S. Time to krill? Antarctica Online: The website of the Australian Antarctic Program: 
http://wv\nA/.antdiv.gov,au/science/bio/issues_krill/krilLfeatures.html 

91 Gaston, K.J. and Blackburn, TM. 1997. How many birds are there'' Biodiversity and 
Conservation 6: 615-625. 

92 Said, M.Y. et al. 1995. African elephant database 1995. lUCN-the World Conservation Union, 
Gland. 



70 WORLD ATLAS OF BIODIVERSITY 



93 SantiapiUai, C. 1997. The Asian elephant conservation: A global strategy. Gajah. Journal of 

the Asian Elephant Specialist Group 18: 21-39. 
9A Nowak, R.M. 1991. Walker's mammals of the world. 5th edition. Volunne II. Johns Hopkins 

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FAQ Species Identification Guide. United Nations Environment Programme, Rome. 

96 Kemf, E. and Philips, C. 1995. Whales in the wild. WWF Species Status Report, WWF - 
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97 Oldfield, S., Lusty, C. and MacKiven, A. 1998. The world list of threatened trees. World 
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98 Harrison, I.J. and Stiassny, M.L.J. 1999. The quiet crisis: A preliminary listing of the 
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(ed.l Extinctions in near time: Causes, contexts, and consequences, pp.271 -332. Kluwer 
Academic/Plenum Publishers, New York. 



Terrestrial biodiversity 71 



5 



Terrestrial biodiversity 



TERRESTRIAL ECOSYSTEMS EXTEND OVER LITTLE MORE than one quarter of the 
Earth's surface but they are more accessible than aquatic habitats and so are far 
better known. The land supports fewer phyla than the oceans but most of the global 
diversity of species, and is characterized above all by an extensive cover of vascular plants, 
with associated animals and other groups of organisms. 

Forest and woodland ecosystems form the predominant natural landcover over most of 
the Earth's surface. These systems generate around half the terrestrial net primary 
production, and forests in the tropics are believed to hold most of the world's species. 
Approximately half the area of forest developed in post-glacial times has since been cleared 
or degraded by humans, and the amount of old-growth forest continues to decline. 

Grassland, shrubland and deserts collectively cover most of the unwooded land surface, 
with tundra on frozen subsoil at high northern latitudes. These areas tend to have lower 
species diversity than most forests, with the notable exception of Mediterranean-type 
shrublands, which support some of the most diverse floras on Earth. 

Humans have extensively altered most grassland and shrubland areas, usually through 
conversion to agriculture, burning and introduction of domestic livestock. They have had less 
Immediate impact on tundra and true desert regions, although these remain vulnerable to 
global climate change. 



THE TERRESTRIAL BIOSPHERE 
Land and water on the Earth's surface 
Land extends over nearly 150 million square 
kilometers (km*) or about 29 percent of the 
total surface of the planet, the remainder 
being covered by oceans. With the continents 
in their present position, more than two thirds 
of the land surface is in the northern 
hemisphere, and the area of land situated in 
the northern hemisphere above the Tropic of 
Cancer slightly exceeds that in the rest of 
the world put together (Table 5.1 1. A small 
proportion of the total land area is occupied by 
inland water ecosystems: takes and rivers 
cover around 2 percent and swamps and 
marshland a similar amount. About half of 
the land surface, approximately 52 percent, 
is below 500 meters (ml in elevation, and 
the mean elevation is 840 m. A minor but 
significant proportion of the land surface is 
mountainous in nature, with alpine land- 



scapes tending to replace trees above 
1 000 m at higher latitudes and above 3 500 m 
in the tropics. 

The land as an environment for living 
organisms 

The environmental conditions prevailing in 
any given place determine what kinds of 
organism can live there. Relevant environ- 
mental conditions include various aspects 
of the physical environment, and the other 



Table 5.1 

Global distribution of land 

area, by latitude bands 



Region 




Land area '^^H 
(million km'l | 


Norttiern tiemisphere 


North of Tropic of Cancer 


74 




Equator nortti to Tropic of Cancer 


26 


Souttiern hemisptiere 


Equator south to Tropic of Capricorn 


23.5 




Soutti of Tropic of Capricorn 


23.5 




(including Antarctica! 





72 WORLD ATLAS OF BIODIVERSITY 



Most terrestrial organisms 
occupy an environment 
where water loss is a 
constant threat. 



species that any given species directly or 
indirectly interacts with. These interactions 
may occur through a range of mechanisms, 
including competition, predation, symbiosis, 
mutualism and parasitism. For many living 
species, interactions with humans are now a 
significant feature of their environment. 

The most fundamental distinction, at least 
for macroscopic organisms, is between ter- 
restrial and aquatic environments. All living 




organisms require water because the basic 
processes of life take place in the aqueous 
medium found in cells. However, most 
terrestrial organisms, in contrast to aquatic 
ones, occupy an environment where water 
loss is a constant threat. The anatomical and 
physiological solutions to this problem are 
many and varied, including epidermal and/or 
cuticular layers with reduced permeability to 
water, water storage tissues, and metabolic 
or behavioral processes that conserve water 
However, the greater the risk of dehydration, 
the more carbon and energy are needed for 
water conservation mechanisms, leaving less 
available for other adaptations. Thus, life 
tends to occur at low density and at low 
diversity in hyperarid environments, whether 
exceptionally hot or exceptionally cold. 

There are a number of other extremely 
important differences between air and water 
as a medium for living things. It takes far less 
energy (about 500 times lessl to raise the 
temperature of a given mass of air by 1 degree 
Celsius than to heat the same mass of water by 
the same amount, and water conducts heat far 



more rapidly than air As a result, while aquatic 
organisms are buffered against rapid fluc- 
tuation in their surroundings, terrestrial 
organisms can be subjected to wide extremes 
in temperature, corresponding to daily and 
seasonal variation in insolation. 

The air surrounding terrestrial organisms is 
much less dense (about 800 times lessl than 
water, and so land organisms must support 
themselves against the full effects of gravity, 
but are not subject to the large forces exerted 
on aquatic organisms by moving water 

Oxygen is freely and uniformly available in 
the atmosphere, but in water is far less 
concentrated and much more variable in time 
and space. On the other hand, mineral 
nutrients dissolve readily in water and it is 
possible for aquatic organisms to extract 
nutrients directly from their immediate 
surroundings (although in many aquatic 
habitats such nutrients are only present in 
low concentrations!. On land, in contrast, 
mineral nutrients occur in soil where their 
distribution and concentration are spatially 
variable, and they are directly accessible 
primarily to microorganisms, plants, fungi 
and algae. 

Global variations in terrestrial habitats 

The wide variations in water availability and 
temperature regimes prevailing on the land 
surface interact in often complex ways with a 
number of other factors, including geology, 
soil, terrain, wind, fire regimes and human 
activities, to generate the immense range of 
environments apparent on the Earth's land 
surface. Classification of these into a manage- 
able system is a major problem in biology, 
one not merely of theoretical interest but of 
considerable importance in the management 
and conservation of the biosphere. The 
problem arises largely from a need to divide 
the natural environment into a series of 
discrete bounded units for the purposes of 
mapping, measuring and monitoring habitats, 
whereas the world in reality appears to form a 
highly variable continuum. 

Where gradients exist between different 
physical regimes (particularly of water and 
temperature], habitat types tend to intergrade 
imperceptibly, and it is impossible without 



Terrestrial biodiversity 73 



^^j 



arbitrary definitions to distinguish, for example, 
grassland with a few trees from open woodland 
with grass ground cover. Even broad categor- 
ies, such as 'forest' or 'wetland', inevitably 
require arbitrary limits to be set, e.g. for the 
density of tree cover necessary before an area 
can be called a forest, or for the duration of 
flooding necessary before an area can be 
classified as a wetland rather than a terrestrial 
system. In such circumstances, it is important 
to keep in mind the inherent variability of 
ecosystems, rather than attach undue signifi- 
cance to their labels. 

Vegetation and chlorophyll 

The global distribution of actively growing 
vegetation can be visualized without classi- 
fication or criteria relating to structure, to 
physiognomy or to species composition. 
Advanced very high resolution radiometer 
lAVHRRI satellite sensors measure the 
reflectance of vegetation, primarily of the 
green photosynthetic pigment chlorophyll, in 
the visible and the near infrared part of the 
spectrum. On land this can be interpreted as 
broadly equivalent to the density and vigor 
of green plant growth, represented as the 
normalized difference vegetation index 
(NDVII. This is plotted, aggregated over one 
year, in Map 5.1. This reveals a clear 
distinction between areas rich in standing 
growth of plants, whether cropland or natural 
vegetation, and areas where standing plant 
growth is sparse or absent - these are 
essentially the drylands and rangelands of 
conventional landuse classifications. 

Spatial variation in chlorophyll density is 
only indirectly related to variation in primary 
production levels; net primary production 
depends further on soil and climate conditions 
and community dynamics within ecosystems 
(see Chapter II. So, while Map 5.1 shows high 
chlorophyll in both tropical and high latitudes, 
net primary production is far higher in the 
former than the latter, where it is restricted 
mainly by seasonally unfavorable climatic 
conditions (compare with Map 1 .21. 

Landcover and ecosystems 

Landcover classification is more directly con- 
cerned with differences in the physical aspects 




of ground cover, mainly for landuse planning 
and management, than with biodiversity or the 
community aspects of vegetation cover Many 
current landcover maps (Map 5.21 are based 
on interpretation of remote-sensing data that 
have the virtue of being quantitative in nature 
and available in time series suitable for 
monitoring applications. Because the source 
data are typically interpreted in terms of 
a classification system developed to take 
account of conditions on the ground, landcover 
maps are subject to some of the definition 
problems mentioned above, no matter how 
generalized the categories are at the highest 
levels (e.g. 'forest', 'cropland', 'urban']. 

One of the most useful ecological 
distinctions to be made is between areas with 
extensive or significant tree cover, and areas 
with few or no trees. Terrestrial plant growth 
IS favored by high soil water availability and 
relatively elevated temperature during all or a 
major part of the year Trees tend to be the 
main plant growth form in such conditions, 
and forest or woodland the main vegetation. 
Conversely, primary production is strongly 
limited by a shortage of soil water. Grasses 
and low shrubs tend to be the main plant 
growth forms in such dryland regions and, 
where vegetation exists, it consists mainly of 
grassland, savannah or shrubland. 

Tree growth remains insignificant in those 
parts of the world where water is present in 
some form, but temperatures are too low 
for growth during all or part of the year 



Grasses and low shrubs are 
the mam plant growth 
forms in dryland regions. 



74 WORLD ATLAS OF BIODIVERSITY 



Map 5.1 

Photosynthetic activity 
on land 

A map illustrating the 
normalized difference 
vegetation index INDVI) 
calculated from AVHRR 
satellite data and 
aggregated over the year 
1998. NDVI values vary with 
absorption of red light by 
plant chlorophyll and the 
reflection of infrared 
radiation by water-filled leaf 
cells, and provide an 
indication of photosynthetic 
activity. The map reveals a 
clear distinction between 
areas rich in standing 
growth of plants, whether 
cropland or natural 
vegetation, and areas where 
standing plant growth is 
sparse or absent. 

Source. Adapted from image provided 
by the SeaWiFS Project, NASA/Goddard 
Space Fllgtil Center and GRBIfvlAGE. 



Box 5.1 Defining ecosystems 



Photosynthetic activity 



High 




I 



Low 



For example, in polar regions there is an 
abundance of water, but because of permanent 
low temperature it is mainly in solid form and 



The word 'ecosystem' was introduced by the plant ecologisl Tansley in 1936, to refer to the 
communities of plants, animals, other organisms and the physical environment of any given 
place. The concept reflected a new and visionary approach to biological research which 
focused on the system-level flow of materials and energy between components at any given 
study site. The spatial boundaries of the system were originally of little or no significance. 
The term is now widely used, particularly in the context of environmental planning, to refer 
to broad biological communities of similar appearance, usually defined by physical, climatic, 
structural or phenological features. Ecosystem diversity is generally understood to refer to 
the range of different kinds of ecosystem, in this sense, within some defined area. In this 
usage, the 'ecosystem' is treated as a map unit and the spatial boundaries of the system 
assume major significance. 



y^ 




S 



so unavailable to living organisms. In tundra 
areas at high latitudes the subsurface is 
permanently frozen and plant growth is 
restricted to the few summer months when 
thawing of the superficial layers makes liquid 
water available. In upper alpine regions, liquid 
water is often present in seasonal abundance, 
but temperatures are too low over the year as 
a whole to allow tree growth. 

Many areas theoretically have sufficient 
rainfall and a temperature regime suitable 
for supporting significant tree cover but do 
not do so. The most important reason for 
this is undoubtedly human intervention but 
in some areas this may also be a result of 
natural causes. Nutrient availability may be 
limiting. Slopes may be too steep to allow 
formation and retention of soil, so that trees 
cannot anchor themselves. Minerals may be 



Terrestrial biodiversity 75 



-^F— 




,^<f-^ ?aira 



present in toxic concentrations, or tlie fre- 
quency and intensity of natural fires or floods 
may be too higti. Large herbivores may also 
prevent establisfiment of significant tree 
cover, although as noted in Chapter 4 many of 
those large herbivores in the Americas and 
Australasia that might have exerted such an 
influence in the past are now extinct. 

Landcover classifications can be used as 
the basis for habitat or ecosystem classi- 
fications Isee Box 5.1], by incorporating 
elements of species composition and com- 
munity structure. Ecosystem maps should 
have more direct application to biodiversity 
conservation and management than maps of 
landcover; hovi/ever the development of such 
maps is constrained by the same problems of 
defining boundaries and developing a consis- 
tent approach to classification. Map 5.7 is a 



representation of global forest cover, derived 
by applying a simple classification of five 
forest and woodland habitat types to the 
landcover data plotted in Map 5.2. Forest 
classification is discussed further below. 

GLOBAL VARIATIONS IN TERRESTRIAL 
SPECIES DIVERSITY 

Just as habitat types show great variation 
across the world's land surface, so does the 
number of species that may be found in any 
given place. The spatial heterogeneity, wide 
range of present physical conditions and 
complex history of the worlds land surface all 
contribute to extreme variation in terrestrial 
biological d'versity. Variation in species 
number is not strictly related to variation in 
habitat type because two areas that are 
structurally similar may have very different 



74 WORLD ATLAS OF BIODIVERSITY 



Map 5.2 

Global land cover 

Map adapted from the 
global landcover 
classification developed by 
ttie University of Maryland. 
Tfie Maryland classification 
includes 13 classes and 
w^as based on AVHRR 
remote-sensing data with a 
spatial resolution of 1 km. 
For presentation purposes 
the data have here been 
generalized to a 4-km grid. 
Also for clarity, the two 
needleleaf forest classes in 
the original classification 
have been combined, as 
well as the two broadleaf 
forest classes, and the 
urban and built-up category 
has been omitted. 

Source: Data from University of 
Maryland Global Land Cover Facility. For 
full ijescription see Hansen . 





€ 



numbers of species present. The most impor- 
tant components of this variation can be 
expressed in three different, though not com- 
pletely independent, ways: 

• the kinds of organisms, particularly pri- 
mary producers, that live in any one place - 
this influences the kinds of habitats found 
there and is a reflection of ecosystem 
diversity; 

• the numbers of different kinds of organisms 
that live in any one place - this is usually 
assessed by measures of species diversity, 
although other classification systems le.g. 
guilds, functional groups, higher taxal can 
also be used; 

• the individual taxonomic identity of the 
organisms that live in any one place - 
this is determined by biogeography and 



influences, among other things, how a 
given area contributes to global biodiversity. 

Measuring species diversity 

Comparing the diversity of different parts of 
the world is complex because of the way 
diversity changes with scale. A vvfide range of 
observations has demonstrated that, as a 
general rule, the number of species recorded 
in an area increases with the size of the area, 
and that this increase tends to follow a 
predictable pattern known as the Arrhenius 
relationship, whereby: logS = c + zlogA where 
S = number of species, A = area and c and z are 
constants (see Figure 5.11. The slope of the 
relationship (z in the equation) varies 
considerably between surveys, although it is 
generally between 0.15 and 0.^0, but some 
survey data do not fit the relationship at all. A 



Terrestrial biodiversity 77 



•^^^ 




common generalization from tliis finding Is 
tfiat a tenfold reduction in an area II. e. loss of 
90 percent of habitat) will result in the loss of 
between 30 percent (with z = 0.151 and 60 
percent (with z = 0.401 of the species originally 
present, or approximately half the species. 

The relative diversity of different sites will 
often partly depend on the scale at which 
diversity is measured. Thus 1 m- of semi- 
natural European chalk grassland may 
contain many more plant species than 1 m' of 
lowland Amazonian rainforest, whereas for 
any area larger than a few square meters this 
could be reversed. In other words, when an 
area is sampled the number of species 
recorded Increases with the size of the area, 
but this rate of Increase varies from area 
to area, I.e. the slope of the Arrhenlus 
relationship is not constant everywhere. 



The reason for this Increase may be 
quite straightforward. When smalt areas are 
sampled they are likely to be relatively 
homogeneous in terms of habitat type. 




Log area 



Figure 5.1 

A typical species-area 

plot 

Note; The data, consisting of 
species counts in a series of 
areas of different size, are 
plotted on logarithmic axes 
resulting in a stralght-tlne 
graph, the slope of which Izl 
indicates the rate at which 
species number changes 
with changing area. 



78 WORLD ATLAS OF BIODIVERSITY 



At a small scale, as sample area Increases, so 
an increasing proportion of the species 
present in that habitat is lil<ely to be included 
in the sample. Beyond a certain point, 
however, as larger areas are sampled so an 
increasing number of different habitats will 
be included in the sample area, each with 
new species that are likely to be included 
in the sample. The species/area relationship 
therefore increasingly reflects habitat hetero- 
geneity at larger scales. Ecologists attempt to 
tal<e account of this by recognizing different 
kinds of biological diversity The diversity 
within a site or habitat is often referred to as 
alpha (al diversity while the differences 
between habitats are referred to as beta Ipl 
diversity. Thus an area with a wide range 
of dissimilar habitats will have a high f,- 
diversity, even if each of its constituent 
habitats may have low a-diversity Differences 
in site diversity over large areas, such as 
continents, are sometimes referred to as 
gamma (yl diversity. 

Measures of diversity can refer simply to 
species richness but can also be more 
sophisticated statistical measures that take 
into account the relative abundance of different 
species in a given place. A variety of different 
measures of this kind has been developed (of 



Box 5.2 Species and energy 

Latitudinal variation in diversity on land is strongly correlated v^ith, and may be largely 
explained by, vanation in incident energy over the Earth's surface. The relationship between 
diversity and productivity, and related measures, has been the subject of long-standing 
debate in ecology, but recent studies have shov\/n that at global or continental scale, 
organismal diversity, particularly as measured at higher taxonomic levels, is strongly 
correlated with available energy". This l<ind of relationship has been demonstrated, for 
example, for flowering plants, for trees, lepidoptera, land birds and land mammals in a 
range of countries and continents", and tor fishes in river basins at a global level'. One 
simplistic explanation for this may be that higher energy availability leads to increased net 
pnmary production (NPPl, and this broader resource base allows more species to coexist. 
While the general relationship appears robust, the details are complex. Energy availability 
can be measured in several ways: as heat energy, as potential (PET) or actual 
evapotranspiration (AETl, or as NPP, and which is the best predictor of diversity has yet to be 
determined. Some measure of the simultaneous availability of water and radiant energy may 
provide the best general predictor of potential macro-scale species richness". More 
complete explanation for richness variation would need also to consider the roles of 
topography history and edaphic factors. 



which H', the Shannon-Wiener function, is a 
commonly used example). With many of these 
measures, an area in which all species are of 
similar abundance would generally be given a 
higher diversity measure than an area with the 
same number of species, a few of which were 
very abundant and the remainder rare. 
Deriving these statistical measures requires 
intensive sampling- for this reason, simpler 
measures of species richness tend to be more 
useful at larger scales. 

Major patterns of variation in global 
species diversity 

Despite the difficulties of establishing strictly 
comparable quantifiable measures, a wealth of 
empirical observations indicates that species 
richness in eukaryotes tends to vary geo- 
graphically according to a series of fairly well- 
defined rules. In terrestrial environments: 

• warmer areas hold more species than 
colder ones; 

• wetter areas hold more species than drier 
ones; 

• areas with varied topography and climate 
hold more species than uniform ones; 

• less seasonal areas hold more species than 
highly seasonal ones; 

• areas at lower elevation hold more species 
than areas at high elevation. 

The single most obvious pattern in the global 
distribution of species is that overall species 
richness increases as latitude decreases 
toward the equator (see Box 5.2). At its 
simplest this means that there are more 
species in total and per unit area in temperate 
regions than in polar regions, and far more 
again in the tropics than in temperate regions. 
This applies as an overall general rule, and 
within most individual higher taxa (at order 
level or higher), and within most equivalent 
habitats. The pattern can be seen in Maps 5.3, 
illustrating vascular plant species diversity, 
and 5. A, which represents country-level 
diversity in terrestrial vertebrates and 
vascular plants. 

There is good evidence that moist tropical 
forests are the most species-rich environ- 
ments on Earth. If current estimates of the 



Terrestrial biodiversity 79 



number of as yet unknown species Isee 
Chapter 21 in the tropical forest microfauna 
are accepted, these regions, extending over 
perhaps 7 percent of the world's surface, may 
conceivably hold more than 90 percent of the 
world's species. If tropical forest small 
Insects are discounted, then coral reefs and. 
for flowering plants at least, areas of 
Mediterranean climate in South Africa and 
Western Australia may be similarly rich 
in species. 

Topographic heterogeneity may be expec- 
ted to play a significant part in determining 
species number for two reasons. First, such 
heterogeneity will increase habitat variability. 
thereby increasing the range of niches that 
can be occupied by different organisms. 
Second, depending on the size and mobility of 
organisms, the chances of geographic Isolation 
and speciatlon increase in topographically 
diverse landscapes. The role of topography 
has been demonstrated statistically at 
continent scale for North American mammals', 
and at both landscape and patch scale for 
vascular plants' '". 

The available Information on distribution 
of species Is geographically very Incomplete, 
and relates to only a small fraction of the 
1.7 million known species. Geographically, 
western Europe has been more thoroughly 
sampled than elsewhere, while large areas 
In the tropics, particularly of South America 
and central Africa are very poorly known. 
Taxonomlcally, the larger mammals, birds, 
vascular plants and a few invertebrate 
groups, such as Odonata. are better known 
than other groups of species. Because of the 
uneven availability of information there has 
been considerable interest In Identifying 
groups of Individual species that may serve 
as surrogates for biodiversity more gener- 
ally, or higher taxa. such as families, that 
might predict patterns of richness In their 
included species". 

Maps 5.5 and 5.6 represent, respectively, 
the distributions of flowering plant families 
(phylum Anthophytal and of terrestrial 
(non-aquaticl vertebrate families Iphylum 
Chordatal. plotted as a global family diversity 
surface for each group. The resulting maps 
share key features, notably a marked latitud- 



inal gradient in family richness, but also have 
striking differences, particularly In areas of 
high diversity. For example. Africa appears 
to be very rich in vertebrate families, most 
notably In moist forest areas around the 
Gulf of Guinea and In the east, including less 
humid woodland and savannah habitats, but 
Is relatively poor In plant families compared 
with other continents in the tropics. For 
flowering plants, this mirrors the sequence 
evident at species level, where there may be 
around 90 000 species In the neotropics. 
40 000 in tropical Asia and 35 000 in tropical 
Africa'-. 

Blogeography and endemism 

While ecological factors Influence which kinds 
of species, and how many of them, can persist 
In a given area, history has already deter- 
mined which actual lineages are present. A 
complete explanation for global variation In 
biodiversity must therefore involve both 
historical events and current ecological pro- 
cesses. The former are implicit in any explan- 
ation of the origin of diversity, the latter In 
explanations of Its maintenance; these being 
two separate, although Intimately linked, 
problems. On land, continental drift resulting 
from plate tectonics, climate change, moun- 
tain building or sea-level change, and prob- 
ably the evolutionary lability of different 
lineages, are among the Important historical 
factors. Geographic features commonly re- 
strict or prevent the further dispersal of 
species: for example, a large river can pre- 
sent a barrier to a terrestrial species, the sea 
Is a barrier to non-flying Island forms and 
land is a barrier to freshwater species. 

Barriers to dispersal explain why the fauna 
and flora of ecologically similar areas In 
geographically separated parts of the world 
tend to be composed largely of different 
individual species. They also underlie the 
phenomenon of endemism. An endemic 
species Is one restricted to some given area, 
which may be a continent or country, or more 
significantly a relatively small area, such as a 
mountain block. Island or lake. Discrete areas 
of complex topography, particularly In the 
tropics, often have high endemism In a 
range of taxonomic groups, possibly because 



WORLD ATLAS OF BIODIVERSITY 



climate change has encouraged speciation by 
isolating different lineages at different times. 

Communities and ecoregions 
The most comprehensive attempts to 
describe and classify habitats try to combine 
elements of all three sources of variation 
outlined above. At fine scales these are 
generally based on community ecology. On 
land this is exemplified by the phyto- 
sociological approach developed principally in 
continental Europe during the 20th century 



UNESCO" 


Closed forest 


Trees > 5 m tall with crowns interlocking 




Woodland 


Trees > 5 m tall with crowns not usually 
touching but with canopy cover > 40% 


US classification 


Closed tree canopy 


Trees with crowns interlocking, with 


standards" 




crowns forming 60-100% cover 




Open tree canopy 


Trees with crowns not usually touching 
forming 10-60% or 25-60% cover 


FAO" 


Forest 


Land with tree canopy cover > 10% and > 0,5 ha 
in area; trees should be able to reach a 
minimum height of 5 m 




Other wooded land 


Land with either a crown cover of 5-10% of trees 
able to reach a height of 5 m at maturity; or 
crown cover of more than 10% of trees not able 
to reach a height of 5 m at maturity; or with 
shrub or bush cover > 1 0% 



Table 5.2 

Different definitions of 

forest cover 



This was intended to describe and classify 
plant communities on the basis of dominant 
and other associated species, by inspection or 
quadrat analysis of vegetation patches, taking 
into account species identity, growth form and 
abundance. One major problem with this is 
that the more precisely a community is 
defined, the more site-specific it becomes, 
and hence the more limited its use in higher- 
level analysis and planning. A more recent 
related approach" attempts to delimit eco- 
regions', these being defined as relatively 
large units of land each of which contains a 
distinct assemblage of natural communities 
and species, with boundaries similar to the 
original extent prior to anthropogenic change. 
These are nested within a hierarchy having 



traditional biogeographic realms and biome 
systems [ecosystem types! as the first and 
second levels. 

FORESTS 

Forests and woodlands probably once covered 
about half of global land area and now cover 
about one quarter They provide habitat for 
half or more of the worlds species. They are 
responsible for just under half of the global 
terrestrial annual net primary production 
[Table l.ll, and they and their soils house 
about 50 percent of the world's terrestrial 
carbon stocks. In addition to carbon storage, 
forests perform many other important 
ecosystem services, such as regulating local 
hydrological and nutrient cycles, and stabiliz- 
ing soils and watersheds. 

Forests also provide a wide variety of 
products, including food and fuel, medicines, 
construction materials and paper, which are 
important both for human subsistence and for 
economic activity. Wood products are one of 
the most economically important natural 
resources. In the region of 3.3 billion cubic 
meters (m-1 of wood is extracted from forests 
and other habitats annually, the equivalent of 
several hundred million trees. Just over half 
of this volume is used as fuelwood and 
charcoal, of which developing countries 
consume 90 percent". The remainder is 
industrial roundwood, which is processed into 
various wood products. Forests are frequently 
important culturally and play a significant role 
in the spiritual life of communities worldwide. 

What Is a forest? 

Despite their importance in a number of 
different human contexts and the large 
amount of research that has focused on forest 
ecosystems, a precise definition of 'forest' 
remains elusive. Although it is generally 
accepted that the term indicates an eco- 
system in which trees are the predominant 
life form, the problem arises because of the 
broad range of systems in which trees occur 
and the difficulty, even, in deciding what 
constitutes a tree. For example, tree species 
may dominate at high altitude, but be barely 
recognizable as trees because of their 
spreading prostrate forms. Savannahs may 



Terrestrial biodiversity 81 



"««/,(, 



possess large numbers of trees, but as they 
can occur at low density in association with 
other life forms it may be difficult to define 
precisely in which areas they are dominant. A 
variety of different definitions of forest have 
been proposed by organizations that evaluate 
and monitor natural resources (Table 5.21. 
Estimates of forest area may vary widely 
depending on the definition adopted. Of sing- 
ular importance is the degree of canopy cover 
used as the threshold for dividing forests from 
non-forests (Table 5.21. As a consequence, the 
precise definitions employed should be borne 
in mind when comparing forest cover data 
provided by different institutions. 

Forest types 

There is great variation in the forms and types 
of forest distributed throughout the world. 
Information about this variation and the 
distribution of forest vegetation types is 
crucial to understanding the different roles of 
forests in supporting biodiversity, in carbon 
and hydrological cycles and other ecosystem 
processes, and in supplying wood and non- 
wood forest products. However, if deriving a 
satisfactory definition of forest is problematic, 
arriving at consensus on how to classify 
forests is even more difficult. 

A number of global classification systems 
have been proposed, but as yet none has 
gained universal acceptance. The UNESCO 
(United Nations Educational, Scientific and 
Cultural Organization) system proposed by 
Ellenberg and Mueller-Dombois'"' is one such 
system. It includes nearly 100 forest and 
woodland subformations' and allows for yet 
finer subdivisions, but many of the charac- 
teristics that separate categories can only be 
determined in the field. Other classifications, 
such as the EROS Data Center seasonal 
landcover regions, with nearly a thousand 
classes, reflect more strongly the nature of 
landcover data obtained from Earth-orbiting 
satellites and the methods used in analyzing 
and classifying them'. This complex system 
has been translated into a much less complex 
one in the International Geosphere-Biosphere 
Programme (IGBP) classification, which 
includes seven forest and woodland types that 
reflect phenology and canopy closure world- 



Country 


Forest definition 




Forest area Ikm'l 


Australia 


Tree canopy cover > 20% 




384 115 




Tree canopy cover > 70% 




30 729 


Senegal 


Tree canopy cover > 10% 








(includes dry woodland] 




78 689 




'Closed' forest (canopy cover 


> iO%] 


3 934 



wide, but provides little other information on 
forest physiognomy, composition or environ- 
ment within the class names'. Map 5.7 
represents the global distribution of a range 
of forest types, based on forest physiognomy 
and phenology, here aggregated into five 
broad categories, discussed further below. 

Temperate and boreal needleleaf forests 
Distribution, types and characteristic taxa 

Temperate and boreal needleleaf forests cover 
a larger area of the world than other forest 
types. They mostly occupy the higher latitude 
regions of the northern hemisphere, as well as 
high-altitude zones and some warm temper- 
ate areas, especially on nutrient-poor or 
otherwise unfavorable soils. These forests are 
composed entirely, or nearly so, of coniferous 
species IConiferophyta). In the northern 
hemisphere, pines Pinus, spruces Picea, 
larches Larix, silver firs Abies, Douglas firs 
Pseudotsuga and hemlocl<s Tsuga dominate 
the canopy, but other taxa are also important. 
In the southern hemisphere coniferous trees, 
including members of the Araucariaceae, 
Cupressaceae and Podocarpaceae, often 
occur in mixtures with broadleaf species in 
systems that are classed as broadleaf and 
mixed forests. 



Forest type Area ImiUlon kmM 


Temperate and boreal needleleaf 


13.1 


Temperate broadleaf and mixed 


7.5 


Tropical moist 


11.7 


Tropical dry 


2.5 


Sparse trees and parkland 


6.9 


Total 


41.7 



Table 5.3 

Sample effects on forest 
area estimates of different 
forest definitions 



Table 5.4 

Global area of five main 

forest types 



Note; Based on forest 
cover as shown in Map 5.7. 
Estimates of this kind 
vary significantly with 
different source data and 
classifications. 



82 WORLD ATLAS OF BIODIVERSITY 



Structure and ecology 

The structure of temperate and boreal 
needleleaf forests is often comparatively 
simple, as conifer canopies are efficient 
[igfit absorbers, reducing the possibilities for 
development of lov^er strata in the canopy. 
The tallest of these forests, the giant redwood 
forests of the west coast of the United States, 
may reach 100 m in height, but most are 
much shorter, and indeed some pine forests 
at high altitude or in arid environments are 
quite stunted. 

The distribution of temperate and boreal 
needleleaf forest is limited at high altitudes 
and latitudes by lack of enough days with 
temperatures suitable for growth, and at 
lower altitudes and latitudes by competition 
with broadleaf species. In about a quarter of 
the area of these forests, deciduous conifers 
of the genus Larix replace the evergreen 
species. This is especially true In far northern 
continental areas with extremely low winter 
temperatures. 

In many areas, wildfire is an important 
factor affecting the dynamics and main- 
tenance of the forest ecosystem. Many coni- 
ferous species produce resins that Increase 
flammability, and many are characterized by 
thick bark that increases the resistance of 
adult trees to fire-induced mortality. A 
number of tree species, such as the jack pine 



Box 5.3 Fire in temperate and boreal forest 

In general, as the frequency of fire increases, the intensity of individual fires tends to 
decrease, because of reductions in the standing amount of fuel. Increased fire frequency 
also tends to Increase the diversity of the herb layer by severely affecting the shrub layer 
Fires are caused by natural events such as lightning strikes and by human activities. 
Changing forest management has significantly altered fire regimes In coniferous forests. 
The 1988 fires that affected over 5 000 km' In and around Yellowstone National Park, 
United States, were attributed to the accumulation of fuel in the forests resulting from a 
long-term policy of fire suppression. 

Climatic variation plays an important part in determining fire occurrence and severity. The 
Yellowstone fires, Canadian fires in 1989, and the 1987 Black Dragon fire In the boreal 
forest region of China and Siberia In 1987, were all ascribed to unusual drought conditions. 
The Black Dragon fire burned over 70 000 km^ and qualified as the largest forest fire in 
recorded history'. There is concern that global climate change may increase the frequency 
and impact of fires in boreal coniferous forests. 



Pinus banksiana. have serotinous cones, 
which depend on the high temperatures of 
forest fires to open and release their seeds. 
Non-coniferous species are generally less 
resistant to fire than conifers, so periodic fires 
are an Important factor In maintaining the 
composition and extent of these forests. 

Biodiversity 

Although tree species richness is low in most 
temperate and boreal needleleaf forests, 
many conifer species are of great con- 
servation concern. A well-known example 
occurs In the giant redwood forests of 
northern California, where the redwood 
Sequoiadendron giganteum is considered 
vulnerable to extinction'. About 22 percent 
(140) of the world's 630 conifer species have 
been assessed as globally threatened". Most 
of the threatened taxa are characteristic of 
mixed forests, particularly in the southern 
hemisphere. Old growth conifer stands, which 
may be many centuries in age, represent an 
irreplaceable gene pool and are an important 
habitat for many other organisms. 

Species richness in these forests is 
commonly Increased by a relatively high 
diversity of mosses and lichens, which grow 
both on the ground and on tree trunks and 
branches. For example, there are at least 100 
species of moss growing in the coniferous 
forests between 1 300 and 2 000 m altitude on 
Baektu Mountain, on the Chinese-Korean 
border^ Mosses and lichens are Important 
sources of food for many animals of coni- 
ferous forest. 

Vertebrate richness is generally lower in 
boreal needleleaf forests than In broadleaf 
temperate and tropical forests. Many species 
are wide-ranging generallsts, often with a 
holarctic distribution, e.g. wolf Canis lupus, 
brown bear Ursus arctos. 

There are a number of animals of con- 
servation concern that are dependent on 
temperate needleleaf forests. The northern 
spotted owl Strix occidentalis caurina requires 
large expanses of old-growth coniferous forest 
in the northwest United States to provide 
nesting habitat and adequate food resources; 
Kirtland's warbler Dendroica kirtlandii needs 
young regrowing jack pine as a nesting habitat. 



Terrestrial biodiversity 83 



"^'^h 



Fire suppression programs have reduced the 
available habitat for this species to critical 
levels. While there is relatively little infor- 
mation available on the conservation status of 
Invertebrates, many common old-growth 
species are known to become much rarer in 
modern managed forests, often through the 
loss of essential microhabltats'. 

Rote in carbon cycle 

Temperate and boreal needleleaf forests make 
a significant contribution to the global carbon 
balance, accounting for more than a third of 
the carbon stored In forest ecosystems (Table 
5.61 and about 8 percent of global annual net 
primary production (Table 1.1|. Furthermore, 
the soils under these forests store large 
amounts of carbon (up to 250 metric tons per 
hectare), some of which may be liberated by 
Increasing decomposition rates related to 
climate change. Of particular note in this 
context are the giant conifer forests of the 
Pacific northwest of the United States. These 
forests may store more than twice as much 
carbon per hectare as tropical rainforests. 

Use by humans 

Global Industrial roundwood production is 
dominated by coniferous species. Pines Pinus. 
spruces Picea, larches Larix, silver firs Abies. 
Douglas firs Pseudotsuga and hemlocks Tsuga 
from the needleleaf forests of the northern 
hemisphere are the major sources of softwood. 
Some conifer species from the southern 
hemisphere and the tropics also provide 
excellent timbers. Large-scale exploitation of 
natural coniferous and mixed forests is taking 
place around the Pacific Rim, notably In North 
America, Russia and Chile. Temperate and 
boreal needleleaf forests are also a principal 
source of pulpwood for paper production. 

Other ecosystem services 

These forests, like others, stabilize soils on 
sloping topography, especially in mountainous 
regions. Recognition of this function in early 
20th century Switzerland was the basis for a 
new program of forest planting to control 
avalanches in the Alps. Coniferous forests 
have high recreational and cultural values, 
especially in regions such as northern Europe. 



Temperate broadleat and mixed forests 
Distribution, types and characteristic taxa 

Temperate broadleaf and mixed forests cover 
some 7.5 million km- of the Earth's surface 
(Table 5.4). They Include such forest types as 
the mixed deciduous forests of the United 
States and their counterparts in China and 



Table 5.5 

Important families and 
genera, and numbers of 
species, in four areas of 
temperate broadleat 
deciduous forest 

Source: Afler Rohng . 



■Family 


Genus 


Common Northeast 
name America 


Europe 


East Asia 


South V 
America^ 


Fagaceae 


Quercus Oak 
Lithocarpus 
Castanopsis 

Cyclobalanopsis Asian oak 
Castanea Chestnut 
Fagus Beech 
Nothofagus Southern beech 


37 
1 

4 
1 


18 

1 
2 


66 
47 
45 
30 
7 
7 


10 


Aceraceae 


Acer 
Dipteronia 


Maple 


10 


9 


66 

1 




Betualceae 


Betulus 
Alnus 


Birch 
Alder 


6 

5 


4 
4 


36 
14 




Salicaceae 


Salix 


Willow 


13 


35 


97 




Juglandaceae 


Car/a 

Juglans 

Platycarya 


Hickory 
Walnut 


11 
5 


1 


4 
4 
2 




Legumlnosae 


Cercis 

Cleditsia 

Gymnodadus 

Maackia 

Robinia 

Acacia 

Albizia 


Redbud 
Honey locust 

Locust 
Acacia 


1 
2 
1 

1 


1 


2 
7 
3 
3 
1 

10 


1 
1 


Magnoliaceae 


Liriodendron 
Magnolia 


Tulip tree 
Magnolia 


1 
8 




1 
50 




Oleaceae 


Fraxinus 
Osmanthus 


Ash 


4 
2 


3 


20 
10 




Rosaceae 


Malus 

Prunus 

Pyrus 

Sorbus 

Kageneckia 

Quillaia 


Apple 

Cherry 
Pear 
Mountain ash 


1 
3 
1 
3 


1 

4 
1 

5 


3 
59 

5 
18 


2 

1 


Tllllaceae 


Tilia 


Basswcod, 
lime, linden 


i 


3 


20 




Lauraceae 


Phoebe 

Sassafras 

Beilschmidia 

Cryptocarya 

Persea 


Sassafrass 


1 




16 
2 




("lyrtaceae 


Amomyrtus 
Myrceungenelli 
Myrceugenia 
Nothomyrcia 










8 
1 


Ulnnaceae 


Celtis 
Ulmus 
Zelkova 


Hackberry 
Elm 


2 
4 


1 
3 


14 

30 

3 




Carpinaceae 


Carpinus 
Ostrya 


Hornbeam 


2 
1 


2 

1 


25 
3 





84 WORLD ATLAS OF BIODIVERSITY 



Table 5.6 

Biomass and carbon 
storage in the world's 
major forest types 

Note: The carbon storage 
figures are over- rather tfian 
underestimates as they 
incorporate no weighting tor 
anthropogenic disturbance or 
for the variation m biomass 
and area among different 
forest classes within the 
broad types- 

Source: After Adams and Huston , 



Japan, the broadleaf evergreen rainforests of 
Japan, Chile, New Zealand and Tasmania, and 
the sclerophyllous forests of Australia, the 
Mediterranean and California. These last are 
characterized by a predominance of often 
evergreen trees with small, hard, leathery 
leaves. 

Trees belonging to the Anthophyta and the 
Coniferophyta grow in mixtures in many of 
these forests, especially in the southern 
hemisphere. For example, the Valdivian and 
Magellanic rainforests of Chile include 
mixtures of Nothofagus Ian anthophytel 
with Podocarpus and members of the 
Cupressaceae Iboth conifers)". Much of 
the forest of New Zealand was originally 
a mixture of the conifers Podocarpus, 
Phyllocladus, Dacryocarpus and Dacrydium. 
with such Anthophyta as Metrosideros, 
Elaeocarpus and Weinmannia. In North 
American mixed forests, pines Pinus, hem- 
locks Tsuga and cypress Taxodium, among 
other conifers, are mixed in various propor- 
tions with oaks Quercus. maples Acer, ashes 
Fraxinus, hickories Carya, beeches Fagusand 
other hardwoods. 

The beech family IFagaceael is generally 
important in temperate broadleaf forests 
(Table 5.51, with such genera as Castanopsis 
and Cyclobalanopsis playing an important 
rolein Japan and China, and many different 
Quercus species being important elements 
of hardwood forests in North America, 
Asia and Europe and of sclerophyllous 
forests in California and the Mediterranean, 
Sclerophyllous forests in Australia are largely 
made up of Eucalyptus species, as are the 
wet forests of Tasmania, which may also 
include Nothofagus'. 



Forest type 


Above-ground 


Estimated total 


Estimated global 




biomass per area 


carbon storage 


total carbon 




(metric tons/lial 


including soil and 


storage 






roots (metric tons/hal 


(petagrams = 10" gl 


Temperate needleleaf 


200-1 500 


300 1700 giant conifer] 


394 


Temperate broadleaf 








and mixed forest 


150-300 


350 


262 


Tropical moist forest 


195-500 


300 


350 


Tropical dry forest 


98-320 


250 


62 



Structure and ecology 

Depending on the precise forest type, these 
forests tend to be structurally more complex 
than pure coniferous forests, having more 
layers in the canopy. It is not uncommon for an 
upper canopy layer to have as many as six 
distinct subcanopy and understorey layers 
below it. The tallest of these forests, the mixed 
forests of southern Chile and some Eucalyptus 
forests of Australia, can reach over 50 m in 
height. On the other hand, some sclero- 
phyllous forests barely reach 5 m. Deciduous 
forests may support rich herb layers, which 
depend on the increased penetration of 
sunlight early in the growing season. 

As in needleleaf forests, fire plays an 
important role in many types of temperate 
broadleaf and mixed forest. Both natural and 
anthropogenic fires are important ecological 
factors affecting the maintenance of forest 
structure and composition, especially in the 
Eucalyptus forests of Australia and the 
sclerophyllous forests of the Mediterranean 
and California. Spatial variation in forest 
structure and composition may be influenced 
by fire and other kinds of natural and 
anthropogenic disturbance. When canopy 
trees die, the resulting gaps in the canopy 
increase light availability locally, and such 
areas may be colonized by a different subset 
of the forest flora. These gap dynamic pro- 
cesses are important in maintaining stand 
diversity. Relatively few old-growth broadleaf 
and mixed forests remain in the temperate 
zones because of the historical exploitation of 
these forests by human populations. 

Biodiversity 

As might be expected from their structural 
diversity, temperate broadleaf and mixed 
forests tend to be richer in species than 
coniferous forests. Southern mixed hardwood 
forests in the United States are commonly 
composed of as many as 20 canopy and 
subcanopy tree species and may include as 
many as 30 overstorey species'". In comparison, 
European forests tend to be less species rich, 
while the deciduous forests of East Asia may be 
the richest of all" (Table 5.51. 

In the late 1990s, some 370 temperate 
dicotyledonous tree taxa worldwide were 



Terrestrial biodiversity 85 



"^K 



considered to be of conservation concern". 
Japan alone has 43 threatened endemic tree 
species, which are mostly characteristic of 
its temperate broadleaf forests'". Fitzroya 
cupressoides, an endangered large conifer 
once important for the local and international 
timber trade, is characteristic of mixed 
forests in southern Chile. It has been over- 
exploited and proves to be highly dependent 
for its regeneration on large-scale natural 
disturbances, such as landslides and light- 
ning strikes. F. cupressoides is nov/ listed in 
CITES (Convention on International Trade in 
Endangered Species of Wild Flora and Fauna! 
Appendix I, and commercial international 
trade is accordingly prohibited. 

While temperate broadleaf forests gen- 
erally support moderate animal diversity, 
species richness is often lower than in 
comparable tropical habitats. However, there 
is considerable geographical variation in 
richness in the northern hemisphere; the 
forests of East Asia are generally the most 
species rich". As in the boreal needleleaf 
forests, many of the mammal and bird 
families of northern broadleaf forests are 
holarctic in distribution and can be found in 
other habitat types. 

In contrast to most northern hemisphere 
forests, the temperate forests of southern 
South America, Australia and New Zealand 
contain several restricted-range mammals 
and birds. Analysis" of habitat requirements 
of Australian mammals indicates that the 
forests of southeast Australia and Tasmania 
are of particular importance for wildlife 
conservation. 

Examples of temperate broadleaf forest 
species of special conservation concern 
include the huemul deer Hippocamelus 
bisulcus of the southern Andes, threatened by 
both habitat loss and hunting; a number of 
New Zealand forest birds including the 
kakapo Strigops habroptilus and some kiwi 
species Apteryx spp., which are threatened 
mainly by introduced predators; Leadbeater's 
possum Gymnobelideus leadbeateri, which is 
threatened by the loss of specific habitat 
within the montane ash forests of Victoria 
lAustralial; the Amami rabbit Pentatagus 
furnessi of Amam\ Island iJapan], threatened 



by habitat loss and introduced predators; and 
the European bison Bison bonasus of central 
and eastern Europe, with low genetic diversity 
and at risk from disease. 



Ttie figure shows tfie living planet index for a sample of 170 forest-occurring birds in Nortli 
America and Europe. The prevailing trend during 1970-99 viias for a small net increase over 
the period. This could be interpreted as reflecting a phase of relative stability in these forests 
during recent decades, following centuries of decline in area, but may also be correlated 
with local increase in forest area through the spread of plantations, and management in 
some forests may have exerted a positive effect. The separate North America and Europe 
samples |123 and 47 species] show extremely similar trends. 



uo 




1970 1975 1980 1985 

Source: UNEP-WCMC. from dala collaled for Lofi". 



Role in carbon cycle 

Above-ground biomass of temperate decid- 
uous forests in Europe, the United States and 
the former Soviet Union ranges from 140 to 500 
metric tons per hectare depending on stand 
age and altitude, among other factors". Soil 
carbon storage in these systems is lower than 
in the needleleaf forests, owing both to climat- 
ically favorable conditions for decomposition 
and to the inherent greater decomposability of 
leaf litter from broadleaf trees. 

One survey" suggests that average soil 
carbon storage in these forests is some- 
where between 135 and 160 metric tons per 
hectare. Thus, global carbon storage across 
the forest types included in this broad cate- 
gory may total as much as 231 petagrams 
(Table 5.61. 



1990 



1995 



/— 



86 WORLD ATLAS OF BIODIVERSITY 



Tropical moist forests 
are often particulary ricfi 
in palms. 



Use by humans 

Temperate broadleaf and mixed forests have 
provided large amounts of timber over tfie 
centuries, but ttiey have largely been replaced 
by tropical forests as the primary source of 
hardwood timber in global trade. Hardwood 
production continues in the temperate zones, 
for products such as wood chips, furniture 
and finishing wood. 

Some of the non-timber products of 
temperate mixed forests include camphor 
from Cinnamomum camphorum in Japan 




(though this has now largely been replaced 
by synthetics], and sweet chestnuts 
Castanea sativa from southern Europe. 
Mushroom production is also a major 
income source in some parts of Europe, 
North America and Asia. 

Other ecosystem services 

Services provided by temperate broadleaf and 
mixed forests Include soil and watershed 
protection. This is especially important in the 
southern Andes" and other areas where steep 
topography is responsible for a high incidence 
of landslides, but it is also a benefit in 
Mediterranean regions where soils are prone 
to degradation. Natural temperate forests 
are important reservoirs of genetic material 
of trees such as eucalypts that are now 
commonly grown as plantation species. These 
forests, perhaps more than any other type, 
experience significant recreational use in 
many areas. 



Tropical moist forests 

Distribution, types and characteristic taxa 

Tropical moist forests cover more than 
11-5 million km' of the humid tropics (Table 
5. A and Map 5.7) and include many different 
forest types. The best known and most 
extensive are the lowland evergreen broadleaf 
rainforests, which make up over half the total 
area. These include, for example: the season- 
ally inundated varzea and igapo forests and 
the terra firme forests of the Amazon basin; 
the peat forests and moist dipterocarp forests 
of Southeast Asia; and the high forests of the 
Congo basin. 

Mountain forests are generally divided 
into upper and lower montane formations on 
the basis of physiognomy. These include 
cloud forest - the middle- to high-altitude 
forests that derive a significant part of their 
water supply from cloud, and support a rich 
abundance of epiphytes. Mangrove forests 
(see Chapter 6] also fall within this broad 
category. 

The high diversity of many tropical forests 
(see below] makes it difficult to characterize 
them taxonomically. However, some plant 
families are more prevalent than others. In 
neotropical moist forests, the legumes 
Leguminosae are particularly abundant and 
are often the most species-rich family. 
Other families that are generally among the 
richest in tree species in lowland neotropical 
moist forests are Moraceae, Lauraceae, 
Annonaceae, Sapotaceae, Myristicaceae, 
Meliaceae, Euphorbiaceae and Palmae". In 
Southeast Asian lowland moist forests the 
dominant family is the Dipterocarpaceae; the 
Myrtaceae is also very speciose". In African 
rainforests legumes are again important, and 
the ten richest families usually include the 
Olacaceae, Sterculiaceae, Dichapetalaceae, 
Apocynaceae, Sapindaceae and Ebenaceae. 
The other most abundant families in both 
Africa and Asia are often the same as those in 
the Americas''\ 

Structure and ecology 

Many tropical moist forests have canopies 
iO to 50 m tall, and some have emergent trees 
that rise above the main canopy to heights of 
60 m or more. Such large-stature forests are 



Terrestrial biodiversity 87 



characteristic of lowland forests and some 
lower montane forests on relatively nutrient- 
rich soils. Another characteristic of these 
forests IS a relatively high frequency of woody 
lianas and, especially in the nootropics, 
palms". Moist tropical forests are also known 
for a high abundance and diversity of vascular 
epiphytes, which take advantage of the higher 
light availability found in the canopy and can 
survive because of abundant rainfall and high 
atmospheric moisture. On more nutrient- 
poor soils and at higher altitudes forest 
stature decreases substantially; communities 
such as those on white sands (bana and 
campinal and in upper montane environments 
lelfin forests] may be no more than a few 
meters tall. With increasing altitude, decreas- 
ing forest stature is accompanied by a reduc- 
tion in the frequency of lianas and palms, and 
an increase in tree ferns ICyatheaceae, 
Blechnaceael and non-vascular as well as 
vascular epiphytes". 

Unlike the other forest types discusaed 
here, tropical moist forests have relatively 
little seasonal limitation to their growth, 
though seasonal drought may be a limiting 
factor, particularly in the semi-evergreen 
formations. However, the tropical moist forest 
environment is an intensely competitive one. 
Though solar energy inputs are high, canopy 
closure and complexity are also substantial, 
resulting in efficient capture of incident 
radiation and understorey light availability 
frequently much less than 2 percent of that 
above the canopy. This in turn limits the 
growth of understorey species and regener- 
ating trees. Some species can tolerate low 
light availability, while others grow or 
regenerate only in gaps in the canopy. Such 
gaps are formed by the death of one or more 
canopy trees and represent a significant 
contribution to overall environmental hetero- 
geneity, which is an important contributor 
to high diversity within tropical forests'". 
Infrequently, catastrophic disturbances such 
as blowdowns caused by hurricanes or 
convective storms may create large areas of 
regenerating forest" ™ and perhaps alter the 
long-term forest composition, as can logging 
and other forms of forest disturbance caused 
by human activities (see belowl. 



Soil nutrients are another limiting re- 
source in many forests. Soils in the humid 
regions of the tropics are notoriously poor in 
nutrients owing, among other factors, to loss 
through leaching by the high annual rainfall 
and to retention in the high-standing biomass. 
The formation of gaps in the canopy allowing 
regeneration of tree species is important for 
increasing the heterogeneity of available 
nutrients as well as the availability of solar 
energy. Forests such as the Amazonian 
varzea, which are seasonally inundated by 
sediment-bearing rivers, are an exception to 
this nutrient limitation. 

Biodiversity 

In numerical terms, global terrestrial species 
diversity is concentrated in tropical rain- 
forests. Many theories have been proposed to 
explain this phenomenon*". Generally speak- 
ing, the wet tropical forests of Africa have a 
lower tree species richness than those of Asia 
and America (Table 5.7). However, there is 
great local variation in species richness. 
Within the Amazon basin, tree species rich- 
ness ranges from 87 species per hectare in 
the east" to 285 species in central Amazonia" 
and nearly 300 species in the west'l 

The high diversity of tree species in 
lowland evergreen rainforests is mirrored in 
the diversity of epiphytes and lianas, which is 
also much higher in neotropical forests than 
in other regions^'. Fifty-three families in the 
Anthophyta and at least nine pteridophyte 
(Filicinophyta and allies) families include 
epiphytes. Of nearly 25 000 species of 
vascular epiphytes, around 15 000 belong 
to the Orchidaceae. Nearly a thousand 
others are members of the pineapple family 
Bromeliaceae, which is primarily neotropical. 
Other groups having a high diversity of 



Table 5.7 

Tree species richness in 

tropical moist forests 

Sources Afler Phillips etat."'. 



Region 



Africa 

Southeast Asia 
Americas 



No. of tree species 

l> 10 cm diameter! 

per hectare 

56-92 
108-2i0 
56-285 




WORLD ATLAS OF BIODIVERSITY 






::?=^ 



Map 5.3 

Diversity of vascular plant 

species 

This map shows the 
species richness of 
vascular plants, plotted as 
a world density surface. It 
is based on some 1 iOO 
literature records from 
different geographic units, 
with richness values as 
mapped calculated on a 
standard area of 10 000 km' 
using a single species-area 
curve. Value categories 
range between extremes of 
more than 5 000 species 
and fewer than 100 species 
per 10 000 km\ 

Source: Data and analysis © Wilhelm 
BarlhloU IBotanic Institute and Botanic 
Gardens. University of Bonn). 
Reproduced by permission, with 
modification to colors. For furtfier 
details see Bartfilott and website 
hltp://www.botanik,uni-bonn,de/system/ 
biomaps,htm#worldmap [accessed 
f^arch 20021. 



Diversity 



High 



Low 



epiphytic species are the cactus family 
Cactaceae, the aroids Araceae, the pepper 
family Piperaceae and the African violet family 
Gesneriaceae. 

Not all tropical moist forests have high 
species richness. Mangrove ecosystems have 
a low diversity of tree species despite their 
sometimes high productivity and high animal 
diversity (see Chapter 6). Extremely nutrient- 
poor soils, such as white sands, lead to the 
development of low-diversity forests including 
bana and campina''. As climate becomes 
more seasonal, tree species richness tends to 
decline [see dry forests, below). Increasing 
altitude also tends to reduce species rich- 
ness, with montane forests typically having 
fewer tree species than lowland ones". 

Regionally, the forests of Asia and South 
America are rich in animal species, generally 




more so than those of Africa"'. Many forest 
animals are largely confined to moist forests, 
e.g. the okapi Okapia johnstoni, but some 
are widespread outside, such as leopard 
Panthera parclus'\ In Africa, the Guineo- 
Congolean forest block contains more than 80 
percent of African primate species, and nearly 
70 percent of African passerine birds and 
butterflies". About half of the 1 100 South 
American reptile species are found in moist 
forests, with around 300 of these endemic to 
the habitat". Amphibian species are partic- 
ularly diverse in tropical moist forests"; 90 
percent of 225 species identified in the 
Amazon basin forests are endemic. 

The importance and diversity of the fish 
communities in forest streams and rivers is 
often overlooked; the Amazon basin has the 
richest freshwater fish fauna known, with at 



Terrestrial biodiversity 89 



■^^^ 




[east 2 500 species, many Important as major 
seed predators and dispersal agents'". Local 
species richness of insect and other arthro- 
pod groups in tropical forest canopies is much 
higher than in temperate forests'". Around 
one third of the animal biomass of the 
Amazon terra firme rainforesi consists of ants 
and termites, and each hectare of soil is 
estimated to contain more than 8 million ants 
and 1 million termites". 

Numerous tropical moist forest species are 
of conservation concern. Notable animals 
include the Sumatran rhino Dicerorhinus 
sumatrensis of Southeast Asia, endangered by 
habitat fragmentation and hunting; the bonobo 
Pan paniscus of the Democratic Republic of 
the Congo, threatened by habitat destruction 
and hunting for food; the Philippine eagle 
Pithecophaga jefferyi of the Philippines, 



reduced to small fragmented populations 
through habitat loss and hunting; and the 
indri Indri indri of eastern moist forest on 
Madagascar, threatened by habitat destruction 
as is the recently rediscovered Edward's 
pheasant Lophura ec/wards/ of Viet Nam. 

Role in carbon cycle 

Lowland evergreen broadleaf rainforests can 
have high above-ground biomass (Table 5.61, 
though not as high as some giant conifer 
forests. Soil carbon, however, is relatively low 
in most tropical moist forests, with the 
exception of the peat forests of Southeast Asia 
and some swamp forests. On this basis it has 
been estimated that the remaining tropical 
moist forests store over 300 petagrams of 
organic carbon, or about one fifth of global 
terrestrial organic carbon. They account for 



90 WORLD ATLAS OF BIODIVERSITY 



Map 5.4 

Biodiversity at country 

level 

Country-level biodiversity. 
represented by an index 
based on species diversity 
in the four terrestrial 
vertebrate classes and 
vascular plants, adjusted 
according to country area. 
Countries at the high end of 
the scale have a higher 
value of the index than 
w/ould be expected on area 
alone. The index is 
unreliable for smaller 
countries (such as Togo and 
Luxembourg in this plot]. 

Note: To reduce ambiguity 
Alaska (United States! has for 
the purposes of this map 
been assigned to the same 
class as adjacent Canada 
rather than the conterminous 
United States, 

Source: Based on national biodiversity 
indices developed by UNEP-WCMC; see 
Appendix 5. 



Diversity 



I 



High 



Low 



nearly a third of global terrestrial annual 
net primary production (Chapter II, and are 
therefore key to the global carbon cycle and in 
regulating global climate. 

Use by humans 

Tropical hardwood species contribute almost 
one fifth of world industrial roundwood 
production^ Of the several thousands of 
species [in more than 200 families in the 
phylum Anthophytal that show commercial 
potential, a few hundred may be found in 
international trade. Important families 
include Dipterocarpaceae, with species of 
meranti and balau Shorea. and keruing 
Dipterocarpus, Meliaceae, with mahogany 
Swietenia and Khaya, and cedar Cedrela and 
Toona; and Leguminosae with rosewood 
Dalbergia and Pterocarpus. 




# 



The exploitation of tropical hardwood from 
moist forests has been the subject of much 
publicity in the past two decades, coinciding 
as it has done with increased rates of 
deforestation and forest degradation. Timber 
supplies from some countries are now widely 
exhausted, generating openings for other 
countries to take over as suppliers. A few 
major producers, however, continue to 
dominate supply. Indonesia, Malaysia, Brazil 
and India accounted for 80 percent of tropical 
log production in International Tropical 
Timber Organization (ITTOI countries in 1999- 
2000-. 

Important non-timber products from tropi- 
cal moist forests include rattans, which are the 
second most important source of export earn- 
ings from tropical forests. A few other craft 
products and some medicinal products, such as 



Terrestrial biodiversity 91 




the bark of Prunus africana. are significant in 
international trade. Brazil nuts Berthoiettia 
excelsa and native rubber Hevea brasiliensis 
are other extractive products from natural 
tropical moist forests that are important in 
international marl<ets, and many tropical moist 
forests provide fruit, bushmeat and other 
products for local markets. 

Other ecosystem services 

Like other forest types, tropical moist forests 
often play an important role in soil and water- 
shed protection. The high rainfall regimes of 
the humid tropics mean that exposed soil is 
particularly liable to leaching of mineral 
nutrients and to erosion. Montane forests, and 
especially cloud forests, serve to intercept 
and store water, thus regulating local and 
regional hydrological cycles". Lowland forests 



are also important in hydrological cycles; it 
has been estimated that about half the rainfall 
in the Amazon Basin is derived from water 
recycled by forest transpiration"'. 

Regenerating forest, or forest fallow, is an 
important part of the cycle of shifting culti- 
vation, which is vital for restoring fertility to 
areas that have been previously cultivated. 
This regeneration can only take place if 
nearby forest cover is adequate to provide a 
source of propagules. 

Tropical dry forests 

Distribution, types and characteristic taxa 

Tropical dry forests are characteristic of areas 
in the tropics affected by seasonal drought. 
Such seasonal climates characterize much of 
the tropics, but less than A million km' of 
tropical dry forests remain. The seasonality of 



92 WORLD ATLAS OF BIODIVERSITY 



rainfall is usually reflected in the deciduous 
habit of the canopy trees, with most being 
leafless for several nnonths of the year 
However, under some conditions, e.g. less 
fertile soils or less predictable drought 
regimes, the proportion of evergreen species 
with leaves highly resistant to water loss 
increases I'sclerophyllous' forest). Thorn 
forest, a dense forest of low stature with a 
high frequency of thorny or spiny species, is 
found where drought is prolonged, and 
especially where grazing animals are plenti- 
ful. On very poor soils, and especially where 
fire is a recurrent phenomenon, woody 
savannahs develop" " (see sparse trees and 
parkland' below). 

Perhaps the best-known tropical dry 
forest tree species is teak Tectona grandis 
IVerbenaceae). a deciduous hardwood charac- 
teristic of the seasonal forests of south 
and Southeast Asia, widely exploited for 
furniture and other uses, and now an important 
plantation species. In Southeast Asia the 
dipterocarps Shorea and Dipterocarpus, and 
Lagerostroemia iLythraceae). and a number of 
species of legumes ILeguminosae) are also 
important components of seasonally dry 
forests'. In Africa, dry forests occur both north 
and south of the equatorial rainforests. In the 
north they are characterized by Afraegete 
IRutaceae). Diospyros (Ebenaceae), Kigelia 
IBignoniaceae) and Monodora lAnnonaceae). 
among other taxa, while in the south the 
characteristic genera are Entandophragma 
iMeliaceae). Brachystegia ILeguminosae), 
Diospyros, Parinari IChrysobalanaceae). 
Syzigium (Myrtaceae) and Cryptosepatum 
ILeguminosae)"". 

In the neotropics. tropical dry forests 
occur in Yucatan and Pacific slopes of Central 
America, on the leeward sides of Caribbean 
islands, in Venezuela. Caribbean Colombia, in 
northeast Brazil and in the Chaco region of 
Bolivia and Paraguay (see Map 5.7). The neo- 
tropical dry forests are quite rich in species 
and include Leguminosae, Bignoniaceae, 
Rubiaceae. Sapindaceae. Euphorbiaceae. 
Falcourtiaceae. and Capparidaceae as the 
families with the largest numbers of 
species. Important genera include Tabebuia 
(Bignoniaceae). Trichitia (Meliaceae). Eryth- 



roxytum (Erythroxylaceae). Randia (Rubi- 
aceae). Capparis (Capparidaceae), Bursera 
(Burseraceae). Acacia (Leguminosae) and 
Coccoloba (Polygonaceae)". 

Structure and ecology 

Tropical dry forests are generally of lower 
stature than moist forests, with canopy 
heights ranging from only a few meters to 
30 m or occasionally AO m". The taller forests 
have multilayered canopies. Dry forests tend 
to have more small trees than moist forests 
and a lower above-ground biomass. The trees 
have a greater proportion of their total 
biomass below ground, as more extensive 
root systems help the trees to obtain water 
from the soil and avoid drought. Dry forests 
have a much lower incidence of epiphytes 
than wet forest, and tend to have both higher 
frequencies and higher diversity of vines 
and lianas". 

Plants with specialized mechanisms for 
avoiding drought or conserving water are an 
important feature of these forests. The 
proportion of deciduous tree species is 
thought to increase steadily with decreasing 
annual rainfall, but factors such as the sub- 
strate and the between-year variation of 
seasonal rainfall patterns are also important. 
Many species have water storage tissues such 
as succulent stems or tubers, and specialized 
photosynthetic mechanisms that conserve 
water are especially common among the 
epiphytes. Most tropical dry forest trees tend 
to flower and sometimes to re-leaf before the 
end of the dry season, and stored water within 
the plant is essential to this pattern. 

As in tropical moist forests, environ- 
mental heterogeneity is linked with in- 
creased species diversity. Gallery areas 
along water courses are one source of such 
variation, and they serve as a refuge for 
animals during the dry season". Termite 
mounds provide an important source of 
environmental variation in African dry 
forests, adding to local topography and 
supplying high-nutrient microenvironments 
to the system, increasing tree species 
richness by ^0-100 percent". Fire is also a 
major factor determining the dynamics and 
extent of tropical dry forests. 



Terrestrial biodiversity 93 



•^Iri. 



Biodiversity 

Though of lower species richness than 
tropical moist forests, dry forests still have 
appreciably more tree species than most 
temperate forests. The richest neotropical dry 
forests, which are not the wettest ones but 
those in western Mexico and in the Chaco of 
southeast Bolivia, have around 90 woody 
species per 0.1 hectare sample". 

Although a comprehensive assessment 
has yet to be made, dry forests are thought to 
have high rates of plant species endemism 
relative to wet forests in the tropics". Sixteen 
percent of the plant species of the Chamela 
dry forest in western Mexico are local 
endemics, and 20 percent of the flora of 
Capeira, Ecuador, are endemic to western 
Ecuador". Many of the dipterocarps in 
Thailand's seasonal forests are national 
endemics and distinct from the species in the 
country's moist forests'. 

Vertebrate species diversity is lower in dry 
forests than in moist forests, but many dry 
forests have high rates of endemism among 
mammals, especially among groups such as 
insectivores and rodents, characterized by low 
body weights, low mobility and short gener- 
ation times". Among neotropical dry forests, 
those of Mexico and the Chaco have the 
highest numbers of mammal endemics (26 
and 22 respectively"). Remaining areas of dry 
forest are often important refuges for once 
widespread species. The Gir forest of Gujarat 
[India] contains the only population of Asiatic 
lion Panthera leo persica, once found 
throughout much of southwest Asia; the dry 
forests of western Madagascar are inhabited 
by around 40 percent of the island-endemic 
lemurs; some, such as red-tailed sportive 
lemur Lepitemur ruficaudatus, are almost 
entirely confined to this habitat. Invertebrate 
species richness tends to be poorly known, but 
in groups such as lepidoptera and hymenop- 
tera richness in some dry forest areas may be 
comparable to adjacent wet forest*'. 

Because of their high degree of endem- 
ism and because degradation and conversion 
have progressed further than in wet forests, 
the biota of tropical dry forests are often 
highly threatened. Hunting, especially for the 
wildlife trade, and habitat conversion are 




important pressures on dry forest animal 
species. 

Threatened dry forest species include Spix's 
macaw Cyanospitta spixii, which is nearly 
extinct globally as a result of trapping and 
habitat loss in Brazil's northeastern caatinga 
region; the Chacoan peccary Catagonus 
wagneri, which was rediscovered in the Gran 
Chaco of central South America during the 
1970s and is threatened by overhunting, 
habitat loss and disease; Verreaux's sifaka 
Propithecus verreauxi of western Madagascar. 
which is at risk from loss of spiny and gallery 
forest habitat; and the Madagascar flat-tailed 
tortoise Pyxis planicauda, which is restricted 
to the western Andranomena forest of 
Madagascar and is believed to be declining as 
a result of habitat destruction. 



Many of ihe endemic 
lemurs of Madagascar are 
found in dry forest. 



M WORLD ATLAS OF BIODIVERSITY 



.^5^=3' -^"a^s^ 



Map 5.5 

Flowering plant family 

density 

Global diversity of flowering 
plants represented as a 
density surface derived 
from distribution maps for 
284 non-aquatic plant 
families. Scale indicates 
number of families present, 
up to a maximum of 182, 
divided into 15 classes. 



Source: Compiled primarily frorr 
Heywood'°° by UNEP-WCMC 



Density 



High 





Low 



Role in carbon cycle 

Their lower biomass means that tropical dry 
forests represent a smaller reservoir of stored 
carbon per unit area than the other forest types 
discussed so far With a total biomass ranging 
from 98 to 320 metric tons per hectare" and soil 
carbon storage in the region of 100 metric tons 
per hectare''. It is unlikely that relatively undis- 
turbed tropical dry forests store more than 250 
metric tons of carbon per hectare (Table 5.61, 
This, combined with the fact that little Intact 
tropical dry forest remains worldwide, suggests 
that the total contribution of seasonally dry 
tropical forests to global carbon storage is far 
less than that of other forest types. 

Use by humans 

Notable among the economically important 
species of seasonally dry tropical forests is teak 



Tectona grandis, which accounts for about 1 
percent of reported global tropical timber 
exports. More than ten species of Thailand's 
seasonal forests are significant for the timber 
trade', with timber species such as mahogany 
Swietenia and several species of Tabebuia 
(Bignoniaceae) characteristic of neotropical 
dry forests. The southern dry forests of Africa 
also contain useful timber species such as 
Entandrophragma spp. Tropical dry forests also 
provide large quantities of fuelwood for local 
populations. A number of food plants are native 
to tropical dry forests and medicinal uses have 
been reported for many dry forest plant 
species*'. Craft products are also Important. 

Other ecosystem services 

Protection of relatively fragile soils is a vital 
ecosystem service provided by tropical dry 



Terrestrial biodiversity 95 



€^ 






forests. Rains may be intense during the wet 
season, and erosion can be a severe problem in 
tropical dry forest areas, where soils are often 
thin and soil formation processes slow". 
Tropical dry forests may also be important 
resources for native pollinators as well as 
nectar sources for domestic bees. Many dry 
forest trees produce conspicuous flowers with 
specialist pollination mechanisms. Their mass 
flowering provides a major nectar resource for 
pollinating insects at the end of the dry season 
when other such resources may well be 
limited" '^ Honey production is one of the liveli- 
hoods being promoted for local communities in 
dry forest areas in Mexico and elsewhere. 

Sparse trees and parkland 

Sparse trees and parl<land are forests with 
open canopies of more than 10 percent crown 



cover They occur principally in areas of 
transition from forested to non-forested 
landscapes. The two major zones in which 
these ecosystems occur are the boreal region 
and the seasonally dry tropics. 

At high latitudes, north of the main zone of 
boreal forest or taiga, growing conditions are 
not adequate to maintain a continuous closed 
forest cover, so tree cover is both sparse and 
discontinuous. This vegetation is variously 
called open taiga, open lichen woodland or 
forest tundra". It is species poor, has high 
bryophyte cover, and is frequently affected by 
fire. It is important for the livelihoods of a 
number of groups of indigenous people, inclu- 
ding the Saami and some groups of Inuit. 

In the seasonally dry tropics, decreasing 
soil fertility and increasing fire frequency are 
related to the transition from closed dry forest 



96 WORLD ATLAS OF BIODIVERSITY 



The large savannah 
vertebrates of Africa are 
largely absent from other 
continents. 



through open woodland to savannah. The 
open woodland ecosystems include the more 
open Srachysteg/a and /sofaer//n/a woodlands 
of dry tropical Africa and parts of both the 
caatinga and cerrado vegetations of Brazil''. 
Open woodlands in Africa are more species 
rich than either closed dry forest or savannah. 
The cerrado supports a high diversity of 
woody plants, though many of them are of 
shrubby habit. 

Animal diversity is generally low in forest 
tundra; few species are restricted to this 
habitat, many also occurring in boreal forest 
or tundra proper The sparsely wooded tropi- 
cal savannahs are generally more species 
rich than temperate forests or grasslands". 
Wooded savannahs vary greatly. Those of 
America are relatively species poor, while 
African savannah sometimes attains a rich- 
ness not far below rainforest in the same 
continent. Sparsely wooded areas in Australia 
are amongst the richest wildlife habitats on 
the continent, sometimes more so than 
adjacent wet forests''''"'. The large savannah 
vertebrates present in such high diversity 
in Africa are largely absent from other 




wmsii^^ 



Air^itlsiJfiilf s'WxHft' 



continents''' (see Chapter 41. The density and 
biomass of tropical savannah soil inver- 
tebrates (mostly earthworms, ants and 
termitesl is generally lower than temperate 
grasslands, but greater (at least in biomassl 
than tropical rainforests'". 

Species of conservation concern include 
the black rhinoceros Diceros bicornis, threat- 
ened primarily by hunting for their horn, and 



the golden-shouldered parrot Psephotus 
chrysopterygius of northern Queensland 
(Australia!, threatened by the burning of 
seeding grasses during the breeding season 
and predation by feral cats. 

Forest plantations 

Forest plantations, generally intended for the 
production of timber and pulpwood, increase 
the total area of forest worldwide. FAQ" 
estimates that forest plantations covered 187 
million hectares in 2000, of which Asia 
accounted for 62 percent. This represents a 
significant increase from the 1995 estimate of 
124 million hectares. Commonly mono- 
specific and/or composed of introduced tree 
species, plantation forests tend to be less 
valuable as a habitat for native biodiversity 
than are natural forests. However, they can be 
managed in ways that enhance their habitat 
value. Plantations are also important pro- 
viders of ecosystem services, such as main- 
taining nutrient capital and protecting water- 
sheds and soil structure as well as storing 
carbon. They may also alleviate pressure on 
natural forests for timber and fuelwood 
production. 

In some countries, wood from plantation 
sources makes up a significant portion of the 
industrial wood supply. For example. New 
Zealand is more than self-sufficient in wood 
production based on plantations". New forest 
plantation areas are increasing globally at a 
rate of 4.5 million hectares per year, with 
particularly high rates of increase in Asia and 
South America'''. In future, forest plantations 
are likely to play an increasingly important 
role in mitigation of greenhouse gases, as 
encouraged by the Kyoto Protocol. 

Changes in forest cover 
About half of the forest that was present 
under modern (i.e. post-Pleistocene) climatic 
conditions, and before the spread of human 
influence, has disappeared, largely through 
the impact of human activities. The spread of 
agriculture and animal husbandry, the har- 
vesting of forests for timber and fuel, and the 
expansion of populated areas have all re- 
duced forest cover. The causes and timing of 
forest loss differ among regions and forest 



Terrestrial biodiversity 97 



> 



types, as do the current trends in ctiange in 
forest cover 

The temperate forests of western Europe 
have diminished by far more than the 
50 percent estimated for forests globally. 
Much of this deforestation occurred between 
7 000 and 5 000 years ago as Neolithic 
agriculture expanded'-. The expansion of 
human populations and increasing demand 
for fuel during classical times and the Middle 
Ages put further pressure on European 
forests. Between Neolithic times and the late 
11th century, forest cover In what is now the 
United Kingdom decreased by 80 percent. 
As European forests dwindled they became an 
Increasingly valuable resource that was more 
carefully managed. Forest cover stabilized 
during the 19th century in much of western 
Europe In response to both improved man- 
agement and reduced demand for forest 
products lowing to the increasing use of fossil 
fuels and changes in construction materials). 
Since the early 20th century, forest cover In 
Europe has expanded, often through the 
establishment of conifer plantations. 

In eastern and central Europe and in 
Russia, forest clearance accelerated during 
the 16th and 17th centuries as sedentary 
agriculture expanded. One estimate" sug- 
gests that around 1 million km' of forest had 
been cleared In the former Soviet Union up to 
1980. Timber exploitation continues to drive 
forest clearance In the coniferous forests of 
Siberia and parts of eastern Europe. 

In North America, Indigenous groups had 
impacts on the forests from at least 12 000 
years ago, but most forest clearance took 
place after European settlement. Forest cover 
In eastern North America reached Its 
minimum around 1860, but then increased 
with the westward movement of the agri- 
cultural frontier and subsequent urbanization 
and Industrialization. Forests west of the 
Appalachians suffered the most severe 
impacts In the late 19th and early 20th 
centuries, but are still under pressure from 
demand for timber and pulp. 

In Oceania, just as in North America, 
Indigenous groups had significant Impacts on 
the forests before the arrival of Europeans. 
This was especially true of the aboriginal 



Region 


Total forest area 


Forest cover 


change 1990-2000 




In 2000 ImlUlon ha| 


Annual change 


Annual rate of 






[thousand ha| 


change l%| 


Africa 


650 


-5 262 


-0.78 


Asia 


548 


-364 


-0.07 


Europe 


1039 


881 


0.08 


North and Central America 549 


-570 


-0.10 


Oceania 


198 


-365 


-0.18 


South America 


886 


-3 711 


-0.41 


World total 


38&9 


-9391 


-0.22 



use of fire In Australia, but European colon- 
ization greatly increased the rate of forest 
conversion. Over 230 000 km' of forest and 
120 000 km- of woodland in Oceania are 
estimated to have been converted to cropland 
between 1860 and 1980". 

In tropical Asia, Africa and Latin America, 
large-scale deforestation was precipitated 
by European colonial activities. Including 
agriculture and timber exploitation. It Is 
estimated that more than 1 million km-' of 
forest and a similar amount of woodland in 
tropical developing countries were converted 
to cropland between 1860 and 1980'''. The bulk 
of this conversion was in south and Southeast 
Asia, where forest area declined by 39 percent 
from 1880 to 1980. 

Globally, tropical dry forest has lost the 
greatest proportion of Its original area of 
the four major types of closed forest, nearly 
70 percent. About 60 percent of the original 
area of temperate broadleaf and mixed 
forests has disappeared, and tropical moist 
and temperate needleleaf forests have lost 
about 45 percent and 30 percent of their 
original area, respectively. 

Current trends in change in forest cover, 
which are shown in Table 5.8, reveal that the 
rates of deforestation continue to be high In 



Table 5.8 

Estimated annual change 

in forest cover 1990-2000 

Note: Figures refer to natural 
forests and plantation forests 
combined. 



Table 5.9 

Global protection of 

forests within protected 

areas in lUCN categories 

l-VI 

Note: Forest areas and 
protection as assessed in 
1999. 

Source: UNEP-WCMc". 



Forest type 




Global protection 


m 




Ikm^l 




i%i ^^ 


Temperate and boreal needleleaf 


675 470 




5.4 


Temperate broadleaf and mixed 


457 535 




7.0 


Tropical moist 


1 382 004 




12.2 


Tropical dry 


413 524 




11.2 


Sparse trees and parldand 


274 401 




5.8 


Total 


3 202 934 




8.3 



98 WORLD ATLAS OF BIODIVERSITY 



Fuelwood and charcoal 
consumption more than 
doubled between 1961 
and 1991. 



the developing countries of the tropics, in both 
absolute and proportional terms. In contrast, 
temperate countries are losing forests at 
lower rates, or indeed showing an increase in 
forest area". 

Pressures on forest biodiversity 

The principal pressures on forests and their 
biodiversity are conversion to other landuses, 
principally forms of agriculture, and logging. 
Conversion of forest to agriculture is the main 
cause of tropical moist forest loss. This is 
largely due to expanding populations and the 
use of shifting cultivation at an intensity that 
does not permit adequate fallow periods. 
Government resettlement programs that have 
moved large numbers of poor farmers have 
increased the rate of land colonization and 
clearance In parts of Southeast Asia and Latin 
America. In some areas, land has been 




converted to ranching principally as a means 
of gaining title In order to permit speculation 
in land values. Thus, population growth, 
poverty and inequitable land tenure are 
among the causes underlying deforestation by 
conversion to agriculture. 

Timber extraction puts great pressure on 
biodiversity in both tropical and temperate 
forests. Global consumption of industrial 
roundwood was more than 1 521 million m' in 
1998" and, on current trends, is projected to 
continue to rise. Although some timber 
species are naturally abundant, a factor that 
can help ensure their survival from com- 
mercial exploitation, many have suffered 



extensive and irreversible population and 
genetic losses. Furthermore, rare species 
that are indistinguishable In the field from 
their commercially important relatives are in 
danger of extinction through overexploitation. 
This particular problem exists, for example, 
among the dipterocarp groups meranti, balau 
and l^eruing. A tew hundred species may be 
traded under these names, and a significant 
proportion of these are geographically and 
ecologically restricted and so at high risl< 
of extinction. 

Furthermore, logging operations create 
access to forest areas that may otherwise 
have remained Isolated. This improved access 
facilitates hunting and other activities that 
exert pressure on forest biodiversity, and may 
ultimately lead to colonization and conversion 
of the land to agricultural use. There is also 
strong evidence that logging can increase the 
probability of wildfire In temperate forests 
and even in tropical moist forests not usually 
subject to burning". 

Particularly in the drier areas of the 
tropics, fuelwood extraction can have serious 
impacts on forests and open woodlands. 
Fuelwood and charcoal consumption more 
than doubled between 1961 and 1991, and is 
projected to rise by another 30 percent to 
2 395 million m= by 2010". About 90 percent of 
the consumption is In developing countries, 
but wood fuel may play an Increasing role In 
some developed countries. Increasing demand 
for wood still further". 

In addition to loss of area, forest conver- 
sion and logging lead to changes In the 
condition or quality of the remaining forest. 
These can include fragmentation of large 
areas of continuous forest. Tropical forest 
fragments are distinct from continuous for- 
ests in both ecology and composition". There 
are physical and biotic gradients associated 
with fragment edges, and forest structure 
undergoes radical change near the edges as a 
result of the impacts of wind and increased 
tree mortality. Some animal species are 
'edge-avolders' and decline in abundance In 
forest fragments, while others become more 
abundant. Some non-forest and even non- 
native species of plants and animals success- 
fully invade forest fragments but not con- 



Terrestrial biodiversity 99 



"^'ik 



tinuous forest. In addition to directly affecting 
canopy composition, removal of large timber 
trees may also affect the availability of seed 
for regeneration and may affect animal 
species that depend on the timber species. 

Other factors that affect forests and their 
biodiversity include acid rain and global 
climate change. So far. most of the effects of 
acid precipitation, which Is caused by indus- 
trial air pollutants, have been documented in 
temperate needleleaf forests and associated 
vi/aterways of Europe and North America. 
The likely impacts of global climate change 
on forests are still being debated, but there 
seems to be general consensus that the 
boreal coniferous forests are particularly 
vulnerable to both range restrictions and 
Increasing fire frequency". 

Another forest type that has been shown to 
be vulnerable to climate change is tropical 
montane cloud forest, which depends upon 
clouds to supply It with atmospheric moisture. 
Research has shown that the mean cloud 
base is moving upwards on tropical moun- 
tains as a result of climatic shifts. The forest 
species are not able to migrate at a 
comparable rate and. In any case, range shifts 
will be limited by the land area existing at 
higher elevations. Local extinctions in cloud 
forest amphibians, including the golden toad 
Bufo pengtenes assessed as critically 
endangered, have been attributed to climatic 
fluctuations that may be linked to long-term 
global climate change". 

NON-FOREST ECOSYSTEMS 

The parts of the Earth that are too cold, too 
dry or too severely affected by fire and/or 
grazing do not support forest or woodland 
ecosystems. However, as can be seen from 
Map 5.1, many of them do support active plant 
growth. Natural non-forest ecosystems in- 
clude tundra (both arctic and montanel, 
grasslands and savannahs, and shrublands. 
Less productive, but with unique elements of 
biodiversity, are the deserts and semi-deserts 
IMap 5.4). 

Tundra 

Tundra Is the vegetation found at high 
latitudes beyond the limits of forest growth; 



the same term is sometimes used for similar 
vegetation at high elevation at lower latitudes, 
but these may be distinguished as polar 
tundra' and 'alpine tundra', respectively. In 
the Arctic, polar tundra occurs north of the 
northern tree line, which Is determined by a 
number of climatic factors Including the 
summer position of arctic air masses" and 
the depth of permafrost (permanently frozen 



In polar tundra systems, 
temperatures are low for 
much of the year 




subsurface soil]. Similarly, alpine tundra 
occurs above the climatic tree line on 
mountains, and Its elevation varies in a 
complex fashion with latitude, continental or 
oceanic climate, and the maximum elevation 
and overall size of the massif™. 

The characteristics of polar and alpine 
tundra environments differ In many respects. 
In high-latitude polar tundra systems, 
temperatures are low for much of the year, 
while permafrost limits both drainage and 
root extension, and the growing season may 
last for as little as six to ten weeks. Rainfall is 
low. usually less than 200 mm per year, and 
at extreme latitudes may be so low that the 
environment Is described as polar desert. 
At high elevation In temperate regions 
temperatures may be similarly low. although 
permafrost Is rare. However, at high altitudes 
in the tropics, although low temperatures 
occur every night, high Insolation causes 
warming during the day so that adequate 
temperatures for active plant growth occur 



100 WORLD ATLAS OF BIODIVERSITY 



<~ ^3,-^^^^ 



Map 5.6 

Terrestrial vertebrate 

family density 

Global diversity of 
terrestrial vertebrates 
represented as a density 
surface derived from 
distribution maps for all 
350 non-aquatic families of 
mammals, birds, reptiles 
and amphibians. Scale 
indicates number of 
families present, up to a 
maximum of 126, divided 
into 15 classes. 



Source: Compiled from multiple sources 
byUNEP-WCMC. 



Density 



Higti 



Lovw 



tfiroughout the year and the diurnal temper- 
ature range is large. 

Despite these environmental differences, 
polar and alpine tundra vegetation have some 
features in common. Both lack trees, but 
contain woody species growing in dwarf or 
prostrate forms, especially in locations with 
less extreme climates. As latitude or altitude 
increases, grasses, sedges, bryophytes and 
lichens increase in importance while shrubs 
decrease. Many plants have tussock or cush- 
ion growth forms. At extremes of latitude or 
elevation a high proportion of bare ground is 
characteristic. 

In the arctic tundra, plants cover 80-100 
percent of the ground", and cover decreases 
along the climatic gradient to polar desert. 
The important woody plants are birches 
Betula, willows Salix and alders Atnus. These 



#S7^ 





are all genera that occur as trees in 
temperate regions but as spreading, prostrate 
or dwarf forms, sometimes less than 20 cm 
tall, in tundra. Other shrub species are also 
important, including Dryas, Vaccinium and 
Empetrum. Interspersed with the shrubs and 
of increasing importance in more extreme 
sites are the sedges Carex and the cotton 
grasses Eriophorum, among other gramin- 
oids, and a high diversity of mosses and 
lichens. Biomass is often much lower above 
than below ground, and annual production is 
low. This combination means that the poten- 
tial of these ecosystems for recovery following 
disturbance is limited". 

Temperate alpine systems are charac- 
terized by many of the same taxa as the arctic 
tundra, but at lower latitudes other groups 
become important. At high altitude in the 



Terrestrial biodiversity 101 





"^ 



tropics, giant rosette plants are a notable 
feature of alpine communities. These dis- 
tinctive plants, whicfi have a number of 
morphological and physiological adaptations 
to the high insolation, large temperature 
fluctuations and desiccating conditions on 
tropical mountains, include Espeletia and 
Puya in the paramos of the Andes, and 
Senecio and Lobelia in the high mountains 
of Africa". 

In comparison with forested ecosystems, 
both polar and alpine tundra are relatively 
species poor It is estimated that species 
richness declines by a factor of between 
three and four between the boreal and arctic 
zones", and species richness in the polar 
desert is one fifth that of the tundra. The 
entire North American Arctic has a vascular 
flora of about 600 species". Bryophytes and 




lichens may add more than 300 additional 
species to the circumpolar flora. The tropical 
alpine systems are richer; Venezuelan 
paramos include more than iOO angiosperm 
species'', but this is still much lower than in 
surrounding forests. 

Because most arctic plant species have 
wide geographic distributions, few are of 
significant conservation concern, but many 
alpine areas are isolated by lowlands with 
contrasting climates so increasing endemism 
in many alpine floras. The floras of isolated 
mountains in North America have significant 
rates of endemism"; about 15 percent of the 
flora above the timber line in the Alps are 
locally endemic species" and about 80 percent 
of the flora at high altitude in East Africa and 
Ethiopia are endemic". Restricted distrib- 
utions tend to make species more vulnerable 



102 WORLD ATLAS OF BIODIVERSITY 



to extinction, while the harsh environment of 
alpine regions adds to the risl<. 

Animal species richness tends to be low; 
groups represented by several species in 
boreal forest are often reduced in diversity by 
up to one third in tundra habitat". In contrast, 
a few groups, particularly water birds and 
waders, are able to exploit the large numbers 
of Invertebrates found in tundra soil and can 
be both diverse and abundant". Although the 
species richness of the most common in- 
vertebrate groups (Collembola and oribatid 
mitesi decreases with increasing latitude, 
their total abundance may increase. 

There are relatively few globally threaten- 
ed species that are completely dependent on 
tundra. An exception Is one of the worlds 
most severely endangered species, the once 
abundant Eskimo curlew Numenius borealis 
of the Americas, which nests almost ex- 
clusively within this habitat. Two globally 
threatened bird species, Stellers [Polysticta 
stelleri] and spectacled elder [Somaleria 
fischeri] remain within the Arctic throughout 
the year Although low In number of species, 
the Arctic is home to most of the world's 
geese and calidrid sandpipers". 

Although the tundra accounts for less than 
2 percent of global annual net primary 
production (Table 1.11, the high below-ground 
biomass and soil carbon in arctic tundra 
means that it makes an important contribution 
to global carbon stocks. Total biomass in 
tundra communities of the Russian Arctic falls 
in the range 7-30 metric tons per hectare, of 
which some 60-70 percent is below ground". 
Sedge-moss communities in the North 
American Arctic may have 15-30 times as 
much biomass below ground as above 
ground", and three to eight times the amount 
of dead as live material may accumulate. 
Tundra soils may store around 200 metric tons 
of carbon per hectare^'. 

Because of Its Inhospitable climate, tundra 
is not subject to severe pressure for 
conversion for other landuses. However, Its 
lack of ecological resilience means that 
disturbances, e.g. those associated with 
settlements or long-distance pipelines, tend 
to have long-lasting effects. It is anticipated 
that the effects of global warming on the 



arctic tundra will be significant, as relatively 
large temperature increases are predicted 
for this zone". These will cause changes in 
the permafrost regime and decomposition 
of accumulated soil organic matter, which in 
turn will release additional carbon dioxide 
into the atmosphere. Evidence suggests that 
the period of active plant growth has recently 
lengthened in parts of North America. There 
is also evidence that species are already 
migrating northwards in response to climate 
change, so that arctic tundra Is likely to be 
compressed Into a much smaller area of 
remaining appropriate climate. 

Grasslands and savannahs 
Grassland ecosystems may be loosely defined 
as areas dominated by grasses (members of 
the family Gramlneae excluding bamboos] or 
other herbaceous plants, with few woody 
plants'". Grassland ecosystems are typically 
maintained by drought, fire, grazing and/or 
freezing temperatures™. In addition, they are 
often associated with soils of low fertility'". 
Savannahs are tropical ecosystems charac- 
terized by dominance at the ground layer of 
grasses and grass-like plants. They form a 
continuum from treeless plains through open 
woodlands to closed-canopy woodland with a 
grassy understorey. Some savannah areas 
therefore meet general definitions of grass- 
land (fewer than 10-15 woody plants per 
hectare"! while others meet the definition of 
woodland. Some polar and alpine tundra 
communities also meet the definition of 
grassland. 

Around 20 percent of the Earth's land 
surface (excluding Antarctica) supports 
grassland, with these regions differing greatly 
In naturalness'". Temperate grasslands make 
up approximately one quarter of this area, and 
savannahs the remainder (tvlap 5.81. The most 
extensive areas of temperate grasslands are 
the prairies of North America, the pampas 
and campos of southern South America and 
the steppes of central Europe, southwest and 
central Asia and Russia. Temperate grass- 
lands are sometimes divided Into formations 
of tall grass, mixed grass and short grass, 
which differ both floristically and ecologically; 
short-grass communities are usually associ- 



Terrestrial biodiversity 103 



•^% 



ated with drier climatic regimes™. Tropical 
grasslands and savannahs include the llanos 
of the Orinoco basin in Venezuela and 
Colombia, the cerrado of central Brazil, and 
the savannahs of tropical and subtropical 
Africa, in addition to many species of grasses, 
the sedges ICyperaceae) and many different 
groups of dicotyledonous herbs are also 
important. 

Key ecological factors in grasslands are 
grazing pressure and the effects of fire. Most 
natural grasslands had, at one time, large 
populations of native grazing mammals. 
These have been replaced to a great extent by 



environments, are often rich in organic matter 
and are therefore particularly vulnerable to 
conversion to cropland, v\/ith replacement of 
native grasses by their domestic derivatives 
Icerealsl and other plants". 

At very tine spatial scales, natural grass- 
lands can be extremely species rich. For 
example, a square meter of 'meadow steppe' 
in the former Soviet Union may have 40-50 
plant species"', a tall grass prairie remnant of 
less than 2 hectares may contain 100 species, 
and 250 hectares may contain 250-300 plant 
species". However, grassland communities 
tend to be similar over large areas, and 



The world's grasslands and 
savannahs support 
distinctive plant and animal 
communities. 




domesticated ungulates, which also exert a 
significant degree of grazing pressure [the 
magnitude depending on stocking densities). 
Grazing tends to increase abundance of less 
palatable species and to increase species 
richness in productive areas, or decrease it in 
less productive areas. At intermediate fre- 
quencies, fire tends to increase diversity and 
suppress invasion by woody species. Frequent 
fires favor grasses, which usually recover 
easily, whereas low fire frequency may allow 
the density of woody species to increase. 

These factors have important conse- 
quences for vegetation structure in grassland 
and savannah systems. A high proportion of 
plant biomass, in the form of roots and 
rhizomes, is located underground; there is a 
high turnover of those parts of the plant above 
ground". One important consequence of this is 
that grassland soils, especially in more humid 



structurally simple, so that at the landscape 
scale diversity is relatively low compared with 
tropical moist forest or t^^editerranean-type 
ecosystems. Grasslands tend to have low 
rates of endemism; however, the climatic 
and soil gradients within them have led 
to substantial ecotypic variation and high 
genetic diversity". 

The world's grasslands and savannahs 
support distinctive plant and animal com- 
munities. Although species diversity tends to 
increase towards the tropics, it tends to be 
moderate or low at the landscape scale and 
above. Little more than 5 percent of the 
world's mammal and bird species are pri- 
marily dependent on grasslands habitats'". 
All these ecosystems hold, or formerly held, 
an array of native herbivores, and these in turn 
support a number of high-profile mammalian 
and avian predators. 



104 WORLD ATLAS OF BIODIVERSITY 



, Box 5.5 Grassland bird trends 
k 

The figure illustrates the living planet index in a sample of 29 grassland birds from North 
Annerica 125 species! and Europe |4|. The clear trend over four decades is dc»/nward, and as 
with forest birds the separate continent samples show a similar overall pattern. 

UOr 



120 



100 



60 



40 



1970 1975 1980 

Source: UNEP-VtfCMC. from data collated lor Loh". 



1985 



1990 



Ttie savannafi communities of East Africa 
are typified by large herds of ungulate 
herbivores, including a remarkable diversity - 
more than 70 species - of antelope and other 
medium- to large-sized bovids. The biomass of 
ungulate herbivores here may rise to 30 metric 
tons per km*, which is the highest recorded in 
any terrestrial environment"'. Most grassland 
invertebrate biomass is found within the soil 
(most commonly nematodes, enchytraeid 
worms and mitesi and may be in the order of 
100 to 1 000 times as great as vertebrate 
biomass; soil invertebrate biomass above the 
soil is often dominated by Orthoptera"'. 

Ivlany grassland birds require targe areas 
of habitat to take full advantage of sparsely 
distributed food resources, and fragmentation 
of natural grasslands has made conservation 
of these wide-ranging species difficult'". 
An analysis of the location of bird species in 
the neotropics indicated that nearly 12 
percent of threatened birds 138 species! are 
confined to grasslands. In this region, the 
grasslands of southern Brazil and northern 
Argentina are particularly threatened as a 
result of agricultural improvement". In North 
America, grassland species have experienced 
the most consistent declines of any group of 



birds monitored in a national survey of 
breeding birds; available data suggest that 
there has been a constant decline in these 
species over the past 30 years'". 

Among notable threatened mammal 
species in grasslands are the greater one- 
horned rhinoceros Rhinoceros unicornis. 
associated with tall riverine grassland in 
northern India and southern Nepal, where it 
is threatened by poaching and further loss of 
its restricted habitat. Other targe mammals 
are the saiga antelope Saiga tatarica of 
central Asian grasslands, and the vicufia 
Vicugna vicugna in arid grasslands and plains 
in the Andes; both of these are threatened in 
parts of their range. Among birds, the short 
grassland habitat of the plains wanderer 
Pedionomus torquatus of southern Australia 
continues to be lost to cultivation; Rudd's lark 
Heterornirafra ruddi inhabits the montane 
grassland plateaus of eastern South Africa 
and is threatened by habitat degradation; 
white the lesser ftorican Syptieotides indica, 
of western India and Nepal, is critically 
endangered through toss of suitable grass- 
land habitat. 

Grassland biomass ranges from around 
2 to over 80 metric tons per hectare". As a 
rule, more than 50 percent of this is below the 
soil surface, and the ratio of root to shoot 
biomass ranges from betow five in warm 
humid grasslands to as much as 30 in the 
desert grassland of Ivlongotia". Usually more 
than half of the below-ground biomass is in 
the upper 10 cm of the soil, and soil carbon 
stocks may be as high as 250 metric tons per 
hectare-'. Annual production reaches 30-50 
metric tons per hectare per year in some 
warm humid grasslands. It is estimated that 
grasslands and savannahs together account 
for about a third of global terrestrial net 
primary production (Table 1.1], so despite their 
relatively low biomass grasslands play an 
important rote in the global carbon balance. 

Because the high below-ground biomass 
tends to increase the fertility of their soils, 
grasslands have been subject to high rates of 
conversion to agriculture. Less than half of 
North American grasslands remain in natural 
or semi-natural states"', the steppes of the 
former Soviet Union have been extensively 



Terrestrial biodiversity 105 



•^% 



irrigated and converted to agriculture and 
much of the pampas has been converted to 
agriculture or grazing land. Some anthropo- 
genic grassland consists of short-term 
monospecific sown pasture, VKhile some 
areas support species-rich semi-natural 
grassland created over centuries by pastoral- 
ists in conjunction with livestock grazing. 

Domestic livestock grazing is the most 
extensive human use of unconverted or 
anthropogenic grassland ecosystems, and 
of most arid or semi-arid ecosystems. 
Livestock have an impact on ecosystems 
through trampling, removal of plant biomass, 
alteration of plant species composition 
through selective grazing, and competition 
with native species. The impact of this on the 
biological diversity of these ecosystems has 
been variable. In some areas where the 
native vegetation is well adapted, the impact 
on plant species diversity has been relatively 
small. In other areas, where vegetation has 
not evolved in the presence of hooved 
herbivores, the changes have been great. 
Sometimes, particularly in tropical and semi- 
tropical grasslands, the dominant com- 
ponent of vegetation has shifted from grass 
to woody plants. 

Overgrazing can lead to reduction in plant 
cover, loss and degradation of soil, and 
invasion by non-native plant species. In 
almost all cases, wild animal diversity has 
been greatly affected (mostly through compe- 
tition and hunting, but also through spread of 
pathogens), so that the biomass of domestic 
livestock greatly exceeds that of native wild 
herbivores. In some areas, feral species (e.g. 
rabbits, camels, donkeys, horses, goatsi may 
also have a marked impact on natural or 
semi-natural ecosystems. 

Shrublands 

Shrub communities, where woody plants, 
usually adapted to fire, form a continuous 
cover, occur in all parts of the world with 200- 
1 000 mm of rainfall". In more arid areas 
including some semi-desert systems, shrubs 
are the dominant life form, but cover is 
discontinuous. The most distinctive and best- 
known shrublands are those of Mediterranean 
climate regions. 



Mediterranean climates are typified by 
cool, wet winters and warm, or hot, dry sum- 
mers. However, no single climatic or bio- 
climatic definition of a Mediterranean eco- 
system has yet been established, so that 
these areas remain rather loosely defined. 
Mediterranean ecosystems encompass a 
wide range of habitat types including forest, 
woodland and grassland, but are typified by 
a low, woody, fire-adapted sclerophyllous 
shrubland (maquis, chaparral, fynbos, malleel 
on relatively nutrient-poor soils. These sys- 
tems occur in five distinct parts of the world: 
the Mediterranean basin; California (United 
States); central Chile; Cape Province (South 
Africa): and southwestern and south Australia. 
Each of these regions occurs on the west 
side of a continent and to the east of a cold 
ocean current that generates winter rainfall. 
They cover around 2.5 million km' in total, 
or between 1 percent and 2 percent of the 
Earth's surface (according to definition). More 
than two thirds of the total Mediterranean- 
type ecosystem area is found within the 
Mediterranean basin. 



'^^^^^r 


Approximate area 


Total no. of ' 




[million km'l 


olant soecies | 


Cape Province, South Africa 


0.09 


8 550 


Southwestern Australia 


0.31 


ca 8 000 


California 


0.32 


5 050 


Chile 


O.U 


ca2100 


Mediterranean basin 


1.87 


25 000 



Differences in vegetation structure between 
regions are in part a consequence of differ- 
ences in the annual distribution of rainfall. 
In South Africa the sclerophyllous fynbos 
community contains an abundance of erica- 
ceous species as an understorey to low 
broader-leaved shrubs including members of 
the Proteaceae and Myrtaceae". 

The Australian heaths are structurally 
similar, with Epacridaceae replacing Ericaceae. 
Californian shrublands, known as chaparral. 
are characterized by Adenostoma (Rosaceae) 
and a high richness of Arctostaphylos species 
and other members of the Ericaceae. The 
shrublands, or matorral, of Chile include many 



Table 5.10 

Estimated plant species 
richness in the five 
regions of Mediterranean- 
type climate 

Source; UNEp". 



106 WORLD ATLAS OF BIODIVERSITY 



Map 5.7 

Current forest distribution 

Map adapted from tfie 
global landcover 
classification developed by 
the University of Maryland. 
The Maryland classification 
includes 13 classes and 
was based on AVHRR 
remote-sensing data with a 
spatial resolution of 1 km. 
The map shown was 
derived by reclassifying the 
Maryland landcover data to 
accord with an ecologically 
based classification of five 
major forest types. For 
presentation purposes the 
data have here been 
generalized to a i-km grid. 

Source; Data from University of 
Maryland Global Land Cover Facility For 
full description see Hansen . 



/ 



Forest type 




Tropical moist 

Tropical dry 

Temperate broadleaf and mixed 

Temperate and boreal needleleaf 

Sparse trees and parkland 



of the same genera as those of California, while 
in the Mediterranean basin itself Ericaceae, 
Cistaceae, Leguminosae and Oleaceae are 
all important. 

Species richness in Mediterranean-type 
ecosystems, particularly among plants, is 
generally high - approaching values for 
moist tropical forest areas - and levels of 
endemism are also very high. Among the five 
Mediterranean-type ecosystems, species 
richness appears highest on the poorer soils 
of South Africa and southwest Australia 
(Table 5.101, and lower on the richer soils of 
California, Chile and the Mediterranean 
basin^'. Countries around the Mediterranean 
Sea hold some 25 000 vascular species 
(about 10 percent of all vascular plants! of 
which around 60 percent are endemic to the 
Mediterranean region. 




sV 



The remaining four Mediterranean-type 
ecosystem regions are all considered to hold 
a disproportionately high floristic diversity in 
relation to their area". 

At fine scale, mean plant richness in the 
fynbos of South Africa is moderate, i.e. around 
16 species per m*. but many species have 
small ranges, and there is a uniquely high 
turnover in the species composition of plant 
communities along ecological and geogra- 
phical gradients. At landscape scale, richness 
accordingly rises to very high values, for 2 256 
species occur in 471 km' on the Cape 
peninsula and the entire Cape floristic region 
(including some non-fynbos vegetation) holds 
some 8 550 species, about 70 percent of which 
are endemic. 

The Mediterranean-type ecosystems in 
general have a relatively high proportion of 



Terrestrial biodiversity 107 




their species categorized as ttireatened. 
The Cape flora, largely within a Mediterranean- 
type ecosystem, occupies only h percent of the 
land area of southern Africa, but accounts for 
nearly 70 percent of the region's threatened 
plant species. About one third of the natural 
vegetation has been transformed by human 
activity; the remaining natural vegetation is at 
risl< from a number of invasive introduced 
woody plants, and the effects of an introduced 
ant (that suppresses native seed-storing ants 
and thus renders seed liable to destruction 
by rodents or firel. Around 10 percent of the 
Californian flora is considered threatened 
(equivalent to approximately one quarter of 
all threatened plants in the United States). 
In Australia, heath habitats, primarily in the 
southwest Mediterranean-type ecosystem 
region, rank third after 'woodland' and 'scrub' 



in numbers of 'endangered' category plants. 
Given their much smaller extent, this indi- 
cates that a far higher proportion of their flora 
is threatened than in either woodland or 
scrub habitats. 

Vertebrate diversity tends to be lower than 
in plants. To take an example, in the Cape 
Mediterranean-type ecosystem, reptile diver- 
sity IS only moderate while bird and mammal 
diversity is relatively low. The absence of large 
mammals in California and the Mediterranean 
basin may be linked to overhunting by 
humans during the late Pleistocene". 

Several threatened animal species rely on 
shrubby or scrub habitat. The Iberian lynx 
Lynx pardinus, found in the light woodland 
and maquls of Spain and Portugal, where 
habitat loss and hunting have led to decline, Is 
possibly the most threatened cat species. 



108 WORLD ATLAS OF BIODIVERSITY 



Map 5.8 

Non-forest terrestrial 

ecosystems 

Map adapted from tfie 
global landcover 
classification developed by 
the University of Maryland. 
The Maryland classification 
includes 13 classes and 
was based on AVHRR 
remote-sensing data with a 
spatial resolution of 1 km. 
The map shown was 
derived by reclassifying the 
Maryland landcover data to 
accord with a highly 
generalized classification of 
non-forest terrestrial 
ecosystem types. For 
presentation purposes the 
data have here been 
generalized to a 4-km grid. 

Source; Data from University of 
Maryland Global Land Cover Facility. For 
full description see Hansen . 




The riverine rabbit Bunotagus monticutaris of 
South Africa is restricted to a small area of 
riverine bush in the central Karoo, where it is 
threatened by further loss of this habitat to 
agriculture. Among birds, the island cisticola 
Cisticola haesitatus is endemic to the island 
of Socotra in the western Indian Ocean, where 
it is threatened by loss of light scrub and 
grassland habitat, possibly through over- 
grazing by goats. 

Mediterranean-type shrublands are not 
notably high in either biomass or net primary 
production, Biomass at mature fynbos sites is 
typically 15-16 metric tons per hectare, and in 
chaparral may be twice that". Combined with 
their relatively low rates of primary production, 
related to both climate and soil fertility factors, 
the incidence of fire tends to reduce the 
accumulation of carbon in these ecosystems 




and their soils. Total carbon storage is probably 
between 100 and 150 metric tons per hectare 
in Mediterranean-type shrublands''. 

The Mediterranean basin itself has for 
many centuries been subject to intense 
human activities, including forest clearance 
and grazing, such that little genuinely natural 
vegetation remains. It has been suggested 
that the plant diversity is locally high because 
of the number of species that have evolved 
as components of successional vegetation in 
response to frequent disturbance, in other 
Mediterranean-type shrublands. expanding 
human populations and conversion of land to 
agricultural or residential use are important 
pressures. These changes are often accom- 
panied by changing fire and grazing regimes, 
and both these changes tend to facilitate 
invasion by non-native plant and animal 



Terrestrial biodiversity 109 




'■'^ i. 



%r 









k^ 




species, which threaten native species pop- 
ulations, especially in California and South 
Africa™. 

Deserts and semi-deserts 

Nearly 10 million km' of the Earths land area 
is hyperarid, or true desert, where rainfall is 
extremely low and unpredictable in space and 
time, so that in some years none falls at all. 
These areas have a ratio of rainfall to 
potential evapotranspiration (P/PETI of less 
than 0.05". The Sahara desert alone makes 
up nearly 70 percent of the world hyperarid 
zone. Other extensive areas are found in the 
Arabian Peninsula and central Asia, with 
smaller areas in southwest Africa, the Horn of 
Africa, western South America and western 
North America. In semi-deserts, areas with 
less arid climates, the vegetation is usually 



more substantial than in deserts, but covers 
no more than 80 percent of the ground. 
Temperate deserts and semi-deserts cover 
nearly 6 million km' in Eurasia and North 
and South America". Polar regions and some 
high mountain areas with a permanently cold, 
dry climate also meet the definition of desert, 
but have completely different ecological 
characteristics from true drylands and are not 
usually considered with them. 

Plant cover in desert ecosystems ranges 
from areas without vegetation to areas with 
low densities of small shrubs and perennial 
grasses, often with populations of annuals 
that vary in density depending on seasonal 
precipitation. Deserts are often character- 
ized by short periods of relatively high pro- 
ductivity during rainy periods, interspersed 
with long periods of very low productivity. As 



110 WORLD ATLAS OF BIODIVERSITY 



Desert species show a 
wide range of adaptations 
to extreme environments. 



Dating back some 
15 million years, 
ttie Namib appears to be 
Earth's oldest desert. 




a result, most herbivorous insects develop 
only one generation per season, leading to a 
time lag in the interaction between herbi- 
vorous insects and their food resources". 
Consequently, most of the primary produc- 
tivity in deserts ends up as dry plant material 
or seeds, which accumulates because micro- 
bial decomposition is limited by the low 
moisture availability. These resources form 
the basis for populations of detritivores such 
as termites, darkling beetles and isopods, 
and seed predators such as ants. These 
groups can support a rich fauna of predatory 
arthropods and reptiles as well as omni- 
vorous birds and mammals. It is often 
assumed that all deserts have low species 
diversity because of the harsh environmental 
conditions, but among animals almost all 



_ 


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■KTl 


mm 


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a. 











terrestrial higher taxa are represented, and 
their species diversity may in some situ- 
ations be comparable to that of more mesic 
habitats". 

True desert species show a wide range of 
adaptations to the conditions of an extreme 
environment. Characteristic plants Include 
the Cactaceae In the Americas and the 
succulent Euphorbiaceae in Africa. Semi- 
desert species include salt bush Atnplex, and 
creosote bush Larrea. Amongst animals, 
groups that are Intrinsically adapted to very 
low moisture environments Include reptiles 
and many arthropods, although species in a 
wide range of other groups have also evolved 
to cope with these conditions. Strategies for 
survival amongst both plants and animals 
often include long periods of dormancy (as 
seeds, in the case of many plants! between 
rainfall events. 

In the particular conditions prevailing In 
the so-called fog deserts (notably the 
Atacama desert and the Skeleton Coast 
desert], different strategies have evolved. 
Here plants and animals make use of the 
regular moisture-laden fogs, which roll in 
from the cold offshore currents, to obtain a 
low but predictable supply of water 

The often overlooked inland water habitats 
of deserts may contain a particularly high 
proportion of locally endemic species; the 
pools of Cuatro Clenegas, tvlexico, for example, 
contain numerous mollusk and fish species 
found nowhere else. 

Many of the larger desert and sub-desert 
vertebrates are threatened; the openness of 
these arid areas means that species such as 
antelopes and other bovids are more con- 
spicuous than forest species and thus more 
vulnerable to overhuntlng. Threatened verte- 
brates include the wild bactrian camel 
Camelus bactrianus with a few remnant 
populations in the Gobi desert of Mongolia 
and China, and Przewalskl's gazelle Procapra 
przewatskii of China's subdesert steppes, now 
restricted by overhuntlng and habitat loss to a 
few small areas surrounding Lake Quinghai. 
The Mexican prairie dog Cynomys mexicanus 
is confined to prairies and intermontane 
basins with herbs and grasses where It is 
threatened by persecution and continuing 



Terrestrial biodiversity in 



habitat loss. The Addax antelope Addax 
nasomaculatus was originally widespread 
from the western Sahara to Egypt and Sudan, 
but as a result of uncontrolled hunting it 
persists only in small and scattered local 
populations and is extinct through much of its 
former range. 

Principal threats to desert environments 
include the activities of domestic livestock 
such as cattle, which cause soil compaction 
through trampling, and can damage vege- 
tation and waterholes. Introduced species, 
such as rabbits in Australia, have been highly 
damaging to some desert environments. 
Other human activities that have affected 
desert habitats include use of off-road 
vehicles, irrigation and afforestation schemes, 
and housing projects. In drylands, most 



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land degradation can be categorized as 
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Combat Desertification, the latter term is 
defined explicitly as land degradation in arid, 
semi-arid and dry sub-humid areas resulting 
from various factors, including climatic 
variations and human activities'. According to 
the above definition, hyperarid lands [true 
deserts! are not susceptible to desertification, 
because their productivity is already so low 
that it cannot be seriously decreased by 
human action. The effects of desertification 
on arid and semi-arid areas promote poverty 
among rural people, and by placing greater 
pressure on natural resources, poverty tends 
to reinforce any existing trend toward 
desertification. 



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the intergovernmental forum on forests. Commonwealth of Australia, Canberra. 
Loh. J. led.) 2000. Living planet report 2000. WWF - World Wide Fund for Nature, Gland. 
Hansen, M. et al. 2000. Global land cover classification at 1 km resolution using a 
decision tree classifier. InternationalJournat of Remote Sensing 21: 1331-1365. 
Barthlott, W. et al. 1999. Terminological and methodological aspects of the mapping and 
analysis of global biodiversity. Acta Botanica Fennica 162: 103-1 10. 
Heyvifood, V.H. led. I 1993. Flowering plants of ttie world. B.T Batsford Ltd, London. 



Marine biodiversity 117 



•^H 



O Marin 



e biodiversity 



MOST OF THE PLANET IS COVERED BY OCEAN waters whose average depth is four 
times the average elevation of the land, making the open sea by far the largest 
ecosystem on Earth. Despite this volume, marine net primary production remains 
similar to or less than that on land because photosynthesis in the sea Is carried out by micro- 
scopic bacteria and algae restricted to the sunlit surface layers (plants are virtually absent). 
The diversity of major lineages (phyla and classes! is much greater in the sea than on land 
or in freshwaters, and many phyla of invertebrate animals occur only in marine waters. 
Species diversity appears to be far lower, perhaps because marine waters are physically 
much less variable in space and time than the terrestrial environment. 

Marine fisheries are the largest source of wild protein, derived from fishes, moltusl<s and 
crustaceans. The world catch from capture fisheries has grown fivefold over the past five 
decades, but appears to have declined during the 1990s despite increased fishing effort. 
More than half of the world's major fishery resources are now in need of remedial 
management, mainly because of excess exploitation. 



THE SEAS 

Oceans cover 71 percent of the world's 
surface. They are on average around 3.8 kilo- 
meters Ikml deep and have an overall volume 
of some 1 370 million kml The whole of the 
world ocean (all contiguous seasi is theor- 
etically capable of supporting life, so that the 
marine part of the biosphere is far larger 
than the terrestrial part. However, as on land, 
life in the oceans is very unevenly distributed 
- some parts are astonishingly productive 
and diverse while others are virtually barren. 

With the present configuration of land 
masses, a major part 137 percent] of the world 
ocean is within the tropics, and about 75 
percent lies between the 45° latitudes. The 
largest continental shelf areas are in high 
northern latitudes (Table 6.21, but about 30 
percent of the total shelf area is in the tropics. 
Within the tropics, the shelf is most extensive 
in the western Pacific (China Seas south to 
north Australia). 

Although knowledge of the functioning 
of the marine biosphere has increased enor- 
mously in the past few decades, overall it 



remains far less well known and understood 
than the terrestrial part of the globe. The 
main reason for this is, quite simply, that 



Table 6.1 

Area and maximum depth 

of the world's oceans and 

seas 

Source: Couper . 



1 Ocean/Se^^^^^^H 


■ Area (km'l 


Depth (meters) J 


Pacific Ocean 


165 384 000 


11524 


Atlantic Ocean 


82 217 000 


9 560 


Indian Ocean 


73 481000 


9 000 


Arctic Ocean 


14 056 000 


5 450 


Mediterranean Sea 


2 505 000 


4 846 


South China Sea 


2 318 000 


5 514 


Bering Sea 


2 269 000 


5 121 


Caribbean Sea 


1 943 000 


7 100 


Gulf of Mexico 


1 544 000 


4 377 


Sea of Okhotsk 


1 528 000 


3 475 


East China Sea 


1 248 000 


2 999 


Yellow Sea 


1 243 000 


91 


Hudson Bay 


1 233 000 


259 


Sea of Japan 


1 008 000 


3 743 


North Sea 


575 000 


661 


Black Sea 


461 000 


2 245 


Red Sea 


438 000 


2 246 


Baltic Sea _. , ,., 


,, 422 000 ,__ 


— „ -'i^O .. _u.. 



118 WORLD ATLAS OF BIODIVERSITY 



S- 



^^^B^HH 


^^_ Open ocean ^^M 


^HrContinentalsh|iy| 


Total area Imillion kml 

Latitude bands 1% of total! 
Polar and boreal 145-90°! 
Temperate [20-45°! 
Tropical [0-20°l 


360.3 

26,6 
36.8 
36.6 


26.7 

40.9 
28.8 
30.3 



Table 6.2 
Relative areas of 
continental shelves and 
open ocean 

Source: Adapted from Longhurst and 
Pauty , after t^oiseev. 



much of it is inaccessible to humans. Study of 
any part below the top few meters requires 
specialized equipment and is expensive and 
time consuming. Knowledge of most of the 
sea IS thus based largely on a range of 
remote-sensing and sampling techniques and 
often remains sketchy. As these techniques 
become more sophisticated, so our under- 
standing of marine ecosystems, particularly 
those away from the coastal zone, is under- 
going constant revision. 

Sea water and ocean currents 
Sea water is a complex but relatively uniform 
mixture of chemicals. Most of the 92 naturally 
occurring elements can be detected dissolved 
in it, but most only in trace concentrations; 
the most abundant are sodium las Na+! and 
chlorine las Cl-1, which occur at a concen- 
tration some ten times higher than the next 
most abundant element, magnesium. The 
term used to quantify the total amount of 
dissolved salts in sea water is salinity, a 
dimensionless ratio, which generally ranges 
between 33 and 37 and averages 35. Most of 
the substances dissolved in sea water are 
unreactive and remain at relatively stable 
concentrations; however those that play a 
part in biological systems can be highly 
variable in time and space. 

The world's sea waters are constantly in 
motion, at all scales from the molecular to 
the oceanic. Large-scale ocean circulation 
plays a vital role in mediating global climate 
as well as influencing the functioning of 
marine ecosystems. It is driven by complex 
interactions between a number of physical 
variables, notably latitudinal variations in 
solar radiation land consequent heating and 
cooling!, precipitation and evaporation, trans- 
fer of frictional energy across the ocean 



surface from winds, and forces resulting 
from the rotation of the planet. 

Surface currents are largely driven by pre- 
vailing winds. The most important features are 
vast, anticyclonic gyres in the subtropical 
regions of the world's oceans. These are pri- 
marily driven by the westerly trade winds in 
the Roaring Forties and circulate clockwise in 
the northern hemisphere and counter- 
clockwise in the southern hemisphere. Also 
important are the eastward-flowing Antarctic 
circum-polar current and equatorial current. 
At smaller scales, eddies and rings are ubiqui- 
tous and are analogous to weather systems in 
the atmosphere; typical oceanic eddies may be 
1 00 km across and persist tor a year or more. 

The density of sea water increases with 
increasing salinity and with decreasing temp- 
erature until it freezes at around -1.9°C. Sea 
water of different density does not mix readily, 
and so the oceans tend to be well stratified 
vertically, with bodies of less dense sea water 
[warmer and less saline! sitting on top of 
cooler, more saline and denser bodies. The 
stratification is rarely stable over the long 
term, however, as the influence of climate 
changes the properties of surface waters and 
causes various forms of vertical mixing. The 
single most important factor controlling large- 
scale deep-water circulation appears to be the 
generation of cold, high-salinity water near the 
surface in the Weddell Sea and off Greenland in 
the North Atlantic. Here, during winter, the sea 
freezes and the floating ice sheet is virtually 
free of salt, leaving the underlying water more 
saline, cold and dense. This sinks to the bottom 
and moves south along the Atlantic floor, some 
passing south of the equator and circulating 
throughout the world ocean, producing what is 
known as the Great Conveyor - the interlocking 
system of major circulation currents in the 
deep sea. Because it originates near the 
surface it is well oxygenated as well as cold, 
and is the major reason why aerobic organisms 
can thrive in the deep sea. 

Major zones of upwelling occur along the 
western boundaries of continents where trade 
winds blow towards the equator, causing 
surface waters to be pushed offshore to be 
replaced by cooler deep waters. There are five 
major upwelling areas: the Humboldt current 



Marine biodiversity 119 



region off the Cfiilean and soutfiern Peruvian 
coast of South America; the California current 
region off western North Annerica; the Canary 
current off the coast of Mauritania in north- 
w/est Africa; the Benguela current region off 
southern Africa; and the northwest Arabian 
Sea. Marine production is much enhanced in 
these regions which typically support major 
pelagic fisheries. Outside these upwelling 
zones, stratification tends to persist through- 
out the year in the tropics and subtropics. 

MAJOR MARINE ZONES 
The continental shelf 

Marine waters around major land masses 
are typically shallow, lying over a continental 
shelf which may be anything from a few 
kilometers to several hundred kilometers 
wide. The most landward part Is the littoral or 
intertidal zone where the bottom is subject to 
periodic exposure to the air Water depth here 
varies from zero to several meters. Seaward 
of this the shelf slopes gently from shore to 
depths of one to several hundred meters, 
forming the sublittoral or shelf zone. Waters 
below low-tide mark in the continental shelf 
region are referred to as neritic. 

The extent, gradient and superficial geo- 
logy of continental shelf areas are determined 
by many factors. Including levels of tectonic 
activity in the Earth's crust. More than 80 
percent of the global volume of river-borne 
sediment Is deposited In the tropics (and an 
estimated 40 percent of it by just two river 
systems: the Huang He or Yellow River and 
the Ganges-Brahmaputra)', and this Is ref- 
lected in the extent of shelf areas In parts 
of the tropics, and in the high turbidity of 
coastal waters In monsoon regions. Most 
shelf areas in the tropics are overlain by 
sands or muds composed of sediment of 
terrestrial origin (terrigenous deposits). 

Shelf regions support the marine com- 
munities most familiar to humans, and many 
of the marine resources of particular value to 
them. Although mangrove and coral reefs are 
two of the best known tropical coastal 
ecosystems, they dominate only a minor part 
of the world coastline: the former mainly In 
deltaic or other low-lying coastal plains, and 
the latter only in shallow waters where 



terrestrial sedimentation is very low. Soft- 
bottom habitats with sparse vegetation are 
probably the most widespread coastal marine 
ecosystem type, and virtually the entire 
seabed away from the coastline Is covered in 
marine sediments (see Box 6.1). 

The deep sea 

At the outer edge of the shelf there is an 
abrupt steepening of the sea bottom, forming 
the continental slope which descends to 
depths of 3-5 km. The sea bottom along the 
slope IS referred to as the bathyal zone. At 
the base of the continental slope are huge 
abyssal plains which form the floor of much of 
the world ocean. Open waters above these 
plains make up the oceanic pelagic zone. 
Given that the oceans cover some 71 percent 
of the globe, and that the shelf area is 
relatively narrow, the oceanic pelagic zone is 
by far the most extensive ecosystem on Earth. 
The plains are punctuated by numerous sub- 
marine ridges and sea mounts which may 
break the surface to form Islands. There are 
also a number of narrow trenches which have 
depths of from 7 000 to 11 000 meters (m). 
These constitute the hadal zone. Ocean 



^Box 6.1 Life in sediments 

Spaces between tfie particles of marine sediments contain water, air, detritus and 
organisms, tfie last divided by size Into two main categories. The microfauna includes 
archeans, bacteria and protoctists. These include a number of pnmary producers at the base 
of the food web in shallow waters, and have a role in interstitial sulfur chemistry and 
oxygenation of sediment. The melofauna includes sediment-living forms between 0,1 and 
1 mm in size, and Is a major component of seabed ecosystems, particularly In the deep sea. 
Four phyla contain only marine melofauna: the Gastrotricha, Gnathostomullda, KInorhyncha 
and Loriclfera. Nematodes are typically the most numerous component, with harpacticoid 
copepods and foraminiferans also important. High concentrations occur around the burrows 
of deposit-feeding mollusks and polychaetes, with high bacterial numbers and elevated 
nutrient flux. They play a key role In the flow of nutrients from the microfauna to larger 
organisms, of which several distinctive species exist In and on the seabed. Sea cucumbers, 
crinoids, polychaete worms, sea spiders, Isopods and amphipods are most abundant. Some 
taxa are only found here. Pogonophorans occur mostly below 3 000 m and many of the 
known species are restricted to the sediments of deep ocean trenches (more than 6 000 m). 
Small-scale variation In food supply, such as the fortuitous appearance of a large animal 
carcass, may enhance spatial structure and provide opportunity for other species to colonize. 
A micro-landscape of hills and valleys is created by the burrowing and fecal mounds of 
echiurid and polychaete worms. 



jea 



120 WORLD ATLAS OF BIODIVERSITY 



Coral reefs develop within 
the photic zone because of 
the symbiotic relationship 
between some coral 
species and zooxanthellae. 



trenches are formed as a consequence of 
plate tectonic processes where sectors of ex- 
panding ocean floor are compressed against 
an unyielding continental mass or island arc, 
resulting in the crust buckling downwards 
IsubductingI and being destroyed within the 
hot interior of the Earth. 

CLASSIFYING THE MARINE BIOSPHERE 

Much less effort has been devoted to catego- 
rizing and mapping units within the marine 
biosphere than to terrestrial environments. 
As on land, basic marine habitat types are 
commonly defined by some combination of 
structural, climatic and community features 
le.g. 'tropical coral reef'l but, probably in part 
reflecting issues of scale and resolution, few 
global maps of marine ecosystem types are 
available. 



marine ecosystem ILMEI units have been 
defined' ', with a view to improving planning 
and management. These are large regions of 
marine space, some 200 000 km' or larger, 
that extend from river basins and estuaries 
out to the seaward boundary of the conti- 
nental shelf, with distinct combinations of 
bathymetry, hydrography and production. 
With similar aims, the conservation organi- 
zation WWF has distinguished 43 large 
marine ecoregions, based broadly on oceano- 
graphic and community features, and also 
intended to be globally representative'. 

THE BASIS OF LIFE IN THE SEAS 

In the sea, as on the land, photosynthesis is 
the driving force behind maintenance of life. 
Because photosynthesis in nature depends on 
sunlight, with few exceptions primary pro- 




The principal global scheme intended to 
classify the entire world ocean within an 
objective oceanographic system is based on 
long-term data on sea surface color 
(obtained by the CZCS radiometer carried by 
the Nimbus orbiting satellite during 1978- 
861' \ These data reflect chlorophyll concen- 
trations and provide a basis for estimating 
primary production rates, and changes over 
time, on a 1-degree grid. These values, 
together with numerous other data sets, have 
been used as the basis of a classification of 
the world ocean into four ecological domains 
and 56 biogeochemical provinces. Taking a 
less quantitative approach, some 64 large 



ductivity is confined to those parts of the 
ocean that are sunlit. Water absorbs sunlight 
strongly so that light intensity decreases 
rapidly with increasing depth. Red wave- 
lengths are most rapidly absorbed, except 
where turbidity is high, while blue-green 
wavelengths penetrate the deepest. Even in 
the clearest waters the latter are completely 
absorbed by around 1 km depth, this marking 
the extreme limit of the so-called photic zone. 
Photosynthesis is thus limited to the contin- 
ental shelf area and the first few hundred 
meters of surface waters (and often much 
less] of the open ocean which together make 
up a very small proportion of the total volume 



Marine biodiversity 121 



of the oceans. The portion of the photic zone 
where sunlight is strong enough to support 
appreciable amounts of photosynthesis is the 
euphotic zone- 
Virtually all other marine organisms. 
including those of the unlit middle depths and 
the deep sea. are dependent ultimately on 
growth of primary producers in areas that 
may be widely distant from them in time and 
space. The most important exceptions to this 
are the archeans and bacteria living around 
hydrothermal vents associated with rift zones 
in the ocean floor. The water here can be 10°C 
warmer than in adjacent areas and these 
microorganisms are able to grow using hydro- 
gen sulfide gas emitted at the vents as an 
energy source, and they in turn are used by 
other organisms. 

With some exceptions primary production 
per unit area tends to be lower in marine 
environments than in terrestrial ones, espec- 
ially if highly managed terrestrial agricultural 
systems are considered. This is because over 
the vast majority of the ocean the euphotic 
zone is far distant from the lithosphere. The 
latter provides essential nutrients for life, and 
these therefore have to be transported to the 
euphotic zone to allow life processes to 
continue. There is also usually a steady loss of 
nutrients and organic compounds from the 
euphotic zone, owing to the sinking of particles 
and bodies into the dark regions of the sea 
where no photosynthesis can take place. 

Continued productivity in the open sea is 
contingent on the replacement of the lost 
nutrients. The latter may originate either on 
land, in the form of river outflow, or from mar- 
ine sediments. Their replacement in the pelagic 
euphotic zone is dependent on the mixing or 
vertical movement of the water column. At 
latitudes higher than A0°, winter mixing allows 
replenishment of the euphotic zone, partic- 
ularly in continental shelf areas. However, in 
permanently stratified subtropical and tropical 
oceanic waters there is little vertical mixing, 
and therefore little influx of new nutrients. 
Productivity in these areas is correspondingly 
low. In zones of upwelling. surface waters are 
regularly replaced by nutrient-rich bottom 
waters and very high levels of productivity can 
be achieved, at least seasonally 



Until the 1980s it had been believed that 
photosynthesis in the pelagic ocean was car- 
ried out only by single-celled phytoplankton. 
between 1 and 100 microns in diameter 
II micron = 0.001 mm|, and also that vast 
expanses of open ocean where phytoplankton 
could not be detected were, in terms of 
productivity, the marine equivalent of deserts. 
New observational techniques have since 
revealed the presence in great abundance of 
exceptionally small and previously unknown 
organisms, collectively termed picoplankton. 
These appear to be predominantly photo- 
synthesizing unicellular cyanobacteria 0.6-1 
micron in size, such as Prochlorococcus. 
Because of their extraordinary abundance 
(some 100 million cells may be present in 
1 liter), and despite their minute size, these 
organisms play a crucial role in the produc- 
tivity of open ocean waters' and have led to 
marked upward revisions in estimates of 
overall marine productivity 

The major role played by microscopic 
organisms in marine productivity has a 
number of important implications for marine 
ecology. Although much remains unknown 
about the structure and dynamics of pelagic 
food webs, it seems that a high proportion 
of marine primary production is used directly 
by microscopic organisms Iboth autotrophic 
and heterotrophic! and cycled back into non- 
living forms (dissolved carbon dioxide and 
organic carbon! rather than supporting 
populations of larger organisms. 

Oceanic primary producers divert a high 
proportion of their energy into reproduction 
rather than accumulating biomass. in con- 
trast to terrestrial primary producers (plants!. 
As a result of this, average standing biomass 
per unit area in the oceans has been esti- 
mated at around one-thousandth that on land. 
Population turnover of oceanic primary pro- 
ducers is also several orders of magnitude 
higher than turnover of major terrestrial pri- 
mary producers [plants!. 

The small size and rapid turnover of 
oceanic primary producers m.ean that there 
are no organisms directly analogous to the 
woody plants that so enrich terrestrial 
environments by providing structurally com- 
plex habitats for other organisms. The nearest 



122 WORLD ATLAS OF BIODIVERSITY 



Table 6.3 

Marine diversity by 

phylum 



Notesr Strictly marine groups 
shown in bold. Estimates are 
from a variety of sources and 
in some cases, e.g. Mollusca, 
differ in detail from ttiose in 
Table 2.1. 



equivalents are the large brown algae known 
as kelp Iphylum Phaeophyta], whose structure 
is less complex and which are much more 
narrowly distributed. Structurally complex 
habitats may in contrast be created by 
animals, particularly corals Iphylum Cnidarial 
and, to a lesser extent, sponges (phylum 
Ponferal, mollusks and serpulid worms 
Iphylum Annelida]. 

BIOLOGICAL DIVERSITY IN THE SEAS 

It is well known that diversity at higher 
taxonomic levels (phyla and classes! is much 
greater in the sea than on land or in 
freshwater Of the 82 or so eukaryote phyla 
currently recognized (see Chapter 2), around 



60 have marine representatives compared 
with around 40 found in freshwater and 40 on 
land. Amongst animals the preponderance is 
even higher, with 36 out of 37 phyla having 
marine representatives (Table 6.31. 

Some 23 eukaryote phyla, of which 18 
are animal phyla, are confined to marine 
environments. Most of these are relatively 
obscure and comprise few species. The major 
exceptions are the Echinodermata, of which 
some 7 000 species are known, and the 
Foraminifera, with around 4 000 known, ex- 
tant species. A number of other important 
phyla including the coelenterates (Cnidarial, 
sponges (Poriferal and brown and red algae 
(Phaeophyta and Rhodophyta, respectively! 



r 



amain Kingdom 



Arcfiaea 
Bacteria 



2 ptiyla 
12 pfiyla 



■H 



inidieii nn. 

of described 
> marine species 



Eukaryota Protoctista 27 phyla, including: 

Chloroptiyta Green algae 

Pfiaeoptiyta Brown algae 

Rfiodophyta Red algae 

others incl. 
Foraminifera 

Animalia 



7 000 

1500 

4 000 

23 000 



Placozoa 




1 


Porifera 


Sponges 


10 000 


Cnidaria 


Coelenterates 


10 000 


Ctenophora 


Comb jellies 


90 


Platyhelminthes 


Flatworms 


15 000 


Nemertina 


Nemertines 


750 


Gnathostomulida 


Gnathostomulids 


80 


Rhombozoa 


Rhombozoans 


65 


Orthonectida 


Orthonectids 


20 


Gastrotncha 


Gastrotnchs 


400 


Rotifera 


Rotifers 


50 


Kinorhyncha 


Kinorhynchs 


100 


Lorlcifera 


Lorlclferans 


10 


Acanthocephala 


Spiny-headed worms 600 


Entoprocta 


Entoprocts 


170 


Nematoda 


Nematodes 


12 000 


Nematomorpha 


Horsehair worms 


<240 



pnyia 


tsiimaieo no. 




of described 




marine specie 


Ectoprocta 


Ectoprocts 


5 000 


Phoronida 


Phoronid worms 


U 


Brachiopoda 


Lamp shells 


350 


Mollusca 


Mollusks 


?75 000 


Priapulida 


Priapullds 


8 


Sipuncula 


SIpunculans 


150 


Echlura 


Echiurids 


UO 


Annelida 


Annelid worms 


12 000 


Tardigrada 


Water bears 


few 


Chelicerata 


Chelicerates 


1000 


Mandibulata 


Mandibulale 






arthropods 


few 


Crustacea 


Crustaceans 


38 000 


Pogonophora 


Beard worms 


120 


Bryozoa 


Bryozoans 


4 000 


Echinodermata 


Echinoderms 


7 000 


Chaetognatha 


Arrow worms 


70 


Hemichordata 


Hemichordates 


100 


Urochordata 


Tunicatesand 






ascidians 


2 000 


Cephalochordata 


Lancelots 


23 


Craniata 


Craniates 


15 000 



Fungi 3 phyla 

Plantae Anthophyta 



500 



50 



Total 



ca 250 000 



Marine biodiversity 123 



•^i< 



''til, 



are very largely marine, each with only a 
small number of non-marine (usually fresh- 
water] species. 

The reason for this predominance of 
marine higher taxa (particularly amongst 
animals) is believed to be because most of 
the fundamental patterns of organization 
and body plan. i.e. the different basic kinds 
of organism that are distinguished as phyla, 
originated in the sea and remain there, but 
only a subset of them has spread to the land 
and into freshwaters. It is noteworthy that 
only a third or so of marine phyla are found 
in the pelagic realm, the remainder being 
confined to sea bottom (benthicl areas - the 
habitat where eukaryotic organisms are 
believed to have evolved. 

In contrast, known species diversity in 
the sea is much lower than on land - some 
250 000 species of marine organisms are 
currently known, compared with more than 
1.5 million terrestrial ones. Much of this 
difference is because of the large numbei of 
described terrestrial arthropods, for which 
there is no marine equivalent. Amongst 
fishes, almost as many freshwater species as 
marine are known, despite the fact that 
freshwater habitats account for only around 
one ten-thousandth of the volume of marine 
ones. Similarly, the most diverse known 
marine habitats (coral reefsl are far less 
diverse in terms of species number than the 
moist tropical forests that are often taken as 
their terrestrial counterparts. 

The apparent lower total species diversity 
of the marine biosphere is likely in part at least 
to be a result of the physical characteristics of 
water, particularly its high heat capacity and 
its ability to mix. Because of these, marine 
environments (particularly deep-water ones) 
tend to show much less variation in time and 
space in their physical characteristics than 
terrestrial ones. This lack of physical variation 
seems to result in a similar lack of ecological 
variation over wide areas. 

In contrast to terrestrial faunas, where 
one phylum - the Mandibulata (insects and 
relatives) - vastly outnumbers all others in 
terms of known species, marine species are 
much more evenly distributed across higher 
taxa. The largest marine phyla - Mollusca and 



1 Class J^l 




r No, of marine species 


Myxini 


Hagfishes 




i3 


Cephalaspidomorphi 


Lampreys 




9 


Elasmobranctiii 


Sfiarks, skates and rays 




900 


Holocephali 


Chimaeras 




31 


Actinopterygii 


Ray-fin fishes 




13 8C0 


Sarcopterygii 


Lobe-fin fisties 




2 


Reptilia 


Reptiles 




60 


Aves 


Birds 




307 


Mammalia 


Mammals 




110 



Crustacea - each comprise far fewer than 
100 000 known marine species, in contrast 
with the Mandibulata, of which around 
1 million terrestrial species have been 
identified to date. The only major eukaryote 
phyla (i.e. those with 10 000 or more de- 
scribed species! that are believed to have 
comparable levels of diversity both on land 
and in the sea are the Platyhelminthes, 
Nematoda, Mollusca and Craniata. As on 
land, vertebrates are by far the best-known 
group of marine organisms. Of the 50 000 or 
so described extant species, more than 
15 000 may be considered marine (Table 6.41, 
the overwhelming majority of which are fishes 
(Table 6.51 and a few tetrapods (Table 6.6). 

Marked latitudinal gradients in species 
richness have been described in a number of 
groups, for example in benthic isopods and 
mollusks", and it is generally true that coastal 
waters within the tropics tend, in parallel with 
terrestrial environments, to be richer in 
number of species than those at higher 
latitudes (although many exceptions to this are 
known, such as penguins, pinnipeds and auksl. 
Although marine biogeography had advanced 
relatively little since the early 20th century, 
improved distribution data and more robust 
methods have recently been developed. These 
have been applied to definition of coastal 
biogeographic regions', and important areas 
for marine biodiversity analogous to those 
delimited on land. For example, 18 centers of 
endemism within the world tropical reef zone 
have been identified using data on reef fish, 
coral, snails and lobsters (Map 6.1); ten 
centers were regarded as hotspots' because of 
their higher threat score'". 



Table 6A 

Diversity of craniates in 

the sea by class 

Note: Because of changing 
taxonomy, incomplete 
information and occupation of 
multiple habitat types, these 
estimates are indicative only. 



124 WORLD ATLAS OF BIODIVERSITY 



mm 


Common fishes No. of 


No. of 


No. of 


JH^ 


—Common fislies No. of 


No. of 


No. off 




families 


genera 


species 


^H 


B families 


genera 


species 


1 








occurring 










occurring 










in marine 










in marine 


■I^H 


^^^^^HH 


B 


k 


tiabitats 




J^B 


B 


1 


habita^ 


Myxiniformes 


Hagfishes 


1 


6 


43 


Atheriniformes 


Silversides 


8 


47 


139 


Petromyzontiformes Lampreys 


1 


6 


9 


Beloniformes 


Needlefishes, sauries, 








Chimaeriformes 


Chimaeras 


3 


6 


31 




flyingfishes, halfbeaks 


5 


38 


140* 


Heterodontiformes 


Bullhead sharks and 
horn sharks 


1 


1 


8 


Cyprinodontiformes 


Rivulines, killifishes, 
pupfishes, four-eyed fishes. 








Orectolobiformes 


Carpet sharks 


7 


U 


31 




poeciliids, goodeids 


8 


88 


13 


Carcharhiniformes 


Ground sharks 


7 


47 


207 


Stephanoberyclformes Gibberfishes, pricklefishes. 








Lamniformes 


Mackerel sharks 


7 


10 


16 




whalefishes, hairyfish. 








Hexanchiformes 


Cow sharks 


2 


i 


5 




tapetails 


9 


28 


86 


Squaliformes 


Dogfishes and 








Beryciformes 


Fangtooths, spinyfins. 










sleeper sharks 


i 


23 


74 




lanterneyefishes, roughies. 








Squatiniformes 


Angel sharks 


1 


1 


12 




pinecone fishes, squirrelfishes 7 


28 


123 


Pristiophoriformes 


Saw sharks 


1 


2 


5 


Zeiformes 


Dories, boarfishes. 








Rajiformes 


Rays 


12 


62 


432* 




oreos, parazen 


6 


20 


39 


Coelacanthiformes 


Coelacanths 


1 


1 


2 


Gasterosteiformes 


Pipefishes, seahorses. 








Acipenseriformes 


Sturgeons 


2 


6 


12 




sticklebacks, sandeels. 








Salmoniformes 


Salmonlds 


1 


11 


21 




seamoths, snipefishes. 








Stomiiformes 


LIghtfishes, hatchetfishes, 










shnmpfishes. trumpetfishes 


11 


71 


238' 




barbeled dragonfishes 


4 


51 


321 


Synbranchiformes 


Swamp-eels 


3 


12 


3 


Ateleopodiformes 


Jellynose fishes 


1 


i 


12 


Scorpaeniformes 


Gurnards, scorpionfishes. 








Aulopiformes 


Greeneyes, pearleyes, 
waryfishes, 
lizard fishes, 










velvetfishes, flatheads. 
sablefishes, greenlings, 
sculpins, oilfishes, poachers. 










barracudinas, lancetfishes 


13 


42 


219 




snailfishes, lumpflshes 


25 


266 


1219* 


Myctophiformes 


Lanternfishes 


2 


35 


241 


Perciformes 


Perches, basses, sunfishes, 








Lampridiformes 


Garfishes, ribbonfishes. 










whitings, remoras, jacks. 










crestfishes, opahs 


7 


12 


19 




dolphinfishes, snappers. 








Polymixiiformes 


Beardfishes 


1 


1 


5 




grunts, damselfishes. 








Ophidiiformes 


Pearlfishes, cusk-eels. 










dragonfishes, wrasses, 










brotulas 


5 


92 


350* 




butterflyfishes, etc. 


148 


1496 


7 371* 


Gadiformes 


Cods, hakes, rattails 


12 


85 


481 


Pleuronectifornnes 


Plaice, flounders, soles 


11 


123 


564* 


Batrachoidiformes 


Toadfishes 


1 


19 


64* 


Tetraodontiformes 


Triggerfishes, 








Lophiiformes 


Anglerfishes. goosefishes, 










puffers, boxfishes. 










frogfishes. batfishes, 










filefishes, molas 


9 


100 


327* 




seadevils 


16 


65 


297 












Mugilitormes 


Mullets 


1 


17 


65* 


Total Iconservative working estimatel 




c: 


uooo 



Table 6.5 

Diversity of fisties in the 

seas by order 

Note: Strictly marine orders in 
bold; ottier orders that are 
mainly marine (more than 
50% of species! marked with 
an asterisk'. 

Source: Afler Nelson*'. 



Algae 

The macro-algae are superficially plant-like 
protoctists that lack the vascular tissue used 
by higher plants to transport w/ater and 
nutrients. They are almost exclusively aquatic; 
three of the four principal groups consisting 
of large-sized species are mainly marine in 
occurrence. These three, the green, brown 
and red algae (seaweeds'!, are all cosmo- 
politan in distribution and occur in a range of 



environments, although some constituent 
families have somewhat restricted ranges. 
There are more marine species of red algae 
IRhodophyta! - around 4 000 - than the 
greens (Chlorophyta, ca 1 0001 and browns 
(Phaeophyta, ca 1 500! combined. 

As with pinnipeds and seabirds, the cold 
and cool temperate regions of the world 
appear to be surprisingly rich in species. On 
present incomplete information, the region 



Marine biodiversity 125 



around Japan (norttiwest Paciticl. the North 
Atlantic, and the tropical and subtropical 
western Atlantic hold the most species of 
marine algae. Southern Australia is not so 
species rich but appears to have the highest 
proportion of endemics. There are few 
species of larger algae in regions of cold- 
water upwelling; small isolated islands and 
polar regions also have few species. In 
contrast, coral reefs support a unique and 
generally diverse algal flora that includes 
many crustose coralline algae (more species 
of which are likely to be discovered). 
Mangrove areas also support a well-defined 
algal vegetation. Sandy coastlines hold few 
species of large algae and often form barriers 
to seaweed dispersal. 

Marine fishes 

Fishes are considered a paraphyletic group 
(see Chapter 21: that is, all living species are 
thought to share a common ancestor but to 
share this ancestor with another group (in this 
case, the tetrapodsl not categorized as fishes. 
Apart from some 50 or so species of generally 
parasitic lampreys and hagfishes in the 
superclass Agnatha, fishes are divided into 
two unequal-sized groups, the cartilaginous 
fishes (Chondrichthyesj including chimaeras 
(class Holocephalil and sharl<s and rays 
(class Elasmobranchiil, and the bony fishes 
(OsteichthyesI, including the 'typical' ray- 
finned fishes (class Actinopterygiil and the 
lobe-finned coelacanths and lungfishes (class 
Sarcopterygiil. Some 60 percent of all l<nown 
living fish species (i.e. around 14 000 species! 
occur in marine habitats. They range in size 
from an 8 mm-long goby Trimmatom nanus in 
the Indian Ocean to the 15-m whale shark 
Rhincodon typus, respectively the smallest 
and largest of all fish species, and occur in 
virtually all habitats, from shallow inshore 
waters to the abyssal depths. 

The elasmobranchs (sharks, skates and 
raysl are an overwhelmingly marine group 
with around 850 living species in ten orders. 
Although far less diverse than the bony fishes, 
the cartilaginous fishes include many of the 
largest fish species, a number of which are top 
predators in marine ecosystems. There tend to 
be more shark species at lower latitudes, but 



at family level richness tends to be higher on 
the edge of the tropics (Map 6.2). The bony 
fishes are a remarkably diverse group, with an 
enormous range of morphological, physio- 
logical and behavioral adaptations. Of the 26 
orders of bony fishes with marine represen- 
tatives (Table 6.5], by far the largest and most 
diversified is the Perciformes. This is the 
largest of all vertebrate orders and dominates 
vertebrate life in the ocean, as well as being 
the dominant fish group in many tropical and 
subtropical freshwaters. 

As with other groups of organisms, the 
majority of fishes in the sea are strictly 
marine, occurring only in salt water. A 



Table 6.6 

Marine tetrapod diversity 

Notesr Birds follow the list of 
seabirds recognized in Croxall 
et al." with the additional 
inclusion of four eider ducks 
and three steamer ducks in 
the family Anatidae and the 
red phalarope Phalaropus 
fulicana (Scolopacidael. 
Taxonomy follows Sibley and 
Monroe". Figures in 
parentheses indicate species 
that breed largely or entirely 
inland. Strictly marine 
families shown in bold. 



Class 


Order 


Family S 


^V 


No. of marine 
species^^ 


Reptilia 


Chelonia 


Dermochelyidae 


Leathery turtle 


1 






Cheloniidae 


Sea turtles 


6 




Squamata 


Elapidae 


Sea snakes and sea kraits 55 






Acrochordidae 


File snakes 


1 






Iguanidae 


Iguanas 


1 


Aves 


Ansenformes 
Ciconiiformes 


Anatidae 

Scolopacidae 

Laridae 


Gulls, terns, skuas, 
auks, skimmers 


7 
1 

120(131 






Phaethontidae 


Tropicbirds 


3 






Sulldae 


Gannetsand boobies 


9 






Phalacrocoracidae 


Cormorants and shags 


36(21 






Pelecanidae 


Pelicans 


2 






Fregatidae 


Frigatebirds 


5 






Spfieniscidae 


Penguins 


17 






Procellariidae 


Petrels, albatrosses, 
shearwaters 


115 


Mammalia 


Cetacea 


Balaenidae 


Right whales 


3 






Balaenopterldae 


Rorquals 


6 






Esclirlclitiidae 


Gray whale 


1 






Neobalaenidae 


Pygmy right whale 


1 






Delpliinidae 


Dolphins 


32 






Monodontidae 


Beluga and nanvhal 


2 






Pliocoenidae 


Porpoises 


6 






Phystereidae 


Sperm whales 


2 






Platanistidae 


River dolphins 


1 






ZIphiidae 


Beaked whales 


19 




Sirenla 


Trichechidae 
Dugongldae 


Manatees 
Dugong 


1 
1 




Carnivora 


Mustelidae 
Odobenidae 


Otters and weasels 
Walrus 


2 

1 






Otarildae 


Eared seals 


U 


i^JbSiA 


B 


Phocidae 


Earless seals _^^ 


17 



126 WORLD ATLAS OF BIODIVERSITY 



•<*- 



Map 6.1 

Coral reef hotspots 

The location of 18 areas 
defined by fiigfi endemisrm 
in reef fisfies, corals, snails 
and lobsters, including the 
ten areas identified as 
'hotspots' on the basis of 
high threat score. 

Source: Adapted from Roberts . 




Endemic-rich areas 

Endemic-rich area at higher risk 
Endemic-rich area 



proportion, however, may also occur in 
inland waters, often passing a particular part 
of their life cycle there. Species that spend 
most of their life in marine waters but ascend 
rivers to breed, such as many salmonids 
Ifamlly Salmonidae, order SalmoniformesI 
and sturgeons (family Acipenseridae, order 
Acipenseriformesl, are referred to as anad- 
romous. Those that breed at sea but spend 
their lives otherwise In freshwater, such as 
most eels In the family Anguillldae (order 
Angulliformesl, are referred to as catadro- 
mous. Species with a wide salinity tolerance 
that may occur In marine, brackish and 
fresh waters (e.g. some sawfishes, family 
Pristidae, order Rajlformesl are referred to 
as euryhaline while those with narrow 
tolerances, be they to marine, brackish or 
fresh water, are referred to as stenohallne. 




t 




Reptiles 

Present-day diversity of reptiles in the seas 
Is low. One Important reason for this appears 
to be that modern reptilian kidneys cannot 
tolerate high salinities and thus the only 
reptiles that have adapted to marine environ- 
ments are those which have developed 
specialized salt-excreting glands. The most 
thoroughly marine reptiles are undoubtedly the 
sea snakes In the subfamily Hydrophlinae 
(family Elapidael. These spend their entire 
lives In the sea, giving birth to live young there. 
Although largely air-breathing like other 
reptiles, they can also absorb some oxygen 
directly from sea water and are thus able to 
remain submerged for long periods. Around 
50 species are known, widely distributed 
In tropical parts of the Indo-Paclfic region. In 
addition the little file snake Acrochordus 



Marine biodiversity 127 




granulatus (family Acrochordidael, from 
northern Australia and Southeast Asia is also 
entirely aquatic, but occurs in brackish estu- 
aries as welt as sea water. 

Five species of sea krait in the subfamily 
Laticaudinae are also largely marine, feeding 
mainly on eels. However they return to land 
to breed, generally on small tropical islands. 
They, too, are confined to the Indo-Pacific 
region. One species of lizard, the Galapagos 
marine iguana Amblyrhynchus cristatus 
(family Iguanidael, feeds underwater on mar- 
ine algae but spends a considerable proportion 
of time on land. Several other reptile species 
regularly enter sea water, most notably a 
number of homalopsine mangrove snakes 
(family Colubridae) from the Indo-Pacific and 
the estuarlne crocodile Crocodyius porosus 
(family Crocodylidael from the same region. 



k 



t 



s 



Undoubtedly the most prominent group of 
marine reptiles is the sea turtles, comprising 
the leathery turtle Dermocheiys coriacea in 
the family Dermochelyidae and six members 
of the family Cheloniidae. Alt species are 
large (ranging from 70-centimeter (cm| adult 
carapace length in Lepidochelys kempii 
to, exceptionally, 250 cm in Dermocheiys 
coriacea] and most are widely distributed in 
tropical and subtropical waters (Map 6.31. Sea 
turtles are almost completely marine; only the 
females emerge to nest on land, mostly within 
the tropics. One species, the loggerhead 
Caretta caretta, nests largely In temperate 
areas of the northern hemisphere. Sea turtles 
typically have a long period to maturity (often 
up to 25 years In the case of the green turtle 
Chetonia mydas] and a long lifespan. Females 
often nest only every two or three years. They 



128 WORLD ATLAS OF BIODIVERSITY 



Map 6.2 

Shark family diversity 

Diversity in sinarks, based 
on the distribution of all 30 
families plotted as a density 
surface. Most sharks are 
coastal in occurrence and 
for illustration purposes the 
family density is shown 
within a band extending out 
400 km from the coastline 
(slightly further than the 
200 nautical-mile EEZ limit). 



Source^ Prepared by UNEP-WCMC: 
family distributions aggregated from the 
species range maps published by 
Compagno 



/ 



Level of diversity 



High 




Low 



habitually return to the same nesting beaches, 
sometimes undergoing protracted migrations 
from feeding grounds. They may lay two or 
three clutches in a season, sometimes consis- 
ting of more than 100 eggs each, depending on 
the species. Nest, hatchling and juvenile 
mortality are often high. 

Birds 

Defining marine birds, or seabirds, is 
somewhat more problematic than defining 
marine species in other groups. All birds 
breed in terrestrial habitats, but a large 
number (almost all of them non-passerinesi 
obtain all or much of their food from aguatic 
or littoral habitats. Some of these, including 
all frigatebirds [Fregatidael, tropicbirds 
(Phaethontidael, gannets and boobies (Sulidael, 
penguins (Spheniscidael and petrels, alba- 




trosses and shearwaters (Procellanidael are 
indisputably marine, in that they obtain all 
their food from marine habitats, almost 
invariably breeding along coastlines and 
spending most or all of their time when not 
breeding out at sea. Many others, however, 
have less clear-cut habits. Some, such 
as a number of cormorants and shags 
(Phalacrocoracidael have both resident in- 
land and marine populations. Others, such as 
a number of gulls and terns (Laridae) and 
some ducks and geese (Anatidael. may breed 
inland but spend the rest of the year living in 
coastal areas or out at sea. Yet others, such 
as sandpipers (Scolopacidael and other 
waders, typically feed in littoral or intertidal 
habitats rather than in the sea itself; many of 
these species also occur inland. 

Adopting a somewhat arbitrary division, 



Marine biodiversity 129 



^% 




and excluding all wading birds with the 
exception of the red phatarope Phataropus 
fulicaria la truly pelagic species outside the 
breeding season), over 300 species of birds 
can be considered wholly or largely marine. 

In common with pinnipeds, seabirds show 
a latitudinal distribution in which diversity is 
much higher at higher latitudes [temperate 
and polar regions! than it is in the tropics. Two 
thirds of all seabirds are confined as breeding 
species to these latitudes, compared with only 
7 percent that are exclusively tropical. This 
stands in sharp contrast to the pattern found 
in most major terrestrial groups Isee Chapter 
51 and many marine groups such as sea 
turtles, mangroves and reef-building corals 
Isee, respectively. Maps 6.3, bA, 6.61 in which 
species diversity increases dramatically with 
decreasing latitude. Diversity is also markedly 





higher in the southern than in the northern 
hemisphere, with over half of all seabird 
species breeding in southern temperate and 
polar latitudes. Dominance of this region is 
even more marked in the Procellariidae, the 
family with the greatest number of truly 
marine species, in which over 60 percent of 
species breed at these latitudes and half are 
confined to it (Table 6.7). 



Table 6.7 

Regional distribution of 

breeding In seabirds 

Notes: Several species breed 
in more than one latitudinal 
band so that overall totals 
exceed actual number of 
seabirds. Numbers in 
parentheses indicate 
approximate number 
confined to the region. 



Breeding ^^^^1 


■ 


Ocean area 


Procellariidae 


Laridae 


Others 


Tote^ 


I distribution 




million km'l 




1 




^Hl 


Northern temperate and pola 


79.5 


24117) 


50 135) 


50 1361 


124138) 


Northern tropical 




75.2 


1515) 


2710) 


19101 


6115) 


Southern tiopical 




78.1 


2518) 


3014) 


26151 


81117) 


Southern temperate 


and pola 


130.1 


70 158) 


341151 


56141) 


1601114) 



130 WORLD ATLAS OF BIODIVERSITY 



Map 6.3 

Marine turtle diversity 

An overview of marine 
turtle diversity, represented 
by the number of turtle 
species nesting in any one 
area. Each symbol shows 
the location of a nesting 
site or area, colored 
according to the number of 
species present, up to a 
maximum of five species in 
some parts of the tropics. 

Source: Prepared using spatial data 
from a GIS database of marine turtle 
nesting beaches maintained at 
UNEP-WCI^C 




Number of species 
nesting in area 



Mammals 

Wholly aquatic mammals (those that never 
normally emerge on to land! are confined 
to two orders, the Cetacea and the Sirenia. 
The Cetacea comprises some 78 species, 
all except five marine, distributed throughout 
the worlds seas. They include the largest 
living animals - the rorquals in the family 
Balaenopteridae. All cetaceans are carniv- 
orous; the baleen or whalebone whales 
(families Balaenidae, Balaenopteridae, Esch- 
richtiidae and Neobalaenidael are filter 
feeders, feeding on organisms several orders 
of magnitude smaller than they are. 

Of four living members of the order Sirenia, 
only one - the dugong Dugong dugon - is 
exclusively marine, occurring widely in coastal 
waters of the Indo-Pacific. One other, the 
Caribbean manatee Trichechus manatus, is 




0-"^ 



• t 



'"Vfl^,,' 







found in both marine and inland waters 
while the other two Ithe Amazonian manatee 
Trichechus inungulsand West African manatee 
Trichechus senegalensis] enter coastal waters 
marginally if at all. One other species, the very 
large Steller's sea cow Hydrodamalis gigas, 
survived in waters around Bering and Copper 
Islands in the North Pacific until the early 18th 
century. All sirenians are herbivores; marine 
populations feed mainly on seagrasses. 

The remaining marine mammals are all 
included in the order Carnivora. Two New 
World otters in the family Mustelidae, the sea 
otter Enhydra lutris from the north temper- 
ate Pacific coast and the marine otter Lutra 
felina from the south temperate Pacific coast, 
feed very largely or exclusively in marine 
waters; other otter species may frequent 
coastal areas but are predominantly inland 



Marine biodiversity 131 




r^— C 
/, 



•1 ._v^ 




•\ 



water animals. Members of the tliree pinni- 
ped families Odobenidae (the walrus], 
Otariidae leared seals! and Phocidae (earless 
seals] are all largely aquatic, emerging on 
land to breed and rest, particularly when 
molting; all are marine with the exception of 
one or two species of Phocidae (the Baikal 
seal Phoca sibirica and, if the Caspian is 
regarded as a lake rather than a sea, the 
Caspian seal Phoca caspica]. One member of 
the family Phocidae, the Caribbean monk 
seal Monachus tropicalis, has become extinct 
this century. All species are carnivorous. 

In contrast to most terrestrial mammal 
families, pinnipeds are considerably more 
diverse and more abundant at higher rather 
than lower latitudes. Of the 32 extant or 
recently extant species, only five occur within 
the tropics (two marginally]. Part of the 






explanation for this undoubtedly lies in the 
greater availability of suitable habitat at 
higher latitudes; as noted above, 70 percent 
of continental shelf waters and just over 60 
percent of the world's marine area are found 
outside the tropics. However, this in itself is 
unlikely to account for the entire difference. 
It is probable that the greater productivity 
of shelf waters at high latitudes, discussed 
above, and of upwellmg areas at mid- 
latitudes (e.g. the Benguela current off the 
western coast of South Africa and the 
Humboldt current off Chile and Peru] plays a 
major part. 

The isolated character of many island 
breeding sites, such as the Galapagos group, 
in temperate and sub-polar parts of the 
southern hemisphere may also have en- 
couraged speciation of pinnipeds here. 



132 WORLD ATLAS OF BIODIVERSITY 



Table 6.8 

Diversity of mangroves 

Note: Where two figures are 
given, second figure indicates 
number of fiybrids. Families 
composed solely of mangrove 
species are stiown in bold. 



Source: Adapted from Duke and 
Spalding e( 3l. . 



COASTAL AND SHALLOW WATER COMMUNITIES 
Mangroves 

Mangrove woodland is indeed a truly hybrid 
terrestrial/marine ecosystem, unique in that 
terrestrial organisms can occur in the canopy 
and marine species at the base". Mangroves, 
or mangals, are a diverse collection of shrubs 
and trees lincluding ferns and palms) which 
live in or adjacent to the intertidal zone and 
are thus unusual amongst vascular plants in 
that they are adapted to having their roots at 
least periodically submerged in sea water A 
wide variety of organisms is associated with 
mangroves including a number of epiphytic, 
parasitic and climbing plants, and large 
numbers of crustaceans, mollusks, fishes 
and birds'l 

Mangrove species are generally divided 
into those found only in mangrove habitats 
and those that may also be found elsewhere 
but which are nevertheless an important 
component of mangrove habitats. Both 
groups come from a wide range of families. 
The former includes around 62 species and 
seven hybrids in some 22 genera (Table 6.81. 
The appearance of mangroves is far from 
uniform: they vary from closed forests 





^ ^^1 


^^^^■(o. of 


Filicopsida 


Adiantaceae 


3 


Plumbaginales 


Plumbaginaceae 


2 


Theales 


Pelliciceraceae 


1 


Malvales 


Bombacaceae 


2 




Sterculiaceae 


3 


Ebenales 


Ebenaceae 


1 


Primulales 


Myrsinaceae 


2 


Fabales 


Leguminosae 


2 


Myrtales 


Combretaceae 

Lythraceae 
Myrtaceae 


4+1 
1 
1 




Sonneratiaceae 


6+3 


Rhizophorales 


Rhizophoraceae 


17+2 


Euphorbiales 


Euphorbiaceae 


2 


Sapindales 


Meliaceae 


2+1 


Lamiales 


Avicennlaceae 


8 


Scrophuliariales 


Acanthaceae 
Bignoniaceae 


2 
1 


Rubiales 


Rubiaceae 


1 


Arecales 


Palmae 


1 



40-50 m high to widely separated clumps of 
stunted shrubs less than 1 m high". They are 
only able to grow on shores that are shelter- 
ed from wave action, and are particularly 
well developed in estuarine and deltaic 
areas; they may also extend some distance 
upstream along the banks of rivers, e.g. 
some 300 km up the Fly River in Papua New 
Guinea. 

Mangrove communities are largely re- 
stricted to the tropics between 30°N and 30°S, 
with extensions beyond this to the north in 
Bermuda and Japan, and to the south in 
Australia and New Zealand" (Map 6.4). They 
occur over a larger geographical area than 
coral reefs (see below) and, unlike reefs, are 
well developed along the western coasts of 
the Americas and Africa. They have a more 
restricted distribution than coral reefs in the 
South Pacific. 

There are two main centers of diversity. 
The eastern group occurs in the Indo-Pacific 
(the Indian Ocean and western part of the 
Pacific Ocean) and is the most species rich'^". 
The western group is centered around the 
Caribbean and includes mangrove com- 
munities along the west coast of the Americas 
and Africa. 

Global mangrove area is believed to slightly 
exceed 180 000 km', divided regionally as 
shown in Table 6.9. Mangroves occur in over 
100 countries (including dependent terri- 
tories) but exist in very small areas in many of 
these. Four countries (Indonesia, Brazil, 
Australia and Nigeria) between them account 
for over 40 percent of the world's mangrove 
area, and Indonesia alone possesses nearly 
one quarter of the global mangrove area". 

Although it is known that mangrove eco- 
systems in most parts of the world have been 
extensively degraded and cleared, it is difficult 
to obtain reliable data on the global extent 
of mangrove loss over time. Mangroves by 
their very nature occupy highly dynamic and 
unstable environments so that even without 
human action the location and extent of 
mangrove cover would be constantly chang- 
ing. One assessment" suggested that more 
than 50 percent of the worlds mangrove 
forest cover had been destroyed. 

Mangroves stabilize shorelines and de- 



Marine biodiversity 133 



crease coastal erosion by reducing the energy 
of waves and currents and by holding the 
bottom sediment in place with their roots. 
They also act as windbreaks and provide 
protection from coastal storms. They are 
generally highly productive ecosystems and 
are important habitats for crustaceans, 
shellfish and finfishes. Most of the larger 
commercial penaeid shrimps are mangrove 
dependent; these and other species are 
harvested both on a subsistence basis and 
commercially, and may provide a major source 
of income in some countries. 

As well as providing habitat for adults of 
many species of finfish and invertebrates, 
mangroves serve as spawning and nursery 
areas for many others, often of major 
economic importance. The wood provides 
building material, used locally in houses, as 
fence poles and to build fish traps, and is also 
harvested on a large scale for production of 
pulp and particle board. In many areas 
mangroves are also an important source of 
fuel, as firewood and charcoal. Mangrove 
foliage may provide an important source of 
fodder for domestic livestock in some coun- 
tries, particularly during dry seasons when 
other sources of greenery are in short supply. 

Salt marshes 

Salt marshes are coastal communities of 
rooted salt-tolerant Ihalophyticl plants of 
terrestrial origin, dominated by grasses, 
herbs and dwarf shrubs. They share many 
characteristics with mangroves but replace 
them geographically in higher latitudes, 
except for some overlap at the extremes of 
mangrove distribution (in the Gulf of Mexico, 
Japan, southern Australia and northern 
New Zealand). Globally, salt marshes are 
estimated to cover around 350 000 km' in 
total". They tend to develop in sheltered areas 
of mud and silty sand that are flat and slow 
draining; as plant growth and sedimentation 
elevates the marshland, so the period of tidal 
submergence decreases and a network of 
creeks develops. The two genera which are 
most prominent as pioneer saltmarsh plants 
are Salicornia, or samphire, and Spartina, or 
cord grass. Species of Puccinetla, Sc/rpus and 
Juncus are also common. 



Salt marshes are highly productive, around 
2 500 g C per m' per year Occurring in highly 
seasonal latitudes, salt marshes rapidly take 
up and accumulate nutrients during the 
growing season. In the autumn, when plants 
die or become dormant, the uptake of nutri- 
ents is greatly diminished and dead organic 
matter may be transported out of the marsh. 
The physical features of salt marshes mean 
that pollutants are not easily or rapidly 
flushed out. Other threats include infilling, 
especially around the North Sea", and in- 
creased erosion through channel dredging. 

Rocky shores 

Rocky shorelines are generally exposed to 
oceanic swells and extreme wave action 
(except for more sheltered fjordlandsl but are 
topographically variable, occurring as wide 
platforms, steep cliffs or other formations, 
depending on local geology™. They provide a 
unique habitat for plant growth, with a stable 
substrate for attachment, and shallow well- 
lit water that tends to be turbulent and 
rich in nutrients. Consequently macro-algae 
communities, dominated by kelps [Laminaria, 
Ecktonia, Macrocystis] and fuccoids [Fucus, 
Ascophyllum] flourish, mainly in temperate 
regions but also in areas of the tropics where 
seasonal upwellings of cold water occur, such 
as Chile and the southern Arabian coast-'. The 
net primary productivity of kelp forests 
is comparable to tropical rainforests, and 
Macrocystis pyrifera, the giant kelp, can attain 
growth of up to 45 cm a day 

Two physical factors - water movement 
and desiccation - have influenced the diversity 
of intertidal rocky shore species. Strong wave 
action has favored use of crevices, dense 
aggregations of individuals, and the evolution 
of strong attachment devices (algal holdfasts. 



Table 6.9 

Current mangrove cover 

Source: Spalding e( a/. . 



Lg^^^^pP 


^^f Mangrove area ^^H 
(to nearest thousand km') 


■|^Uftotal| 


■ 


South and Southeast Asia 
Americas 


75 000 
49 000 


42 
27 




West Africa 


28 000 


16 




Australasia 


19 000 


10 




East Africa and Middle East 


10 000 


6 




Total 


181 000 




m 



13« WORLD ATLAS OF BIODIVERSITY 



Map 6.4 
Mangrove diversity 

The location of current 
mangrove forest, togetfier 
witfi contours representing 
gradients of mangrove 
species ricfiness. Note 
tfiat grapfiic presentation 
at tliis scale enormously 
exaggerates actual forest 
area. 



Source; Reproduced with modification, 
from Spalding 




Mangrove locality and 
species richness 

• Mangrove forest 

High diversity 



Low diversity 



cementation of barnacles, the byssus threads 
of mussels and the adhesive feet of gastro- 
pods and echinodermsl. The need to avoid 
being washed off the rocks means that most 
organisms on rocky shores are sessile or have 
limited motile ability. Competition for space is 
therefore intense and organisms inhabiting 
rocky shores occupy well-defined zones. It 
also means that they can be particularly 
vulnerable to disturbances such as oil spills, 
especially if these occur during calm periods 
which extend the residence time of pollutants. 
However, because they are generally exposed 
and high-energy environments, rocky shores 
tend to be less vulnerable than most to 
pollution. These features also limit some 
destructive human activities, such as 
construction, responsible for degrading other 
marine habitats. 







r-qr 



Seagrasses 

Seagrasses are a mixed group of flowering 
plants Inot true grasses) that are adapted to 
live submerged in shallow marine and estuar- 
ine environments at a wide range of latitudes. 
About 58 species are recognized by many 
authorities, in four families, all within the 
monocotyledons. They occur from the littoral 
region to depths of 50 or 60 m but are most 
abundant in the immediate sublittoral area. 
There are more species in the tropics than in 
the temperate zones, and of the 12 seagrass 
genera seven are confined to tropical seas 
and five to temperate seas". Most seagrass 
species are similar in external morphology, 
with long thin leaves and an extensive 
rhizome root system which enables them to 
fasten to the substrate. A variety of substrates 
are occupied from sand and mud to granite 



Marine biodiversity 135 



fr'' 




rock, but the most extensive beds occur on 
soft substrates". 

While species diversity (see Map 6.51 is 
highest in Southeast Asia las with mangroves 
and corals) there are two important centers in 
temperate regions: in mainland Japan and in 
southwest Australia. Secondary centers of 
diversity include East Africa, the Red Sea 
and the Mediterranean. Although seagrasses 
themselves are not a diverse group, they 
support considerable diversity in some assoc- 
iated species, including an estimated 450 
species of epiphytic algae''. 

In many areas seagrasses form extensive 
but simple communities, referred to as sea- 
grass beds or seagrass meadows. Seagrass 
beds have high productivity and contribute 
significantly to the total primary production of 
inshore waters. Seagrasses are particularly 



important in nutrient-poor (oligotrophicl 
waters because they can extract some nutri- 
ents from sediments, unlike other marine 
primary producers. Many commercially impor- 
tant species are dependent on seagrass beds, 
often as nursery habitat, providing shelter 
from predators and adverse sea conditions. 
These include mollusks (such as the queen 
conch Strombus gigas], shrimp, lobster, holo- 
thurians and many finfish (such as grunts, 
Haemulidae; rabbitfish, Siganidae; emperors, 
Lethrinidae; and snappers, Lutjanidael. A 
small but important number of threatened 
species depend on seagrasses, including 
sirenians, the green turtle Chelonia mydas 
and many species of seahorse (Syngnathidae). 
In addition to such direct values, seagrass 
beds also play an important role in binding 
sediments, providing some protection from 



136 WORLD ATLAS OF BIODIVERSITY 



Map 6.5 

Seagrass species diversity 

This first global map of 
seagrass diversity indicates 
tfie extent of seagrass 
fiabitat (inventory sites) and 
stiovtfs contours of species 
ncfiness. Diversity contours 
not sfiown for parts of West 
Africa because species 
inventory incomplete. 

Source: Prelimtnarv plot, compiled using 
multiple sources, from a seagrass atlas 
in preparation at UNEP-WC(»1C. 



Seagrass locality and 
species richness 

• Seagrass 

High diversity 




Low diversity 



coastal erosion, and may help remove excess 
nutrients and toxins from coastal waters. 

A preliminary estimate'' suggested that 
seagrass beds extend over some 600 000 km' 
globally, but a precise figure is not yet 
available. Although seagrasses can be highly 
dynamic ecosystems, with individual seagrass 
beds undergoing significant shifts in distri- 
bution over relatively short periods, there is 
considerable evidence that there have been 
major net losses over the past century. 
Extensive losses along the Atlantic coasts of 
Europe and North America in the 1930s 
followed disease caused by a marine slime 
mold; some evidence suggests that other 
environmental impacts (possibly increased 
turbidity in coastal waters) may have increas- 
ed susceptibility to disease. More recently, 
nutrient loading has continued to increase in 



'^M'^ 






•^^' '•?-> 




v^, - 



i: 



coastal waters worldwide, often leading to 
enhanced growth of phytoplankton, epiphytes 
and macroalgae, all of which can out- 
compete seagrasses for available sunlight. 
Sedimentation also has a major impact, 
reducing the passage of light through the 
water column and physically smothering 
seagrass plants. To a lesser degree, toxic 
pollutants and physical disturbance from 
activities such as trawling and dredging have 
also played a role in seagrass losses". 

Shallow tropical coral reefs 

The term coral reef applies to a variety of 
calcium carbonate structures developed by 
stony corals. They are tropical shallow-water 
ecosystems, typically with high biodiversity 
and largely restricted to coastal seas between 
the latitudes of 30°N and 30°S", They are 



Marine biodiversity 137 



V^gi- 



"V, 






^k: 



-r^-.-: ii.'-^ 



> \ 




Vj: 



>^-. ^ 




most abundant in shallow, well-flushed 
marine environments characterized by clear, 
warm, low-nutrient waters that are of oceanic 
salinity". There are two basic categories: 
shelf reefs, which form on the continental 
shelves of large land masses, and oceanic 
reefs, which are surrounded by deeper waters 
and are often associated with oceanic islands. 
Within these two categories there are a num- 
ber of reef types: fringing reefs, which grow 
close to shore; barrier reefs, which develop 
along the edge of a continental shelf or 
through land subsidence in deeper waters, 
and are separated from the mainland or 
island by a relatively deep, wide lagoon; and 
atolls, which are roughly circular reefs around 
a central lagoon and are typically found in 
oceanic waters, probably originating from the 
fringing reefs of long-submerged islands. 



Two other less clearly defined categories are 
patch reefs, which form on irregularities on 
shallow parts of the seabed, and bank reefs, 
which occur in deeper waters, both on 
continental shelf and in oceanic waters". 

The global extent of coral reefs is not 
l<nown with certainty. A recent estimate, 
derived by measuring the total reef extent in 
a comprehensive set of national maps, sug- 
gests a world total of 28^ 300 km-. This is 
equivalent to a little more than 1 percent of 
the total world ocean shelf area. New 
information and mapping techniques may 
increase the accuracy of such estimates, but 
the total is unlikely to exceed 300 000 km'. 
A regional breakdown is provided in Table 
6.11. Although coral reefs occur in around 
110 countries and territories, just five 
countries - Indonesia, Australia, Philippines, 



138 WORLD ATLAS OF BIODIVERSITY 



Table 6.10 

Diversity of stony corals in 

the order Scleractinia 

Note: Includes only the reef- 
forming scleractinians with 
zooxanthellae. 

r- w 2' 

Source: Veron . 



France (overseas departments and territories 
in the Indian Ocean and Pacific) and Papua 
New Guinea - account for over half of the 
global total. 

Five different orders within the phylum 
Cnidaria (coelenteratesl include species with 
calcified skeletons, or stony corals'. Many of 
these are reef building Ihermatypicl and some 
are solitary lahermatypicl. The great majority 
of hermatypic corals belong to the order 
Scleractinia. the true stony corals, although 
not all scleractinians are reef-builders. 
Together with sea anemones and sea fans, the 
Scleractinia mal<e up the class Anthozoa. 
Most scleractinian coral polyps have symbiotic 
algae Izooxanthellael within their tissues; 
these use the nitrates, phosphates and carbon 
dioxide produced by the coral, and through 
photosynthesis generate oxygen and organic 
compounds that provide much of the polyps' 
nutrition. The zooxanthellae give corals their 
color and, because they photosynthesize, 
restrict the corals that contain them to the 
photic zone-'. Corals without zooxanthellae 
typically do not form reefs and can exist in 
deeper colder waters (see deep-water reefs, 
below]. Because of difficulties with synonymy 
and in defining species, it is difficult to 
estimate precisely how many extant species 



^l^y^^^^HPi 


Acroporidae 


4 


262 


Agaricidae 


6 


43 


Astrocoenidae 


i 


13 


Caryopiiylliidae 


1 


1 


Dendrophyllidae 


i, 


U 


EupiiylUdae 


5 


14 


Faviidae 


2i 


125 


Fungiidae 


13 


56 


Meandriniidae 


7 


8 


Merulinidae 


5 


12 


Mussidae 


13 


50 


Oculinidae 


4 


15 


Pectiniidae 


5 


28 


Pocilloporidae 


3 


30 


Poritidae 


5 


92 


Rhizangiidae 


1 


1 


Siderastreidae 


6 


28 


Trachypfiyllidae 


1 


1 



of reef-building coral there are. A recent 
synthesis-' deals with around 800 zooxan- 
thellate scleractinians (see Table 6.101; 
another" lists more than 1 300 scleractinians 
in all, and 260 calcified hydrozoans. 

Not all reefs are constructed primarily 
by corals. Within the red algae (phylum 
Rhodophytal and the green algae (phylum 
Chlorophytal in particular several genera 
grow heavily calcified encrustations which 
bind the reef framework and in places are the 
main contributors to shallow reef growth. 

Coral reefs are among the most productive 
and diverse of all natural ecosystems. The 
mam center of diversity for reet-building 
corals is Southeast Asia, with an estimated 
minimum of 450 species found associated 
with reefs around the Philippines, Borneo, 
Sulawesi and associated islands. This area 
IS part of a single, vast, Indo-West Pacific 
biogeographic province that extends from the 
Red Sea in the west to the Pitcairn Islands in 
the east. Many coral genera and a significant 
number of species are found throughout the 
region, although overall diversity in the pro- 
vince decreases on leaving this center. In the 
east of this region, the central and eastern 
Pacific forms a series of somewhat distinct 
subregions, characterized by a number of 
genera and species (particularly in Hawaii] 
not found further west. This area also shares 
many species with the Indo-West Pacific 
province but overall has much lower diversity 
than most of the latter The Atlantic, including 
the Caribbean and the Gulf of Mexico, forms a 
distinct province with few species in common 
with the Indo-West Pacific. It is also very 
depauperate compared with most of the latter 
(Map 6.6]. 

Deep-water reefs 

Beyond the coral reefs of shallow tropical 
waters, a few species of coral that lack 
zooxanthellae form deep-water reefs on hard 
substrates in high-current areas associated 
with topographic rises, such as ridges and 
pinnacles. The best known is Lopheiia 
pertusa, a colonial coral that forms structures 
ranging from patches a few meters in width 
to reefs many hundreds of meters in size, at 
depths of 100-3 000 m and temperatures of 



Marine biodiversity 139 



i-SX. Although best known through trawling 
and oil exploration activities in the northeast 
Atlantic, these deep-water reets are global in 
occurrence. 

The complex matrix of living and dead 
branches of Lophelia increases spatial 
heterogeneity above that of the surrounding 
seabed and provides a habitat for many 
species. Boring sponges, anemones, bryo- 
zoans, gorgonians, polychaetes, barnacles and 
bivalves occur in large numbers" and their 
diversity is comparable to some shallow- 
water tropical systems'". Although targe 
aggregations of fish are associated with 
Lophelia reefs, and reef areas support higher 
catches than adjacent seabed", the species 
diversity of fish and coral is much lower than 
in tropical coral reefs (some 23 species of fish 
have been recorded on Lophelia reefs in the 
northeast Atlantic!. 

The reefs are delicate structures easily 
destroyed by demersal fishing gear which 
routinely operate to depths of 2 000 m. The 
total destruction of some Norwegian reets 
has already been documented and an esti- 
mated 30-50 percent of others have been 
damaged. Slow growth, in the region of 
4-25 mm per year, severely limits the ability 
of Lophelia to recover The trawl fishery for 
orange roughy Hoplostethus atlanticus and 
oreos Allocyttus niger on sea mounts south 
of Tasmania has been responsible for sub- 
stantial destruction of Solenosmilia variabilis 
reefs, with some reduced to more than 90 
percent bare rocks". If recovery ever occurs it 
will take hundreds of years. 

OCEANIC PELAGIC COMMUNITIES 

There is a fundamental distinction between 
the processes and patterns observed in open 
oceans, dominated by global winds and large- 
scale vertical and horizontal movement of 
water masses, and those observed nearer to 
coasts, where shelf bathymetry, coastal winds 
and local input of nutrients, pollutants and 
sediments generate a diversity of smaller- 
scale phenomena. 

The oceanic pelagic zone is dominated by 
the activity of plankton in the euphotic surface 
waters. Plankton are by definition drifting or 
weakly swimming organisms, comprising a 



1 Region ^^^^^^^^^^H 


Shallow reef area 
Ito nearest thousand km'l 


% of world total jH 


Indo-Pacific 


261 200 


91.9 


Red Sea, Gulf of Aden 


17 400 


6.1 


Arabian Sea, Persian Gulf 


4 200 


1.5 


Indian Ocean 


32 000 


11.3 


Southeast Asia 


91700 


32.3 


Pacific 


115 900 


40.8 


Wider Atlantic 


2U00 


7,6 - 


Caribbean 


20 000 


7.0 


Atlantic (excl. Caribbean] 


1600 


0.6 


Eastern Pacific 


1600 


0.6 


Total 


284 300 





wide range of small to microscopic animals, 
protoctists and bacteria. Free-swimming 
pelagic organisms, predominantly fishes but 
also cetaceans and cephalopod mollusks 
(squid), are collectively termed nekton. These 
organisms, when adult, are predators of 
plankton or smaller nekton. They in turn - as 
vertically migrating fishes or larvae, and as 
dead organic material - provide food for deep- 
sea and benthic organisms. With few excep- 
tions, the only other food source for creatures 
in the aphotic zone is the 'rain' of organic 
matter, such as feces, molted crustacean 
exoskeletons, and a variety of other organic 
material derived from plankton in the surface 
waters of the ocean. Plankton and larger free- 
swimming organisms tend strongly to con- 
centrate along major circulation currents 
(gyresl, contact zones and upwellings, and 
this can give rise to significant local variation 
in diversity. 

The marked vertical gradients within the 
pelagic zone - of light, temperature, pressure, 
nutrient availability and salinity - lead to 
vertical structuring of pelagic species assem- 
blages. Several zones based on changes in 
species composition with depth have been 
recognized, including epipelagic (usually 
taken as from the surface to a depth of 200- 
250 m and including the euphotic zone); 
mesopelagic, which underlies the epipelagic 
zone to a depth of 1 000 m or so; and below 
this the bathypelagic which changes in a 
somewhat less well-defined fashion to 
abyssopelagic at around 2 500-2 700 m depth. 



Table 6.11 
Coral reef area 

Source: Spalding ef at. 



uo WORLD ATLAS OF BIODIVERSITY 



Map 6.6 
Coral diversity 

The location of coral reefs, 
togettier with contours 
representing gradients of 
species richness annong 
reef-building scleractmian 
coral species. Note that 
graphic presentation at 
this scale enormously 
exaggerates actual reef 
area. 

Source: Reproduced by permission, wilt 
modification, from Veron . revised by 
Veron, pers. comm. November 2001. 



Coral locality and 
species richness 



Coral 

High diversity 




Low diversity 



These zones, however, tend to fluctuate in 
time and space. As well as seasonal changes 
in water characteristics, many components of 
the epipelagic and mesopelagic nel<ton under- 
go marked diet migrations (i.e. on a 24-hour 
cycle], ascending to surface waters at night to 
feed and descending, sometimes over 1 l<m, 
during the day. Many species of nel<ton, 
particularly cetaceans and larger fishes, are 
also highly migratory, ranging over enormous 
expanses of ocean in more or less regular and 
predictable patterns. 

It has generally been assumed that bio- 
mass in the pelagic zone everywhere below 
the euphotic zone is low. However, recent 
studies have indicated that biomass of tropical 
mesopelagic animals may be surprisingly 
high. Study of the mesopelagic fauna has been 
limited to date, as it requires the use of 





expensive high-seas research vessels; i<now- 
ledge of taxonomy, distribution and biology of 
most of the species concerned remains very 
incomplete'. One study" recognized around 
160 fish genera in 30 families as important 
components of the fauna. Most species are 
small (less than 10 cm in length) and often 
bizarrely shaped. Estimates based on a variety 
of surveys carried out indicate that global 
biomass of this stocl< may be large: a figure of 
650 million tons (some six to seven times total 
current marine fisheries landings! has been 
suggested, although this should be regarded 
with extreme circumspection'. 

From available data, it would appear that 
the mesopelagic biomass is greatest in the 
northern Indian Ocean, and particularly in the 
northern Arabian Sea, one of the five major 
upwelling zones. Surveys here indicated 



Marine biodiversity 141 




extremely high blomass 125-250 g per m') in 
the Gulf of Aden and Gulf of Oman as well as 
off the western coastline of Pakistan. These 
figures are around an order of magnitude 
higher than those recorded elsewhere in the 
tropics, indicating either a great overestimate 
for the northern Indian Ocean or an under- 
estimate elsewhere. Alternatively this region 
genuinely is ten times as productive as the rest 
of the tropical ocean system. Although this 
appears as yet unresolved, it is nevertheless 
apparent that there is substantial global 
mesopelagic fish biomass ar.d that the Arabian 
Sea is particularly rich in these species. 

DEEP-SEA COMMUNITIES 

Approximately 51 percent of the Earth's 
surface is covered by ocean over 3 000 m in 
depth, so deep-sea communities are preva- 



lent over a significant proportion of the planet. 
All deep-sea habitat is in the aphotic zone, 
well below the distance sunlight can pene- 
trate. As deeper and deeper levels are 
reached biomass falls exponentially^'. 

Despite their enormous volume, the deep 
oceans appeared to be relatively simple 
ecosystems and to make little contribution to 
global species diversity, but discoveries 
during the past decade or so have shown that 
in some regions species diversity in the 
benthic community increases with increasing 
depth. This was revealed by novel sampling 
techniques, principally the epibenthic sled^^ 
The rate of discovery of new species and the 
proportion of species currently known from 
only one sample both suggest that a great 
number remain to be discovered"". As 
with arthropods in tropical moist forests. 



142 WORLD ATLAS OF BIODIVERSITY 



estimates for the number of unknown species 
vary widely, witfi some suggestions tfiat there 
may be as many as 10 million undescribed 
species in the deep sea". Others consider that 
the true figure is more likely to be around 
500 000". 

Ocean trenches 

Ocean trenches are typically close to land 
masses and tend to have high rates of 
sedimentation, a significant amount of which 
is of organic origin and an important available 
food source for trench communities. Several 
trenches also underlie highly productive cold- 
water upwelling zones, the organic fallout 
from which contributes greatly to productivity 
there. The water within trenches generally 
originates from the surrounding bottom 
water, which is derived from cold surface 
water at high polar latitudes and is relatively 
well oxygenated"'. 

Trenches tend to be isolated linear systems 
that because of their seismic activity form a 
habitat that is unstable and unpredictable 
compared with the relative environmental 
stability of the adjacent abyssal plains"'. 
Trench faunas are not rich in species but are 
often high in numbers of endemic species. 
There are some 25 genera restricted to the 
ultra-abyssal [hadall zone, representing some 
10-25 percent of the total number of genera 
present, and two known endemic hadal 
families: the Galatheanthemidae (Cnidarial 
and Gigantapseudidae (Crustacea). The latter 
family contains a single species: Giganta- 
pseudes adactylus. The greatest number of 
endemic species known from a single trench 
is a sample of 200 from the Kurile-Kamchatka 
trench; this maybe compared with 10 endemic 
species known from the Ryukyu and Marianas 
trenches. 

Hydrothermal vents 

Hydrothermal vent communities were first 
discovered in 1977, at a depth of 2 500 m on 
the Galapagos Rift, but contribute to what 
might be one of Earths most archaic eco- 
systems. They are now known to be assoc- 
iated with almost all known areas of tectonic 
activity at various depths. These tectonic 
regions include ocean-floor spreading centers, 



subduction and fracture zones and back-arc 
basins". Cold bottom water permeates 
through fissures in the ocean floor close to 
ocean floor spreading centers, becomes 
heated at great depths in the Earth's crust 
and finds its way back to the surface through 
hydrothermal vents. The temperature of vent 
water varies greatly, from around 23°C in the 
Galapagos vents to around 350°C in the vents 
of the East Pacific Rise, and they may be rich 
in metalliferous brines and sulfide ions'". 
Most species live out of the main flow at 
temperatures of around 2°C, the ambient 
temperature of deep-sea water The biomass 
of vent communities is usually high compared 
with other areas of similar depth, and dense 
colonies of tube worms, clams, mussels 
and limpets typically constitute the major 
components. 

Hydrothermal vent communities are of 
particular interest because they flourish 
in the dark at high pressures and low 
temperatures", and unique because they are 
supported by chemolithoautotrophic archeans 
and bacteria, notably Thiomicrospira species 
(phylum Proteobacteria), which form dense 
microbial carpets in the rich hydrothermal 
fluid and derive their energy chiefly from 
oxidizing hydrogen sulfide"". Many of the 
eukaryote vent species filter-teed on these 
microorganisms, whilst others rely on sym- 
biotic sulfur bacteria for energy". 

The overall species diversity at vents is low 
compared with other deep-sea soft sediment 
areas", but endemism is high. More than 20 
new families or subfamilies, 50 new genera 
and nearly 160 new species have been 
recorded from vent environments, including 
brine and cold seep communities"". Vent 
communities are separated by gaps of 
between 1 and 100 km, and although they may 
persist only for several years or decades, sites 
of vent activity move relatively slowly allowing 
dispersal of vent organisms". 

Two further seep patterns are known. Cold 
sulfide and methane-enriched groundwater 
seeps occur near the base of the porous 
limestone of the Florida escarpment, as well 
as in the Gulf of Mexico and elsewhere. These 
support a community similar in taxonomic 
composition to the hydrothermal vents of the 



Marine biodiversity 143 



''% 



east Pacific. Tectonic subduction zone seeps 
are more diffuse and lower in temperature 
than hydrothermal vent seeps and are rich in 
dissolved methane. They are known to occur 
off Oregon and in the Guaymas basin in the 
Gulf of California, and cold seeps occur at a 
depth of 1 000 m in Sagami Bay near Tokyo 
and in the subduction zones of the trenches 
off the east coast of Japan. 

HUMAN USE OF AND IMPACT ON THE OCEANS 

The seas provide many biological resources 
used by humans. A wide range of animal 
species, notably fishes, mollusks and crus- 
taceans, contribute to marine fisheries and 
these provide by far the most Important 
source of wild protein, of particular Impor- 
tance to many subsistence communities 
around the world. Marine algae are also an 
Increasingly important foodstuff, notably in 
the Far East, with current annual world 
production of around 2 million metric tons. 
Marine organisms are also proving extremely 
fruitful sources of pharmaceuticals and other 
materials used In medicines. Relatively minor 
although locally Important uses include 
exploitation of coastal resources for building 
materials (e.g. coral limestone, mangrove 
poles] and other industrial products (e.g. 
tannins from mangroves). 

Access to marine resources is not equitably 
distributed amongst the world's nations. Most 
obviously, some 39 states are landlocked, i.e. 
have no seaboard (five of these border the 
Inland Caspian Seal. Those that do have sea- 
boards show great variation in length of 
coastline, and area of territorial waters and 
exclusive economic zones lEEZsl, both abso- 
lutely and relative to their land areas. They also 
show great variation In their capacities to 
exploit marine resources, both on the high seas 
and within their territorial waters and EEZs. 

Human activities, directly and indirectly. 
are now the primary cause of changes to 
marine biodiversity. Approximately one third 
of the world's human population lives in the 
coastal zone Iwithin 60 km of the seal and 
indications are that this proportion will rise 
during the 21st century". Pressures exerted 
by the human population on the marine 
biosphere are substantial and Increasing. 




-:^# tr^^ 



-^ ^s«. 




Most identified threats relate to coastal 
and Inshore (continental shelf] areas. How- 
ever, threats to the oceanic realm are 
undoubtedly Increasing: fisheries and their 
attendant physical effects, such as habitat 
alteration owing to dredging and trawling, 
have entered deeper continental slope waters 
having previously been largely confined to the 
epipelagic zone, and deep-water oil and gas 
mining is planned. Even abyssal and hadal 
areas are susceptible to human Impact. A 
small, steady increase In abyssal temperature 
of 0.32°C In 35 years has been attributed to 
global climate change brought about by 
human activities. Ocean waste dumping and 
the potential for deep-water mining and 
mineral extraction are also causes for 



Many species of fishes, 
mollusks and crustaceans 
provide humans with 
their largest source of 
wild protein. 



u« WORLD ATLAS OF BIODIVERSITY 



Just five species of finfish. 
among them tfie Atlantic 
herring, account for a 
quarter of the global catch. 



concern, as are the changes in biomass and 
species composition in the waters above 
these regions". 

The following five activities have been 
identified as the most important agents of 
present and potential change to marine bio- 
diversity at genetic, species and ecosystem 
levels": 

• fisheries operations; 

• chemical pollution and eutrophication; 

• alteration of physical habitat; 

• invasions of exotic species; 

• global climate change. 

WORLD MARINE CAPTURE FISHERIES 

World marine capture fisheries have grown 
fivefold in the past half-century, with annual 
landings increasing from nearly 18 million 
metric tons between 1948 and 1952 to around 
87 million metric tons during the period 199A- 
97, with a fall caused mainly by El Nino in 1998 
and a subsequent recovery in 1999. Marine 
capture fisheries made up just over 70 percent 
of current recorded global production of 
aquatic resources in the late 1990s, the 




l^".'M 



remainder being accounted for by inland 
capture fisheries Isee Chapter 7| and 
aquacutture"". With capture fisheries appar- 
ently remaining more or less stable, the 
increase in total marine production during the 
1990s was due to a continuing increase in 
aquaculture production. 



Composition of marine fisheries 
Marine fisheries encompass a wide range of 
organisms, including algae, invertebrate ani- 
mals in various phyla and vertebrates including 
fishes (often termed finfishes in fisheries analy- 
sis], reptiles, mammals and birds (although by 
convention the last of these groups is not 
normally considered in fisheries analysis). 

The Food and Agriculture Organization of 
the United Nations IFAOI recognizes in total 
just under 1 000 species items' (species, 
genera or families! that feature at least 
periodically in national catch statistics. 
However, globally important marine fisheries 
are confined to relatively few groups, with 
over 80 percent of landings by weight being 
finfishes and virtually all the remainder 
mollusks and crustaceans. 

In terms of major species groups, by far the 
most important are the herrings and anchovies 
in the order Clupeiformes, which in 1998 
accounted for over 22 million metric tons, or 
around 25 percent of marine landings. These 
are followed by cod, hake and haddock 
(GadiformesI, and jacks and mullets (some Per- 
ciformes and MugiliformesI, with production of 
more than 9 million and nearly 8 million metric 
tons respectively The most important inver- 
tebrate group overall is cephalopod mollusks 
(squid, cuttlefish and octopus! of which some 
3.A million metric tons were reported landed. 

In terms of individual species, for several 
years during the 1990s just five species of 
finfish, anchoveta Engraulis ringens, Alaska 
pollock Theragra chalcogramma, Chilean jack 
mackerel Trachurus murphyi, Atlantic herring 
Clupea harengusar\6 chub mackerel Scomber 
japonicus, together made up around one 
quarter of marine landings. Each accounted for 
over 2 million metric tons annually, and among 
several others around this level, Japanese 
anchovy Engraulis japonicus and Skipjack 
tuna Katsuwonus pelamis were increasingly 
important. In most years by far the most im- 
portant single species is the anchoveta 
Engraulis ringens, whose fishery (off the west 
coast of South America) was nearly 13 million 
metric tons in 1994, constituting by far the 
largest single-species fishery the world has 
ever seen; but just under 8 million in 1 997 and 
less than 2 million in 1998. 



Marine biodiversity 145 



^* 



Distribution of marine fisheries 

The geographical distribution of marine 
fisheries is determined both by the distri- 
bution of harvestable fish stocl<s and by a 
range of complex socioeconomic factors. The 
former is largely determined by variations in 
productivity, which, as noted above, are them- 
selves largely determined by nutrient availa- 
bility, so that overall the most productive 
fisheries areas are on continental shelves at 
higher latitudes and in upwelling zones at 
lower latitudes. As a generalization, the latter 
are associated with pelagic fish stocks and 
the former more with demersal or semi- 
demersal (deep-water or bottom-dwelling) 
stocks, although pelagic stocks play an in- 
creasingly important role even here. 

As might be expected purely on the basis 
of its size, the Pacific Ocean is by far the most 
important major fisheries area, accounting 
for over 60 percent of marine landings. The 
northwest Pacific alone - an area with 
extensive continental shelf development - 
accounts for nearly half this total. 

The various upwelling zones are not all of 
equal importance in fisheries. That associated 
with the Humboldt current off Peru and Chile 
is the single most productive, while those 
associated with the California, Benguela and 
Canary currents are of somewhat lesser 
importance, although each is still a major 
fisheries area. The Arabian Sea upwelling 
appears to be anomalous, in that it evidently 
supports major populations of mesopelagic 
li.e. middle-depth) rather than epipelagic 
species. Not only are the former generally 
considered of low value, with an identified 
market only as animal feed, but capture and 
processing requires expensive, advanced 
technology They thus remain virtually un- 
exploited at present and are considered along 
with the Antarctic krill stocks to be the major 
unexploited fisheries resource left. 

Trends in marine fisheries 

National fishery statistics are collated by the 
FAG. These data are the principal source of 
information on global fishery trends, although 
it is widely acknowledged that they are 
variable in quality During the 1950s and 
1960s, total landings increased steadily as 




■ 1996 ^Sl 1998 

Million metric tons 
2 4 6 8 10 

i ' \ I 1 1 

Alaska pollock (Theragra chalcogrammal 
4.5 
I 4.0 

, Atlantic herring IClupea harengusi 
2.3 

2.4 

I Japanese anchovy lEngraulisjaponicusI 
1.3 
I 2.1 

Cack mackerel ITrachurus murphyil 
4.4 
2.0 

Skipjack tuna IKatsuwonus pelamisi 
1.6 
1.9 

I Chub mackerel IScomber japonicusi 
2.2 
1.9 




Largehead hairtail ITrichiurus lepturusl 

1.3 

HH 1.4 

Atlantic cod ICadus morhuaj 

1.3 

^1 1.2 

Yellowfin tuna IThunnus albacaresl 

1.1 

^1 1.2 

Blue whiting IMicromesistius poutassoul 

D_6 
HI 1.2 



new stocks were discovered, while improved 
fishing technology and an expansion of fishing 
effort enabled fuller exploitation of existing 
stocks of both pelagic and demersal species. 
Long-range fleets increased in size during 
this period and, as traditional fishing grounds 
in the North Atlantic and North Pacific 
became fully exploited, moved into new fish- 
ing grounds closer to the tropics and in the 



Figure 6.1 

Species contributing most 

to global marine fisheries 



U6 WORLD ATLAS OF BIODIVERSITY 



Figure 6.2 

Marine fisheries landings 

by major group 

Note: 1984-99 data are 
for capture fisheries only: 
pre-1984 data include 
aquacuUure. 



50 



W 



S30 



= 20 



10 



southern hemisphere. By concentrating their 
efforts in the richest ocean areas, these fleets 
were largely responsible for the rapid 
increase in world catches. 

At the beginning of the 1 970s, the Peruvian 
anchoveta fishery alone contributed some 20 
percent of marine fisheries production. These 
stocks collapsed around 1972, at the same 
time as the important South African pilchard 
fishery in the Atlantic, seemingly in asso- 
ciation with an ENSO lEl Nino Southern 
OscillationI event. There was a sharp drop in 
overall marine fisheries production, after 
which the global catch increased more slowly 
than before, reaching the early 1970s level 
by the end of the decade. Landings of most 
demersal fish stocks remained relatively 
constant, however, implying that they were 
close to full exploitation. Long-range fleets 
continued to expand in importance. 

The 1980s once again saw a period of 
continuous growth (averaging 3.8 percent a 
year! in reported world landings. As in the 
1970s landings of demersal stocks were 
generally static or declining so that shoaling 
pelagic species provided most of the increase 
in fish production. In fact, just three pelagic 
species [Peruvian anchoveta. South American 
sardine Sardinops sagax, and Japanese 



Pelagic 

Demersal 

Species not elsewhere included 

Crustaceans 

Mollusks lexcl. cephalopodsl 

Cephalopods 




1950 



1999 



sardine Sardinops melanostictus] and one 
semi-demersal species (Alaska pollock) 
accounted for 50 percent of the increase in 
world landings during the 1980s". Most of 
this increase appears to have been because 
of favorable climatic effects on stock sizes 
rather than new fishery developments or im- 
proved management practices". 

Following a sharp decline at the end of the 
1980s, FAQ data indicate slow net growth in 
marine capture fisheries through the 1990s. 
Four of the five most important fishes in 
fisheries in the late 1990s are pelagic, the 
exception being the Alaska pollock. This 
dominance of pelagic over demersal species 
is reflected in overall fisheries figures, with 
pelagic landings well over twice demersal 
landings globally. This contrasts sharply with 
the situation in the early 1950s when pelagic 
landings were only some 30 percent greater in 
volume than demersal landings (Figure 6.21. 

This increasing dependency on pelagic fish 
stocks is symptomatic of a major crisis in 
global marine fisheries. In general demersal 
fishes are more valuable per unit weight than 
pelagic species so that all else being equal 
the former are preferentially harvested. 
The increased importance of the latter in the 
past AO years is indicative of the growing 
overexploitation of fisheries stocks worldwide 
- as valuable demersal stocks have been 
depleted so attention has turned to the 
intrinsically less valuable pelagic stocks. 

Of some 441 fishery stocks for which status 
data were available in 1999, FAO considered 
that only 4 percent were underexploited, and 
with a further 21 percent assessed as 
moderately exploited, around 25 percent 
of stocks analyzed were above the level of 
abundance thought to correspond to maxi- 
mum sustainable yield (ivlSYI level (or have a 
fishing capacity below this level). The remain- 
ing 75 percent of stocks were considered to 
require strict control of fishing capacity and 
fishing effort in order for them to recover to 
MSY biomass". The proportion of stocks in 
this condition has increased between 1974, 
when first systematically reviewed, and 1999 
(Figure 6.3). In terms of ocean regions, the 
situation worsened steadily in the North 
Atlantic and North Pacific until the early 



Marine biodiversity 147 



•^*l* 



1990s, when there were signs of possible 
stabilization, mainly in the former. Stocks 
appear still to be in decline in the tropical and 
southern parts of these oceans, with some 
possible stabilization in the tropical Atlantic, 

Recent analysis of fishery data suggests that 
the widespread overexploitation of marine 
fisheries is probably more serious than the 
global catch statistics indicate because mis- 
reporting by countries with particularly large 
fisheries, coupled with wide fluctuation in 
Peruvian anchoveta stocks (linked with El Nino 
events], can produce spurious trends at global 
level. When more realistic estimates of the 
catch in China, for example, are substituted for 
reported catch figures believed to be incorrect, 
the global catch appears to have declined by 
0.36 million metric tons annually since 1988, 
rather than increased by 0.33 million metric 
tons (Figure 6.A1. The declining trend is much 
steeper if the pelagic Peruvian anchoveta are 
excluded^". 

There are three major reasons for the 
declining state of many marine fisheries. First, 
and most fundamental, most fisheries have 
traditionally been regarded as an 'open access' 
resource, so that, in effect, it pays any one 
fisher to harvest as much as possible at any 
given time because, if they do not, somebody 
else will. Secondly, technological innovations 
have made fishing much more efficient. 
Thirdly, there has been high investment in the 
world's commercial fishing fleet (partly a 
consequence of the nature of fisheries as an 
open access resource but also for complex 
socioeconomic and political reasons!. 

Bycatches and discards 

The effects of overfishing are compounded by 
the wastefulness of many marine capture 
fisheries. FAO estimated in 1994 that global 
marine fisheries bycatch and discards 
amounted to 18-40 million metric tons (mean 
27 miUionI (Map 6.7|. This represented just 
over 25 percent of the annual estimated total 
catch (i.e. landings represent around 75 
percent of actual catchl. Although figures are 
not available, it is generally assumed that the 
great majority of discards die. Further losses 
are caused by the mortality of animals which 
escape from fishing gear during fishery 




Overexptoited ^ 
depleted + 
recovering 



1970 



1975 



1980 



1985 



1990 



1995 



2000 



2005 



operations, but it is impossible at present to 
estimate the importance of this. Shrimp fishing 
produces the largest volume of discards 
(around 9 million metric tons annually). 

Bycatches include non-target, often low- 
value or 'trash' species, as well as undersized 
fish of target species. Non-target species 
may include marine mammals, reptiles (sea 
turtles! and seabirds, as well as finfishes 
and invertebrates. Of particular concern in 
recent years has been mortality of marine 
mammals, especially dolphins, in pelagic drift 
nets, of sea turtles in shrimp trawls and 
more recently of diving seabirds, especially 
albatrosses, in long-line fisheries. Discarding 



Figure 6.3 

Global trends in the state 

of world stocks since 1974 



Source: Adapted (rom Figure iO in 
FAo". 



Figure 6.4 

Trends in global fisheries 

catch since 1970 

Source: Adapted from Watson and 
Pauly=°. 




1970 



1975 



1980 



1985 



1990 



1995 



2000 



us WORLD ATLAS OF BIODIVERSITY 



Map 6.7 

Marine fisheries catch and 

discards 

The location of the fishery 
areas recognized by FAO for 
statistical purposes is 
shown on this map, with 
symbols representing the 
approximate late 1990s 
yield from capture fisheries 
and the volume of 
discarded catch, most of 
which IS presumed not to 
survive. Each symbol 
represents approximately 
1 million metric tons. 

Source: Data from FAO , 




Fisheries catch 

<C^^^^!^ 1 million metric tons landed 

^ \ {{{h" i 1 million metric tons discarded 



may be a side-effect of management systems 
intended to regulate fisheries (e.g. non- 
transferable quotas may cause discarding of 
over-quota catch; species-specific licensing 
may cause discard of non-licensed but still 
commerciaUy valuable speciesl. 

Solutions to bycatches and discards will 
be found essentially through improvement in 
the selectivity of fishing gear and fishing 
methods. Much of the research in this has 
been carried out in higher latitudes and is 
not readily transferable to multispecies 
tropical fisheries, where the tropical shrimp 
trawls still produce high rates of bycatch. 
Improved use of bycatch either as fishmeal 
or human food is also a possibility; however, 
this does not address the problem of 
mortality of potentially threatened species 
(sea turtles, seabirds, cetaceans), nor the 



wasteful capture of immature specimens of 
harvestable species. 

A further problem in the efficient use of 
marine resources is post-harvest loss. It is 
almost impossible to estimate this accurately, 
but FAO believes it to exceed 5 million metric 
tons per year li.e. around 5 percent of harvest!. 
Most significant are physical losses of dried 
fish to insect infestations and loss of fresh fish 
through spoilage. These problems are partic- 
ularly significant in developing countries. 

AQUACULTURE 

One major response to the growing crisis in 
marine capture fisheries has been the rapid 
rise in various forms of aquaculture (Figure 
6.5). The latter may be defined as the rearing 
in water of organisms (animals, plants and 
algae) in a process in which at least one 



Marine biodiversity 149 




phase of growth is controUed or enhanced 
by human action. The animals used are gen- 
erally finfishes, mollusks and crustaceans, 
although a number of other groups such as 
sea squirts ITunicatal, sponges (Poriferal and 
sea turtles are cultured in small quantities. 
Seaweeds of various kinds are also cultured, 
some in large amounts. Most of the species 
grown in any quantity are low in the food 
chain, being either primary producers, filter- 
feeders or finfishes that in their adult stages 
are either herbivores or omnivores. 

FAO notes that aquaculture is the worlds 
fastest growing food production sector, 
annual output having increased at an average 
rate of some 10 percent in the period 1984-98 
(compared with less than 2 percent for 
capture fisheries) (see Figure 6.5). In 1999 
aquaculture provided around one quarter of 



recorded global fisheries production. Of the 
total 32.9 million metric tons recorded in 
1999, almost 20 million originated inland, and 
nearly 13 million were produced in marine 
and brackish environments". In 1996. some 
7.7 million metric tons of algae and plants 
were produced, almost all of this seaweed, 
chiefly Japanese kelp Laminaria japonica, 
nori Porphyra tenera and wakame Undaria 
pinnatifida. The first of these was, in terms of 
volume, the most important of all aquaculture 
species, with around i.4 million metric tons 
produced. 

In marine and brackish (usually estuarine) 
environments, by far the m.ost important 
animal group in terms of volume is the moll- 
usks, whose 1997 recorded production of 
some 8.6 million metric tons made up more 
than 75 percent of all animal production in 



150 WORLD ATLAS OF BIODIVERSITY 



10 



■^i. 



Seaweeds 
MoUusks 

Diadromous fishes 
Marine fishes 
Crustaceans 
Miscellaneous 
aquatic animaLs 




1984 

Figure 6.5 

Marine aquacuUure 

production 



1998 

these environments. Around 50 moUusk 
species are produced in significant quantity, 
almost all bivalves. As with most culture 
systems, production is heavily skewed to a 
small number of species, with 65 percent of 
production composed of just three: the Pacific 
cupped oyster Crassostrea gigas, Japanese 
carpet shell Ruditapes philippinarus and 
Yesso scallop Pectenyessoensis. The Far East 
dominates production, with around 75 percent 
of that recorded taking place in China and 
most of the remainder in Japan. 

Although production of marine crusta- 
ceans accounts for only 10 percent or so by 
volume of marine and brackish water animal 
aquacuUure, it has disproportionately high 
economic importance, and is also the sector 
that has given rise to most environmental 
concerns. Between 1984 and 1998 annual 
production grew nearly sixfold, from less than 
200 000 to over 1 million metric tons. The 
great majority of production takes place in 
tropical and subtropical Asia and is dominated 
by Penaeus species; globally this genus pro- 
duces over 90 percent of aquacuUure crusta- 
cean supply by weight. Three species of 
Penaeus account for around three quarters of 
crustacean production. The giant tiger prawn 
P. monodon is the most widely cultivated and 
accounts for nearly half; the whiteleg shrimp 
P. vannamei is cuUured in the Americas and 
accounts for around 15 percent of estimated 
global supply (around 70 percent of this 



originating in Ecuador); and the fleshy prawn 
P. chinensis is cultured in China and currently 
accounts for around 10 percent of production, 
having declined considerably since the early 
1990s when around 200 000 metric tons were 
produced annually. Other marine crustaceans 
cultivated include other Penaeus species, 
some Metapenaeus, and spiny lobsters 
Panulirus. These groups, however, make an 
insignificant contribution to global supply. 

Growth in crustacean aquacuUure has been 
fuelled by the high value of the product: the 
market in 1996 was estimated to be worth 
nearly US$7.5 billion, or around one quarter of 
the total value of marine and brackish water 
aquacuUure". The great majority of production 
takes place in low-income countries - the five 
countries producing over 100 000 metric tons 
annually being China, Thailand, Indonesia, 
Ecuador and Bangladesh - and is aimed mainly 
at the export market (primarily to Europe, the 
United States and Japan) and to a lesser extent 
at the domestic luxury market. Pressure is 
high to produce maximum returns on invest- 
ment so that increasingly intensive farming 
methods are used. These are widely acknow- 
ledged to be having adverse social and 
environmental impacts in the countries of 
production, as well as leading to increasing 
difficuUies in maintaining supply, owing to the 
spread of major diseases. The last of these 
accounts for the major decline in the Chinese 

leshy prawn industry during the 1990s. 

mpacts include: 

loss of mangrove habitat; 
abstraction of freshwater; 
introduction of pathogens and other 
damaging non-native species; 
escape of cultured non-native species; 
pollution; 

diversion of low-quality or cheap fish food 
resources (may lead to more efficient use 
of bycatches and trash fish but also to 
more indiscriminate catch fisheries); 
• diversion of effort from other forms of aqua- 
culture (notably milkfish Chanos chanos]. 

The aquarium trade 

Up to 2 million people worldwide (about half in 
the United States and a quarter in Europe! are 



Marine biodiversity 151 



■"^'* 



thought to keep marine aquanunns, most of 
which are stocl<ed with wild-caught species. In 
1997 a total of 1 200 metric tons of coral was 
traded internationally, with 56 percent impor- 
ted by the United States and 15 percent by the 
European Union. Approximately half of this was 
live coral for aquariums, a tenfold increase on 
the amount of live coral traded in the late 
1980s^'. Qualitative estimates of trade suggest 
that 14-30 million fish may be traded per year, 
representing some 1 200 species, about two 




thirds of which are from coral reefs. Aquarium 
species are typically gathered by local fishers 
using live capture techniques or chemical 
stupefactants (such as sodium cyanidel which 
are non-selective and adversely affect the 
health of specimens as well as killing non- 
target organisms. Inappropriate shipping 
methods and poor husbandry along the supply 
chain often cause high mortality among the 
fish and invertebrates collected. 

While the current impacts of the aquarium 
trade remain poorly known, the industry has 
considerable potential to contribute to sus- 
tainable development. It is relatively low in 
volume but very high in value - a kilo of 
aquarium fish from one island country was 
valued at almost US$500 in 2000, whereas 
reef fish harvested for food were worth only 
US$6*'. Aquarium species are a high-value 
source of income in many coastal com- 
munities with limited resources, with the 
actual value to the fishers determined largely 
by market access. In Fiji many collectors pay 



an access fee to the villages to collect on their 
reefs, but by selling directly to exporters they 
can have incomes many times the national 
average. By contrast, in the Philippines there 
are many middlemen, and collectors them- 
selves typically earn only around US$50 per 
month. Targeting mostly non-food species, 
aquarium fisheries could in principle provide 
an alternative economic activity for low- 
income coastal populations and an important 
source of foreign exchange for national 
economies, as well as an economic incentive 
for the sustainable management of reefs. 
The application of international certification 
schemes may provide an important tool for 
achieving this. 

OTHER (vlAJOR IMPACTS ON THE MARINE 
BIOSPHERE 

Alteration of physical habitat 
Physical alteration of habitats through human 
action chiefly affects coastal and inshore 
areas. Impacts here can be severe, although 
few attempts have been made to quantify 
them on a global basis. Major causes include 
coastal development, particularly landfilling 
and construction of groynes and jetties, aqua- 
culture, dredging of channels for navigational 
purposes, extraction of materials such as 
sand and coral, stabilization of shorelines, 
and destructive fishing methods such as 
beam-trawling, use of explosives and muro- 
anni (using rocks on ropes to drive fishes into 
netsl. Upstream activities, such as defores- 
tation and dam construction can greatly alter 
sediment loads in rivers, affecting patterns of 
sediment deposition in estuarine areas. 

Chemical pollution and eutrophication 

Human activities have increased inputs of a 
huge range of organic and inorganic chem- 
icals into marine ecosystems. Such inputs 
may enter by direct discharge (e.g. in sewage 
outflow pipes], via river and stream outflow, 
as land runoff, through the atmosphere or 
from seagoing vessels. Because virtually all 
such input originates on land, as with most 
other human Impacts on marine ecosystems, 
areas most affected are coastal and inshore 
regions, particularly enclosed or semi- 
enclosed water bodies. In oceanic regions. 



The aquarium trade targets 
mostly non-food species 
and many invertebrates 
such as crabs, anemones 
and shrimp. 



152 WORLD ATLAS OF BIODIVERSITY 



Worldwide, human 
activities have increased 
inputs of nitrogen and 
phosphorus in rivers and 
coastal waters fourfold 



mixing of the enormous volume of sea water 
generally ensures that inputs become rapidly 
diluted. 

Major categories of input include nutrients 
of various l<inds (e.g. nitrates and nitrites, 
phosphates, dissolved organic matter], per- 
sistent organic pollutants IPOPsl, including a 
range of chlorinated hydrocarbons, and heavy 
metals such as cadmium (Cd), copper (Cu|, 
mercury IHgl, lead (Pbl, nickel (Nil and zinc 
(Zn|. Quantifying these inputs and assessing 
their impact is problematic, particularly 
because many occur naturally in sea water 

Many POPs and heavy metals can act as 
toxins above certain concentrations, inducing 
mortality or morbidity or impairing repro- 
ductive success, particularly in cases where 
they become increasingly concentrated to- 
wards the top of food chains. Their overall 
impact on marine ecosystems remains un- 
certain. More easily observable is the impact 




of eutrophication resulting from the increased 
input of organic and inorganic nutrients 
(particularly nitrogen and phosphorus) into 
coastal waters, mainly through fertilizer 
runoff and sewage disposal. It is believed that 
human intervention has increased river inputs 
of nitrogen and phosphorus worldwide into 
coastal areas by more than fourfold over 
background levels. These inputs lead to 
increases in productivity in coastal waters, 
often in the form of algal blooms. These 
blooms may themselves be noxious; they also 
typically cause the euphotic zone to reduce 
in vertical extent and are implicated in the 



development of hypoxic (low dissolved oxygen 
concentration] and anoxic (zero dissolved 
oxygen] zones. A shallowing of the euphotic 
zone may cause die-off of photosynthesizing 
benthic algae in shallow-water areas. This 
has occurred, for example, in the Black Sea 
where the euphotic zone had decreased from 
50-60 m vertical extent in the early 1960s to 
around 35 m by 1990, leading to a decrease of 
up to 95 percent in living biomass of benthic 
macrophytic algae such as Phyllophora, 
formerly an important harvested resource. 

Hypoxia and anoxia result from the 
activities of oxygen-respiring bacteria below 
the euphotic zone feeding on accumulated 
dead algae and other organisms and waste 
matter raining down from above. Hypoxia re- 
sults in the emigration of mobile aerobic 
species and mortality of sedentary ones. This 
may have catastrophic impact on local 
fisheries. Most hypoxic zones vary in extent 
through the year and from year to year and 
some are only seasonal, disappearing when 
winter mixing causes re-oxygenation of 
bottom waters. They may be very extensive - 
the hypoxic zone to the west of the Mississippi 
delta covered some 16 000 km' in 1997, having 
covered some 9 000 km' in 1989. Over 50 such 
zones have been identified worldwide to date; 
some appear to be at least in part induced by 
natural phenomena while others are believed 
entirely anthropogenic. 

Invasions of exotic species 

As on land, the breakdown of biogeographic 
barriers in the sea appears to be having a 
major, and increasing, impact on marine 
ecosystems. The chief source is the deliberate 
or accidental translocation of organisms, but 
the construction of marine corridors between 
previously isolated areas has had a major 
impact on geographically restricted areas. 
These factors have resulted in a rapid spread 
of alien species in all the worlds oceans. 

Translocation can result from deliberate 
introduction of harvestable species, acciden- 
tal escapes from aquacuUure and aquarium 
operations, transport in ballast water of ships 
and release of fouling organisms that adhere 
to the hulls of ships and boats. The extent of 
introductions of this kind has yet to be fully 



Marine biodiversity 153 



assessed but is certainly large. In many cases 
(see Box 6.21 the introduced species appear to 
be having a major impact on native biota, 
although In general it is difficult to separate 
the effects of a particular species from 
general ecosystem deterioration. 

Deliberate introductions include the plant- 
ing of mangroves and Nips palms along 
coastlines, and the cultivation of fish, crus- 
taceans, mollusks and algae In many coastal 
regions. Atlantic salmon that have escaped 
from aquaculture are reportedly affecting vs/ild 
stocks In the northeast Pacific, and their 
pathogens have themselves moved into the 
wild populations of closely related species. 
Ballast water is commonly pumped Into the 
hold of ships as a means of controlling balance 
and position In the water, and is liable to be 
flushed out far distant from where It was taken 
on. It has been estimated that on any one day 
the ballast water of the world's ocean fleets 
contains around 10 000 different species". 

The principal artificial corridors are [he 
Suez Canal lopened In 1869) and the Panama 
Canal lopened In 19U1. The 165-km long Suez 
Canal Is a continuous seawater channel with 
a water level at the Red Sea end some 1.2 m 
higher than at the Mediterranean end, leading 
to a constant northward flow of water It Is 
estimated that to date some AOO-500 marine 
species have migrated through the canal (so- 
called Lessepsian migrants'", after Ferdinand 
de Lesseps who planned the canall and 
established themselves In the Mediterranean, 
while a far smaller number have moved in the 
other direction. New species are believed to 
arrive in the Mediterranean at the rate of four 
to five annually. Because the Panama Canal 
has a separate freshwater section, some 25 m 
above sea level, migration through It has been 
limited to date. 

Global climate change 

Human-induced climate change is liable to 
Impact directly on marine and coastal areas 
by warming (particularly of the surface 
layers], by sea-level rise (associated both with 
thermal expansion and the melting of terres- 
trial Ice caps and glaciers!, and through 
change In the gases dissolved in surface 
waters. These impacts are well understood. 



and measurable changes are already appar- 
ent. A more complex array of secondary 
changes may also occur, including changes 
to ocean stratification and surface mixing, 
changes to patterns of surface current, and 
perhaps to global systems such as the El Nino 
Southern Oscillation. 

Tropical coral reefs appear particularly 
sensitive to temperature change. Reef- 
building corals are adapted to stable thermal 
conditions and In most areas appear to be 
growing close to their upper temperature 
limit. Temperatures tittle more than 1°C 
above the normal maximum for a period of 
a few weeks are sufficient to drive a stress 
response known as coral bleaching. During 
a particularly strong El Nino event in 1998, 
warmer waters around the Seychelles and the 
Maldives Induced a bleaching event In which 
60-90 percent of all corals in the area died, 
equivalent to 5 percent of the world coral reef 
area. Although this event was linked to an 
extreme climatic perturbation, it occurred 
at a time of rising global temperatures and 
provides an indication of the impact of 
potential future climate change'". More subtle 
changes associated with the gradually 
changing background conditions, particularly 
temperatures, have been recorded in the 



Box 6.2 Marine introductions 



Leidy's comb jelly Menopsis leidyi was introduced from the American Atlantic into the waters 
of the Black Sea in 19B2, presumably In ballast water Unchallenged by natural predators, 
this species proliferated to a 1988 peak estimated at around 1 000 million metric tons wet 
weight (about 95 percent of the entire wet weight biomass in the Black Seal. The species 
depleted the natural zooplankton stocks, with subsequent algal blooms and decline of the 
fishing industry in the Black Sea". 

. The Asian clam Potamocorbula amurensis spread through the northern San Francisco 
' estuary (United StatesI following its introduction, possibly in ballast water, in 1986. The 
species reaches high density, up to 2 000 Individuals per m', and has caused sharp declines 
In the abundance and extent of several plankton species; its impact on fisheries is not yet 
clear With more than 200 Introduced species, this bay may be the most Invaded aquatic 
habitat In North America. 

The green alga Cauterpa taxifolia is thought to have escaped from an aquanum In the 
western Mediterranean and is spreading rapidly In the coastal waters of Spain, France and 
Italy, with severe Impact on the native seagrass beds and on coastal fisheries". It has 
recently been reported In the coastal waters of California^'. 



154 WORLD ATLAS OF BIODIVERSITY 



*K< 



Much less readily observed, 
marine species are in 
general more difficult to 
monitor and assess tfian 
terrestrial ones. 



distribution of pelagic seabirds along the 
Californian coast"; the faunal composition of 
intertidal communities'"; and penguin distri- 
bution in the Antarctic". 

The mean sea level has risen 18 cm during 
the past 100 years and further increases 
could have a severe impact on coastal 
communities. Rising sea levels will lead to 
the inundation of some coastal lands, whilst 
in many other areas they will alter patterns 
of coastal erosion, and they may increase 
groundwater salinization. While many inter- 
tidal habitats are highly adaptable, the 
growing human presence in most coastal 
areas will prevent the natural migration of 
these habitats, leading to overall losses of 
saltmarsh, mangrove, or even beach and 
rocky shore habitats. 

The biomass of the world's oceans is very 
low compared with terrestrial environments, 
but because of the rapid turnover in oceanic 
carbon cycles, marine phytoplankton Icyano- 
bacteria and algael play an important role 
in removing dissolved carbon dioxide from 
solution, and are intermediates in the trans- 
port of organic carbon to the deep ocean. 
Once in the deep ocean, this carbon is effec- 
tively removed from exchange with the 
atmosphere tor millennia. In this way, marine 
photosynthesizers help to buffer the rising 
concentration of carbon dioxide in the 
atmosphere, but this service, and marine 
primary productivity, may be affected if ocean 




temperatures rise (warmer waters hold 
less carbon dioxide in solution] and if the 
broad patterns of ocean circulation change 
significantly". It is thought that increasing 
atmospheric temperatures may affect the 
generation of cold, oxygen-nch bottom waters 
beneath Arctic and Antarctic ice sheets, with 
major implications for deep-sea biota and 
for global patterns of seawater circulation, in 
particular the Great Conveyor, driven by 
bottom water generated in the North Atlantic. 

THE CURRENT STATUS OF MARINE 
BIODIVERSITY 

Because they are usually much less readily 
observed, marine species are in general 
much more difficult to monitor and assess 
than terrestrial ones. Assessment is based 
on sampling and, in the case of harvested 
species, often on the basis of catch rates, 
although the latter may vary in response to 
a wide range of factors in addition to changes 
in the population of the species concerned. 
An exception lies with those groups such as 
pinnipeds, sea turtles and seabirds that nest or 
breed on land. Because many of these tend to be 
colonial species and because they tend to breed 
in open habitats (beaches, cliff tops, ice sheets), 
they may be easier to monitor than many other 
species, either terrestrial or aquatic. In the case 
of large, commercially valuable fish stocks, 
monitoring at large scale has in some cases 
been carried out tor many years, so that 
estimates of the stock level are obtainable. 

Threatened and extinct species 

The only major marine species groups 
(classes or above) that have been com- 
prehensively assessed in terms of threatened 
species status to date are mammals and 
birds. In addition, sea turtles and a number of 
fish families and genera (e.g. the sturgeons 
in the order Acipenseritormes and the sea- 
horses Hippocampus in the order Syngnathi- 
formes) have also been assessed. Other 
threatened marine species have been identi- 
fied on more of an ad hoc basis. Data are 
summarized in Table 6.12. 

Relatively speaking, far fewer marine 
species are known to have become extinct 
since 1600 than either terrestrial or fresh- 



Marine biodiversity 155 



^'h 



water ones. Cataloged extinctions comprise 
two marine mammals (the Caribbean monk 
seal Monachus tropicaiis and Steller's sea 
cow Hydrodamalis gigas] and five seabirds 
(three island petrels, Pallas's cormorant 
Phalacrocorax perspicillatus and the great 
auk Alca impennis]. In addition five coastal 
or island duck species have disappeared at 
various times from the late 17th century 
onwards; however there is In most cases 
insufficient information to determine whether 
these species were predominantly marine 
or terrestrial. 

As a gross generalization, marine species 
appear to be somewhat less extinction prone 
as a result of mankind's activities than fresh- 
water or terrestrial ones. There are arguably 
two main reasons for this. First, because of 
the size of the world ocean and the fact that 
people do not actually live In it, the marine 
biosphere remains as a whole considerably 
more buffered from human Intervention than 
terrestrial and inland water areas. Second, 
marine species on the whole appear to be 
more widespread than terrestrial or inland 
water ones. In the open ocean, there are vast 
areas with apparently similar habitat con- 
ditions and there are few barriers to dispersal 
so that many species have circumglobal 
distributions. In addition, many forms that as 
adults are sessile (e.g. sponges and coralsl or 
sedentary (many mollusks and crustaceans! 
have planktonic larvae that are often widely 
dispersed In water currents. For this reason, 
many coral reef species, for example, are 
found in suitable habitat throughout the Indo- 
Paclfic region. In addition, many of the most 
heavily exploited fish species have high 
fecundity (In the case of some tunas amoun- 
ting to several million eggs In a single 
spawning), so that they have at least poten- 
tially high population growth rates unparallel- 
ed in terrestrial vertebrates. 

There are of course significant exceptions to 
all these. Coastal regions In many parts of the 
world, and enclosed or semi-enclosed marine 
areas such as the Baltic, Black and Yellow 
Seas, are often under intense pressure from a 
range of human activities. A number of marine 
species do appear to have restricted ranges 
(e.g. the Hawaiian coral reefs are relatively rich 



in species found nowhere else while many 
southern hemisphere seabirds are apparently 
confined to a small number of breeding sites) 
and significant numbers have low or very low 
reproductive rates (many chondrichthyme 
fishes, marine mammals and seabirds). 



Marine species which are 
easily exploitable or have 
high economic value may 
suffer catastrophic declines 
if exploitation is not strictly 
controlled. 




Until recently by far the most important 
human activity affecting marine species was 
uncontrolled exploitation. Where species are 
either easily exploitable or are highly sought- 
after [i.e. have high unit value], or both, they 
may suffer catastrophic declines. This Is the 
case with sea turtles and a number of marine 
mammals and birds that are or have been 
harvested principally at their terrestrial 
breeding sites (which are often colonial), as 
well as with the great whales and the dugong, 
which although strictly marine are air- 
breathing and therefore spend some time at 
the sea surface (when they may be spotted). 
Most of these species have relatively low 
reproductive rates, so that even if they are 
ultimately afforded protection population 
recovery rates may be very slow. 

A recent synthesis of a wide range of 
information, including paleoecologlcal and 
archeologlcal data relating to early human 



156 WORLD ATLAS OF BIODIVERSITY 



Table 6.12 

Taxonomic distribution 
and status of threatened 
marine animals 

Note: Only the birds and 
mammals have been 
comprehensively assessed for 
species at risk; numbers refer 
to species (units such as 
subspecies and geographic 
populations appear in the Red 
List database but these are 
not tabulated here). 



Source: Status categories from 2000 
Red List database, www.redlist.org 
(accessed February 20021. 



communities, suggests that overfishmg of 
larger vertebrates and mollusks is charac- 
teristic of indigenous and colonial human 
use of coastal ecosystems, and the first of 
what is typically a series of impacts'-, 
tvlassive losses in biomass and abundance 
appear to have occurred, on a scale largely 
unsuspected, and seemingly amounting to 
the loss of entire trophic levels of consumer 
organisms, with radical consequences for 
ecosystem status. Overfishing is likely to be 
followed by impacts of pollution, mechanical 
habitat loss, introduced species and climate 



change. Loss of filter-feeding organisms 
that maintain water quality is liable to be 
followed by eutrophication, hypoxia and 
disease, as exemplified by conditions in 
Chesapeake Bay following the collapse of 
the oyster fishery in the early 20th century. 
Synergistic interactions of this kind are 
making the effective, long-term manage- 
ment of marine resources one of the major - 
and most intractable - problems currently 
facing humankind. 

Land-breeding species may also be sus- 
ceptible to other threats, such as predation, 



Phylum ^^^^^^^^1 


F ' Common name 'Critically 


'Endangered' 


'Vulnerable' 


_ class or order ^^^^^^H 


|^^_ endangered' 




^ 


Cnidaria 








Anthozoa 


Stony corals 




2 


Mollusca 








Bivalvia 


Bivalves 




4 


Gastropoda 


Gastropods 1 


2 


6 


Craniata- fishes 








Carcharhiniformes 


Ground sharl<s 1 


3 


5 


Hexanchiformes 


Cow/ sharks 




1 


Lamniformes 


Mackerel sharks 




3 


Orectolobiformes 


Carpet sharks 




2 


Pristiformes 


Saw/fishes 2 


5 




Rajiformes 


Rays 


2 


1 


Rhinobatiformes 


Giiitar fishes 1 




1 


Squaliformes 


Dogfishes and 
sleeper sharks 




1 


Squatiniformes 


Angel sharks 


1 


2 


Coelacanthiformes 


Coelacanths 1 






Acipensenformes 


Sturgeons 2 


10 


6 


Batrachoidiformes 


Toadfishes 




5 


Clupeiformes 


Herrings and anchovies 


1 


1 


Gadiformes 


Cods, hakes, rattails 1 




2 


Gasterosteiformes 


Sticklebacks 




1 


Lophiiformes 


Anglerfishes, etc. 1 






Ophidiiformes 


Pearlfishes, cusk-eels, bnatulas 




1 


Perciformes 


Perches, etc. 7 


5 


33 


Pleuronectiformes 


Plaice, flounders, soles 


1 


1 


Salmoniformes 


Salmonids 1 


i 


6 


Scorpaeniformes 


Gurnards, scorpionfishes, etc. 1 


2 


1 


Siluriformes 


Catfishes 


1 




Syngnathiformes 


Pipefishes, seahorses, etc. 1 


1 


35 


Tetraodontiformes 


Triggerfishes, etc. 




3 



Marine biodiversity 157 



coastal development and pollution. It is note- 
worthy in this context that the family 
Procellariidae contains nearly three times as 
many threatened species (36 out of 115 
species, or 28 percent! as the average bird 
family, in Vi^hich 11 percent of species are 
threatened, and nearly six times as many 
critically endangered species as would be 
expected at random. It is almost certainly the 
tendency of these birds to nest on islands, 
whose biotas have in general suffered enor- 
mously more from mankind's influence in the 
past few centuries (see Chapter A], rather 



than their seagoing habits that has led 
to this. 

For truly marine species (chiefly finfishes 
and invertebrate animals! the situation appears 
somewhat different. Even when these have 
been exploited to the point of stock collapse, 
as has occurred for example with the cod 
Gadus morhua stocks off Newfoundland in the 
North Atlantic, the species concerned do not 
appear to have become imminently threatened 
with biological extinction. This is in part 
because once stocks are reduced below a 
certain level it is often no longer economically 



phylum and 


Family 


Common name 


'Critically 


'Endangered' 


'Vulnerable' 


■ class or order ^^^ 




endangered' 




^ 


Cranlata - Reptilla 












Squamata 


Iguanidae 


Iguanas 






1 


Chelcnia 


Dermochelyidae 


Leathery turtle 










Cheloniidae 


Sea turtles 


2 


3 


1 


Craniata - Aves 












Anseriformes 


Anatldae 


Ducks 




1 


1 


Cfiaradriiformes 


Alcldae 


Auks, puffins 






4 


1^ 


Charadriidae 
Laridae 


Plovers 

Gulls, terns, skuas, 
auks, skimmers 




2 
1 


1 

5 


Ciconiiformes 


Ardeidae 


Egrets, herons 






1 


Pelecaniformes 


Fregatidae 
Pelecanidae 


Frigateblrds 
Pelicans 






1 
1 




Phalacrocoracldae 


Cormorants and shags 






8 




Sulidae 


Gannetsand boobies 






1 


Procellarilformes 


DIomedeldae 
Hydrobatldae 


Albatross 
Strom petrels 


2 


2 


12 
1 


• 


Pelecanoidldae 


Diving petrel 




1 




Procellariidae 


Petrels, shearwaters 


10 


6 


20 


Sptienisciformes 


Spheniscidae 


Penguins 




3 


7 


Craniata - Mammalia 












Carnivora 


Muslelidae 


Otters, etc. 




2 






Otarlidae 


Eared seals 




1 


5 


Cetacea 


Phocidae 
Balaenidae 


Earless seals 
Right whales 


1 


1 
1 


1 


■§ 


Balaenopteridae 


Rorquals 




3 


1 


^k 


Delphlnidae 


Dolphins 




1 


1 


^K 


(i^onodontidae 


Beluga 






1 


■ 


Phocoenidae 
Physeteridae 


Porpoises 
Sperm whales 


1 




1 
1 


Sirenia 


Dugongidae 


Dugong 






1 




Trichechidae 


Manatees 






3 


l^ 













158 WORLD ATLAS OF BIODIVERSITY 



*i<m 



UO 



120 



100 



60 



W 



1970 



1975 



viable to continue harvesting them. Generally, 
the residual population at this stage is still 
large enough to allow recovery if harvesting 
ceases, particularly in the case of species with 
high fecundity and therefore high potential 
intrinsic rates of increase. Exceptions to this 
are species that have low fecundity, partic- 
ularly if they also have a long period to 
maturity, with limited ranges, and which may 
either have high unit value or be caught as 
bycatches. 

In the case of bycatches, because the 
fishery is not directed at the species 
concerned, its intensity will not decrease as 
population levels decrease so that it may 
theoretically be possible at least locally to 
extirpate species, particularly if they are 
habitually caught before they reach maturity. 
Examples include several sawfish species 



1980 



1985 



1990 



1995 



1999 



Figure 6.6 

Marine population trends 

Note: A simplified 
representation of the average 
population change in a 
sample of 217 marine 
species, see text. 



(family PrIstidaeL These are large, slow- 
growing, predominantly inshore species that 
give birth to relatively small numbers of live 
young. Population densities appear to be 
naturally low and animals are widely caught 
as bycatch in inshore fisheries before they 
are large enough to reproduce. As a result 
five species are classified as 'endangered' and 
two as 'critically endangered'. In addition, it is 
possible that trophic shifts may occur when 
populations of some species are severely 
reduced, inhibiting recovery of these popu- 
lations when exploitation ceases. This has 
been suggested in the case of some great 



whale populations that have not apparently 
recovered as rapidly as projected following the 
cessation of their harvest. 

The marine living planet Index 

An impression of the overall trend in a large 
sample of species for which population 
indicators are available can be derived from 
the WWF living planet index"'. This approach 
IS designed to represent the change in the 
average species' in the sample from one five- 
year interval to the next, starting in 1970. 
The marine sample represents 217 aquatic 
and coastal species of mammals, birds, 
reptiles and fishes, and the overall trend is for 
a significant decline In population levels over 
the last three decades of the 20th century 
(Figure 6.61. The sample is dominated by the 
stocks and species that humans have an 
interest in monitoring, most of the fishes 
among these being of commercial importance 
as a fisheries resource. These should also 
be stocks that humans have an Interest In 
managing as well as possible. That the index 
has declined In every five-year interval since 
1970 is evidence that such management Is 
failing, as confirmed by the picture painted 
above of global marine capture fisheries. 

Assessing the status of marine and coastal 
ecosystems 

Threatened species inventories and the marine 
living planet index can give a very general 
overall Impression of the status of marine 
biodiversity. Assessing marine ecosystem 
health' is much more problematic. However, 
snapshots can be obtained from examining 
particular ecosystems, such as mangroves 
and coral reefs". In the former, an overall 
assessment can be made on the basis of the 
area destroyed or severely degraded. In the 
latter areal measures are more problematic. In 
part because reef extent is much more difficult 
to measure than mangrove extent and, of 
greater importance, because the vast majority 
of a reef Is composed of non-living calcareous 
deposits. Measures of the change in extent of 
these give little insight Into the state of the 
living component of the reef. For this reason 
other measures, such as estimates of inci- 
dence of coral disease, may be feasible. 



Marine biodiversity 159 



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60 



61 



162 WORLD ATLAS OF BIODIVERSITY 



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2002). 



Inland water biodiversity 163 



/ Inland water bi 



odiversity 



INLAND WATERS h4AKE UP A MINUTE PROPORTION Imuch less than a hundredth of 
1 percent! of the world's water resource. Despite this, they encompass a wide range 
of habitat types and contain a disproportionately high fraction of the world's biodiversity. 
Freshwater is also a vital resource for human survival. In consequence, Inland water 
ecosystems are placed under many, often conflicting, pressures, with Increasingly adverse 
consequences for their biodiversity. There are indications that, overall, a higher proportion 
of inland water species are In decline than marine or terrestrial forms. 



INLAND WATERS 

The hydrosphere is estimated to contain about 
1 386 million cubic kilometers Ikml of water, 
almost all of which (97.5 percent) is saline 
water making up the world ocean, leaving 
some 35 million km' 12.5 percent) of fresh- 
water. A major proportion (about 69 percent] 
of this freshwater is locked up in the form of 
ice and permanent snow, where it is unavail- 
able to living organisms. The Earths liquid 
freshwater is mostly in subterranean ground- 
waters, with a small proportion in soils and 
wetlands, and the smallest proportion of all - 
about 0.01 million km^ or 0.3 percent of all 
freshwater - makes up the world's lakes and 
rivers on which inland water biodiversity 
depends (Table 7.1). 

There are very large regional differences in 
the concentrations of water in all its forms 
(e.g. about twice as much atmospheric water 
in equatorial as in temperate latitudes], and in 
the occurrence of different types of inland 
waters (e.g. South America has an enormous 
concentration of river water but few large 
lakes, while the converse is true for Africa). 
Over the oceans, evaporation exceeds input 
from rivers and rainfall, and over land precipi- 
tation exceeds evaporation. This excess on 
land amounts to about ^3 000 km^ annually 
and this represents the global runoff that 
replenishes the world's rivers, lakes and 
marine waters, and which humans draw on. 



together with groundwater, to meet their 
domestic, agricultural and industrial needs. 

Chapter 5, on terrestrial ecosystems, 
outlines some of the principal differences 
between terrestrial and aquatic environments 
as they determine the living conditions of 
organisms in them, (n general, freshwater sys- 
tems and the organisms within them are far 
more strongly affected by daily and seasonal 
changes as a result of weather patterns and 
climate conditions than are marine aquatic 
environments, in parallel with change in the 
broader terrestrial environment. For example, 
upland streams may receive an influx of cold 
meltwater in spring, and be subject to strong 
insolation during midsummer; the volume, 
speed and transparency of river water is 
liable to change radically following rainfall in 



Table 7.1 

Components of the 
hydrosphere 

Source; Anon . Shiklomanov 





Area 


% total 


Volume 


% total 


1 
% 


m^l^ 


Imilllon km') 


area 


(million km^l 


water 


freshwater 


Earth's surface 


510 










Land 


U9 


29 








World ocean 


361 


71 


1351 


97.5 


- 


Freshwater 


_ 


_ 


34.65 


2.5 


100 


Ice and permanent 


snow 16 


- 


23.8 


1.76 


68.9 


Groundwater 


- 


- 


10.4 


0.77 


29.9 


Wetlands, soil viiater 


permafrost 2.6 


- 


0.3 


0.02 


0,9 


Lakes and rivers 


1.5 


- 


0,01 


0.0007 


0.3 



164 WORLD ATLAS OF BIODIVERSITY 



Box 7.1 Saline and soda lakes 



Some coastal or high altitude lakes and lagoons combine high concentrations of dissolved 
chemicals with high water temperatures, sometimes as high as 70°C. Few kinds of species 
thrive in these harsh alkaline or saline conditions. Communities typically include 
cyanobacteria, diatoms, a few small invertebrate animals such as brine shrimps [Artemia], 
with flamingos often being the only large organisms present. Flamingos, of which five living 
species are recognized, are filter-feeders, with the bill and mouthparts specialized to extract 
small particles from water All share the same basic morphology, but tend to extract prey of 
different sizes: e.g. in sub-Saharan Africa the lesser flamingo feeds almost exclusively on 
cyanobacteria and other microorganisms, while the greater flamingo (often found in the 
same lakes), feeds mainly on small macroscopic invertebrates, such as brine shrimps and 
other crustaceans. Fishes are often absent from lakes where flamingos are abundant, and 
flamingos are rarely abundant where fishes are common. This may be because the two 
compete with each other for their food supply but, in the inhospitable lakes where they 
abound, flamingos have no real competitors. Their early adaptation to an extreme 
environment that no other large animal was capable of exploiting may explain why they have 
survived relatively unchanged in morphology for so long (fossils are known from the Eocene, 
about 50 million years before the present). 



ttie catctiment basin. The soil and surface 
geology of watershed areas have a strong 
influence on the chemical composition of both 
river and lake waters and, for example, buffer 
or reinforce the effect of acid rain. 

Inland water habitats 

Although the terms 'inland water' and 
'freshwater' are often used more or less 
interchangeably they are not equivalent. 
A considerable number of inland waters are 
saline, some much more so than sea water. 
Conversely, waters of the deltaic regions of 
some major river systems (most notably the 
Amazon] may be fresh a considerable dis- 
tance out to sea. 

Despite their vastly smaller extent, inland 
aquatic habitats show far more variety in their 
physical and chemical characteristics than 
marine habitats. They encompass systems as 
varied as the world's great lakes and rivers, 
small streams and ponds, temporary puddles, 
thermal springs and even the minute pools 
of water that collect in the leaf axils of certain 
plants, such as bromeliads. Chemically they 
range from almost pure water to highly 
concentrated solutions of mineral salts, 
toxic to all but a few specialized organisms 
(see Box 7.1). 



Inland water habitats can be divided into 
running or lotic and standing or lentic sys- 
tems. They may also be divided into per- 
manent water bodies, periodically (usually 
seasonally) inundated, and ephemeral or 
transient. Each of these has its own distinct 
set of ecological characteristics. 

There is not necessarily a rigid dividing line 
between an inland aquatic habitat on the one 
hand and a terrestrial or marine habitat on 
the other. Any temporarily inundated area, 
such as a river floodplain, is effectively a 
hybrid or transitional system, being at some 
times essentially aquatic, at other times 
terrestrial. Similarly there are many areas 
that consist of shifting mosaics of land and 
shallow water, or areas of saturated vege- 
tation, such as sphagnum moss bogs that are 
strictly neither land nor water. These trans- 
itional areas are often collectively termed 
'wetlands'. Similarly, estuarine areas are 
transitional areas between inland and marine 
systems. 

Lotic systems: rivers and catchment basins 
A river system is a complex but essentially 
linear body of water draining under the 
influence of gravity from elevated areas of 
land toward sea level. The typical drainage 
system consists of a large number of smaller 
channels (streams, rills, etc.) at higher ele- 
vation merging as altitude falls into pro- 
gressively fewer but larger channels, which in 
simplest form discharge by a single large 
watercourse. Most such systems discharge 
into the coastal marine environment. Some 
discharge into lakes within enclosed inland 
basins; a few watercourses in arid regions 
enter inland basins where no permanent 
lake exists. 

The source area of all the water passing 
through any given point in the drainage 
system is the catchment area for that part of 
the system. In parallel with the hierarchical 
aggregation of tributaries of the major river 
system, sub-catchments aggregate into a 
single major catchment basin; this is the 
entire area from which all water at the final 
discharge point of the system - i.e. usually 
the sea - is derived. Strictly, the watershed 
is the line of higher elevation dividing one 



Inland water biodiversity 165 



catchment basin from another, but this 
term is increasingly used as a synonym 
of catchment. 

The speed and internal motion of river 
water depends largely on water volume and 
the shape of its channel. These factors typi- 
cally differ greatly through the river system, 
from narrow, steep and fast upland feeder 
streams to broad, level and slow downstream 
reaches. In virtually all river systems water 
volume also varies seasonally. Some rivers in 
arid or semi-arid catchments flow for only 
part of the year, or in extreme cases only once 
every several years. 

Large rivers may span many degrees of 
latitude and pass through a wide range of 
climatic conditions within their catchments. 
Variations in water flow and underlying geo- 
logy also create a wide range of habitats 
within any river and often within a short 
distance. Different organisms are typically 
adapted to different parts of any given 
river system. 

River systems can change course radically 
as a result of deposition and erosion of their 
channel, and the uplift and erosion of 
watershed uplands. Despite the dynamic 
physical state of these systems, large rivers 
rarely disappear, and although direct evi- 
dence is scarce, indications are that some 
have been in continuous existence for tens of 
millions of years. This is consistent with the 
fact that running waters include represen- 
tatives of almost all taxonomic groups found 
in freshwaters, and that several invertebrate 



taxa occur only in running waters or attain 
greatest diversity there. By far the largest 
river catchment in the world is the Amazon 
(with the Ucayalil in South America which 
covers just under 6 million km' and is nearly 
60 percent larger than the next largest, the 
Congo in central Africa. Unsurprisingly, the 
former is the major repository of the world's 
freshwater biodiversity. Between them the 
20 largest river catchments cover around 
A5 million km', or about one third of the 
world's ice-free land surface'. 

Lentic systems 

Lakes and ponds 

The great majority of existing lakes, of which 
around 10 000 exceed 1 km' in extent, were 
formed as a result of glacial activity, with most 
of the rest a result of tectonic activity'. Tectonic 
lakes are formed either as a result of faults 
caused by deep crustal movements or by 
volcanism. In the case of the former, a lake 
may form in a depression caused by a single 
fault, or in a depressed area between two or 
more faults - these being graben lakes - or in 
a rift valley Most volcanic lakes form in craters 
or calderas of volcanoes while a few (usually 
short-livedl may form behind dams caused by 
lava flows. Glacial lakes occupy basins caused 
by the scouring action of ice masses. 

Most of the world's existing lakes are 
glacial and geologically very young, dating 
from the retreat of continental ice sheets at the 
start of the Holocene. around 11 500 years 
before present. All such lakes are expected to 




The majority of lakes 
were formed as a result 
of glacial activity. 



u« WORLD ATLAS OF BIODIVERSITY 



Table 7.2 

Physical and biodiversity 
features of major long- 
lived lakes 

Notes: Lakes ordered by 
volume. A few other lakes 
have notable endemism 
among fishes, mollusks, 
crustaceans or other groups. 
Among these are lakes Inle 
IMyanmarl. Lanao 
IPhilippmesI, Malili 
(Indonesia] and the Cuatro 
Cienegas basin (Mexico] - but 
their ages are not yet firmly 
established. Qualitative 
remarks (e.g. very high', 
Tow'] in the Biodiversity 
column are related to long- 
lived lakes, not to lake 
systems in general. All 
biodiversity data are 
approximate and subject to 
change with new survey data 
or different taxonomic 
opinion, 

i Evidence indicates that the 
lake dried out completely, or 
nearly so. around the late 
Pleistocene, 10-12 000 years 
ago". 

Source: Collated from data in Martens 
et at. . Fish estimates for East African 
Lakes from Snoeks°^. 



Lake 


Country Age 


Max. 


Vol. 


^^^^^^^^^M 




(million 


deptit 


(km'l 


^^^^^^^^M 




years) 


(ml 




'^V^^H 


Baikal 


Russia 25-30 


1637 


23 000 


Very high sp, richness; exceptional 


Largest, deepest. 








endemism in fishes and several 


oldest extant freshwater 








invertebrate groups 


lake (20% of all liquid fresh 








Total animal spp.: 1 825, endemic: 982 


surface water on Earth] 








Fishes: 56 spp., 27 endemic 


Tanganyika 


Burundi 20 
Tanzania 
Zambia 
DR Congo 
(former Zaire] 


1470 


18 880 


Very high sp. richness; high 
endemism, especially high among 
cichlid fishes 

Total animal spp.: 1 470. endemic: 632 
Fishes: 325 spp., including 250 cichlids 
of which 98% endemic 


Malawi 


Ivlalawi >2 

Iwlozambique 

Tanzania 


780 


8 400 


Very high sp. richness; high 
endemism, especially high among 
cichlid fishes 

Fishes: 845 spp., including 800 cichlids 
of which 99% endemic 


Victoria 


Kenya >4?' 


70 


2 7i0 


High sp, richness, especially of 


World's second largest 


Tanzania 






fishes; exceptional endemism 


freshwater lake (area] 


Uganda 






among cichlid fishes -many fish 
endemics depleted or extirpated 
following introduction of Nile perch 
Fishes: 545 spp,, including 500 cichlids 
of which 99% endemic 


Titicaca 










One of world's highest 


Bolivia 3 


280 


890 


Moderate sp. richness and endemism 


altitude lakes 


Peru 






(highest among fishes] 

Total animal spp,: 533, endemic: 61 

Fishes: 29 spp., 23 endemic 


Biwa 


Japan 4 


104 


674 


Moderate sp. richness and 
endemism (highest in gastropod 
mollusks and fishes] 

Total animal spp.: 595, endemic: 54 ! 
Fishes: 57 spp., 4 endemic i 


Olirid 


Albania 3 


295 


50 


Moderate sp, richness; 5 


Fed mainly by 


Macedonia 






exceptional endemism in several ' 


subterranean karst 


IFYR] 






groups (planarians, oligochaetes, '■ 


waters 








gastropod mollusks, ostracod 

crustaceans] 

Fishes: 17 spp., up to 10 endemic 



Inland water biodiversity 167 



fill slowly with sediment and plant biomass. 
and to disappear within perhaps the next 
100 000 years, along with any isolated biota. 
Lakes may also be caused by the dissolution of 
soluble rocks, most notably limestone in karst 
regions which is gradually dissolved by dilute 
acids in water running through it, and by 
changes in the course of rivers in floodplain 
regions, which result in ox-bow and scroll 
lakes. 

Only about ten existing lakes are known 
with certainty to have origins much before the 
Holocene (Table 7.2P, and most of these 
occupy basins formed by large-scale sub- 
sidence of the Earths crust, dating back to 
at most 20 million (Lake Tanganyikal or 30 
million (Lake Baikal) years before the present. 

There is good evidence that some extinct 
lake systems in the geological past were very 
large and very long-lived under different 
climatic and tectonic conditions. In general, 
the long-lived lakes are of particular interest 
in terms of biodiversity because they tend to 
be rich in species of several major groups of 
animals and many of these species are 
restricted to a single lake basin. 

Wetlands 

As indicated above, the distinction between a 
wetland, an aquatic system and a terrestrial 
system may be essentially arbitrary. However, 
a number of mixed shallow-water and terres- 
trial habitat types share several characteristics 
and are habitually grouped as wetlands. 
Wetlands in this sense are typically hetero- 
geneous habitats of permanent or seasonal 
shallow water dominated by large aquatic 
plants and broken into diverse microhabitats\ 
The four major broad habitat types are: 

Bogs 

Bogs are peat-producing wetlands in moist 
climates where organic matter has accum- 
ulated over long periods. Water and nutrient 
input is entirely through precipitation. Bogs 
are typically acid and deficient in nutrients 
and are often dominated by sphagnum moss. 

Fens 

Fens are peat-producing wetlands that are 

influenced by soil nutrients flowing through 



the system and that are typically supplied 
by mineral-rich groundwater Grasses and 
sedges, with mosses, are the dominant vege- 
tation. Fens are typically more productive and 
less acidic than bogs. 

Marshes 

Marshes are inundated areas with herb- 
aceous emergent vegetation, commonly dom- 
inated by grasses, sedges or reeds. They may 
be either permanent or seasonal and are fed 
by ground or river water, or both. 

Swamps 

Swamps are forested freshwater wetlands on 
waterlogged or inundated soils where little or 
no peat accumulation occurs. As with marshes, 
they may be either permanent or seasonal. 

Biogeography and important areas 
Freshwater lineages that originated in contin- 
ental water systems may show general 
patterns of distribution similar to terrestrial 
groups, corresponding more or less to broad 
biogeographic realms. Lineages of marine 
origin may remain restricted to peripheral 
systems corresponding to the area where the 
ancestral forms moved into freshwater 

Unlike many terrestrial species, which can 
disperse widely in suitable habitat, the spatial 
extent of the range of strictly freshwater 
species tends to correspond to present or 
formerly continuous river basins or lakes. 
These species include fishes and most moll- 
usks and crustaceans. Watersheds between 
river basins are the principal barriers to their 
dispersal between systems, and their ranges 
are extended mainly by physical changes to 
the drainage pattern (e.g. river capture 
following erosion or uplift can allow species 
formerly restricted to one system to move into 
another!, or by accidental transport of eggs by 
waterbirds, or by flooding. 

In many instances, the range within a 
system will also be restricted by particular 
habitat requirements (variations in water 
turbulence or speed, shelter, substrate, etc.]. 
These frequently differ at different stages in 
the life cycle (for example in fishes the 
conditions and sites required for egg depos- 
ition and development, for early growth of fry, 



168 WORLD ATLAS OF BIODIVERSITY 



Continent 


Area name ^^^^H 


m 


Taxonomic group 


^^1 


Africa 


L. Malawi 


Fishes 


Mollusks 




Africa 


L Tanganyika 


Fishes 


Mollusks 


Crabs 


Africa 


L.Victoria 


Fishes 


Mollusks 




Africa 


Madagascar 


Fishes 


Mollusks 


Crabs 


Africa 


Niger-Gabon 


Fishes 




Crabs 


Africa 


Upper Guinea 


Fishes 


Mollusks 


Crabs 


Africa 


Lower Congo 


Fishes 


Mollusks 


Crabs 


Australia 


SE Australia & Tasmania 


Fishes 


Mollusks 


Crayfish 


Australia 


SW Australia 


Fishes 




Fairy shrimp 


Eurasia 


SE Asia and lower 










Ivlekong River 


Fishes 


Mollusks 


Crabs 


Eurasia 


Balkans (southwest 1 


Fishes 


Mollusks 




Eurasia 


L. Baikal 


Fishes 


Mollusks 




Eurasia 


L. Biwa 


Fishes 


Mollusks 




Eurasia 


L Inle 


Fishes 


Mollusks 




Eurasia 


L Poso 


Fishes 


Mollusks 




Eurasia 


Malili Lakes 


Fishes 


Mollusks 




Eurasia 


Sri Lanka 


Fishes 




Crabs 


Eurasia 


Western Gtiats 


Fishes 


Mollusks 


Crabs 


Nortfi America 


East Mississippi drainage 
(Ohio, Cumberland, 










Tennessee rivers) 


Fishes 


Mollusks 


Crayfish 


North America 


Mobile Bay drainage 


Fishes 


Mollusks 


Crayfish 


North America 


Western USA 


Fishes 


Mollusks 


Fairy shrimp 


South America 


L. Titicaca 


Fishes 


Mollusks 




South America 


La Plata drainage 


Fishes 


Mollusks 




South America 


Amazon basin 


Fishes 


? 


Crabs 



Table 7.3 

Partial list of global 

hotspots of frestiwater 

biodiversity 

Notes: This table lists areas 
of special importance for 
diversity in fishes and either 
mollusks or crustaceans or 
both. See text and Appendix 
6. Six of the seven long-lived 
lakes in Table 7.2 also 
appear here 

Source: See sources cited in 
Appendix 6, 



and for feeding and breeding of adults are 
often different). 

Many cave or subterranean freshwater 
aquatic species (e.g. of fishes, amphibians 
and crustaceans) have restricted ranges, 
perhaps consisting of a single cave or aquifer, 
and limited opportunities for dispersal, 
depending on the surrounding geology and 
the consequent morphology of the water 
system occupied. 

Analysis of data from some 151 river 
basins indicates that there is a strong cor- 
relation between the spatial extent of a river 
catchment and the number of fish species 
therein. The size' of a river can be represen- 
ted by the area of the basin or by the volume 
of water flowing through the river system in 
any given period; the latter is a better 
predictor of fish species richness than is 
basin area. When area is taken into account. 



there is also a strong relationship between 
species richness and the latitude of the basin. 
Recent analysis suggests that latitude may be 
a surrogate measure for energy availability 
and productivity within the basin' ^ factors 
known to be well correlated with variation in 
terrestrial diversity (Chapter 5). No taxonomic 
class restricted to inland waters has yet been 
mapped globally at species level but, at a 
higher taxonomic level, a density surface of 
freshwater fish families has been developed 
(see Map 7.1] with a view to providing an 
indication of global variation in inland water 
diversity analogous to those available for 
terrestrial groups (Chapter 5). 

A recent analysis of areas important for the 
maintenance of global freshwater biodiversity' 
was based on the expert view of a number of 
regional and taxonomic specialists. The analy- 
sis was designed to make effective use of 
readily available information and. although 
preliminary, yielded the first global overview of 
freshwater biodiversity hotspots'. fvtaps 7.2, 7.3 
and 7.A show, respectively, important areas for 
freshwater fishes, mollusks and selected crus- 
tacean groups. Further details of all these 
areas can be found in Appendix 6. Table 7.3 
lists the sites and areas that have been 
identified as of special importance for more 
than one of the above groups. It is not a 
comprehensive global listing because it omits 
several large but imprecisely defined areas of 
known high diversity, and it omits diverse taxa 
not covered in the assessment (e.g. amphi- 
pods, copepodsl; nor does it mention sites of 
key importance mainly for one group of 
animals. Although the Amazon basin is a vast 
region rather than an identifiable site, it has 
such an exceptional diversity of fishes that it 
could not reasonably be excluded from a list of 
globally important areas. 

More detailed continental reviews are now 
also available for Asia, including discussion of 
taxonomy, hotspots and policy', tor Latin 
America' and North America'. In order to help 
prioritize investment, the conservation organi- 
zation WWF has selected 53 freshwater eco- 
regions, based on a combination of biogeo- 
graphic region, water body type, biodiversity and 
representativeness' ". Perhaps more so than in 
other biomes, these freshwater ecoregions 



Inland water biodiversity 169 



■'^^ 



have a firm objective basis because they 
correspond broadly with catchment basins. 

BIOLOGICAL DIVERSITY IN INLAND WATERS 

At high taxonomic levels the diversity of 
freshwater organisms is considerably lower 
than on land or in the sea. Only one extant 
eukaryote phylum IGamophyta - green conju- 
gating algael is apparently confined to fresh- 
water habitats. The number of species overall 
is low in absolute terms in comparison with 
marine and terrestrial groups, but species 
richness in relation to habitat extent is 
relatively high. For example, about 10 000 (40 
percent! of the 25 000 known fish species are 
freshwater forms'. Given the distribution of 
water on the Earth's surface this is equivalent 
to one fish species for every 15 km^ in fresh- 
waters compared with one for every 100 000 
km-' of sea water. This high diversity of 
freshwater fishes relative to habitat extent is 
undoubtedly promoted by the extent of isol- 
ation between freshwater systems. Many 
lineages of fishes and invertebrates have 
evolved high diversity in certain water sys- 
tems, and in some cases, species richness 
and endemism tend to be positively correlated 
between different taxonomic groups". 

As is the case with terrestrial habitats, 
species richness increases strongly toward 
the equator, so that in most groups of 
organisms, there are many more species in 
the tropics than in temperate regions, 
although in a tew specific cases (e.g. fresh- 
water crayfish! this appears to be reversed. 

Protoctlsts 

The larger algae comprise some 5 000 species 
in three major groups (the green, brown and 
red algae!, the great majority of which are 
marine or brackish water forms ('seaweeds']. 
The green algae Chlorophyta includes one 
order of around 80 species (Ulotrichales) that 
is mainly freshwater However, one major 
group usually associated with the green algae 
- the stoneworts (charophytes) - is almost 
entirely freshwater The stoneworts include 
some 440 species, most of which are endemic 
at continent level or below; they tend to be very 
sensitive to nutrient enrichment and have 
declined in many areas'^ 



Fungi 

There are more than 600 species of fresh- 
water fungi known, currently more from 
temperate regions than from the tropics, 
although probably only a small fraction of 
existing species have been described, and the 
tropics have been little sampled''. Virtually all 
described freshwater fungi are ascomycotes 
with few basidiomycotes and zygomycetes 
having been identified. They occur wherever 
vascular plant material is available as a 
substrate. They appear to be important as 
parasites, endotrophs and saprotrophs of 
emergent aquatic macrophytes, as decom- 
posers of submerged allochthonous woody 
debris, and as a food resource for inver- 
tebrates' '-. Most are very small, with 
sporomes (fruiting bodies! less than 0.5 mm 
in diameter. 




Plants 

Wetland or aquatic species occur with some 
frequency in the non-vascular plant phyla, 
which generally prefer moist habitats, and 
among the ferns and allies. Mosses in the 
order Sphagnales (a single family Sphag- 
naceae and genus Sphagnum] often grow 
submerged, and are key components of 
peat bogs. Many groups of damp-loving 
(hygrophilousj terrestrial mosses (e.g. 
Thamnium, Bryum, Mnium] have aquatic 
forms. Several genera of Bryales are aquatic 
or have aquatic species. A number of 



Species richness in 
inland waters, in relation 
to habitat extent, is 
relatively high. 



170 WORLD ATLAS OF BIODIVERSITY 



Map 7.1 

Freshwater fish family 

diversity 

Family richness of typical 
bony fishes (Actinopterygii) 
in inland waters, plotted as 
a world density surface. It is 
based on generalized range 
maps of 157 families. Color 
depth represents the 
number of families, up to a 
maximum of kit, potentially 
present at any point. Two 
families of cartilaginous 
fishes lElasmobranchiil that 
together have a very few 
inland water species are 
omitted. About ten or so 
families of bony fish that 
occur in coastal and 
estuarine waters, but do 
not extend significantly into 
inland freshwaters, are 
omitted. Several families 
range more or less widely 
in inland waters and also 
occur in coastal and 
estuarine waters around 
the continents, but this 
peripheral part of the range 
is in most cases not 
represented. 

Source: Produced by UNEP-WCMC 
using range maps prepared from 
information in Berra . 



Density 




High 



Low 



liverwort species growing otherwise in wet 
terrestrial situations may also live sub- 
merged, sometimes at considerable depths. 
Truly aquatic liverworts include several 
species of Riccia and Ricciocarpus natans 
[Ricciaceae; MarchantialesI that live free- 
floating on the surface of eutrophic lakes. At 
least 16 species of Riella in the Riellaceae 
(JungermannialesI are aquatic forms charac- 
teristic of temporary waters in semiarid 
regions, reaching highest diversity in north- 
ern and southern Africa. Among the lyco- 
phytes, most of the 60 or so species of 
Isoetes (family Isoeteacael are aquatic, some 
of great limnological importance, and Stytites 
is an endemic member of the littoral 
community of Andean lakes. Sphenophytes 
Ihorsetailsl often occur in moist situations, 
including around water margins. Equisetum 







icr-'v- 






Jlfi>c 




t 



\ 1 




fluviatite, for example, is a notable emergent 
littoral form of north temperate lakes. 

The Filicinophyta include several aquatic 
forms. The genus Ceratopteris (four species, 
family Pteridaceael has the only truly aquatic 
(floating! homosporous ferns; some are culti- 
vated ornamentals, others are edible. A 
few other species, e.g. Microsorum pteropus 
(Polypodiaceael and Microlepia speluncae 
(Dennstaedtiaceae) can grow in water. Among 
heterosporous ferns, the family Marseliaceae 
comprises three genera and 55-75 species 
which are either amphibious or fully aquatic. 
All members of the families Salviniaceae (one 
genus and about ten species) and Azollaceae 
(one genus and around six species] are 
floating aquatic ferns. The latter supports the 
symbiotic nitrogen-fixing Anabaena azollae 
(phylum Cyanobacteria). 



Inland water biodiversity 171 






ry- 



■V 




\ 



Vascular plants are essentially terrestrial 
forms, and existing aquatic species are 
derived from terrestrial ancestors; several 
different lineages include aquatic species and 
this transition has therefore occurred several 
times. It has been estimated that at most 
1 percent of angiosperms, i.e. up to 2 700 
species, are aquatic". Around U angiosperm 
families consist largely or exclusively of in- 
land water forms (Ceratophyllaceae, Hippuri- 
daceae, Hydrostachydaceae, Nymphaeaceae, 
Podostemaceae, Trapaceae, Butomaceae, 
Hydrocharitaceae, Lemnaceae, Limnochari- 
taceae, Najadaceae, Pontederiaceae, Potamo- 
getonaceae, Zannichelliaceael. 

Most inland water plant species are 
relatively widespread, ranging over more than 
one continental land mass; many are cosmo- 
politan, occurring around the world and on 



, r^ k 




remote islands. Of the widespread forms, 
some are essentially northern temperate 
species extending to a great or lesser extent 
into the tropics; some are mainly tropical". 
The Podostemaceae is particularly note- 
worthy for its many monotypic genera, and a 
large number of narrowly endemic species, in 
at least one instance with several forms 
restricted to different stretches of a single 
river Tropical South America, Madagascar, 
Sri Lanka, India, Myanmarand Indonesia hold 
such localized species". 

Inland water animals 

Animal species are considerably more diverse 
and numerous in inland waters than plants. 
Most of the major groups include terrestrial 
or marine species as well as freshwater 
forms. Apart from fishes, important groups 



172 WORLD ATLAS OF BIODIVERSITY 



Table 7.4 

Insects of inland waters 

Notes: Data refer to number 
of species; all estimates 
approximate. 

i Partially aquatic as adult 

and sometimes as nympfi. 

ii Number m parentfieses 

refers to fully aquatic 

Nepomorpha. 

ill All tfiese species are 

parasitoids as larvae. 



Source: Collated from data in 
Hutctiinson . 



With inland water species include crus- 
taceans (crabs, crayfishes, shrimps, as well 
as planktonic forms such as filter-feeding 
Cladocera and filter-feeding or predatory 
Copepodal, moUusks (including mussels 
Bivatvia, and snails Gastropoda], insects 
(including stoneflies Plecoptera, caddisflies 
Trichoptera, mayflies Ephemopteral, sponges, 
flatworms, polychaete worms, oligochaete 
worms, numerous parasitic species in various 
groups, and numerous microscopic forms. 
Information is incomplete for many groups, 
but crustaceans and mollusks have speciated 
profusely In certain freshwater systems, with 
a tendency to form local endemic species. 
Because of the feeding mode - attached 
bottom-living filter-feeders - bivalves can 
help maintain water quality but tend to be 
susceptible to pollution (their larvae are 
parasitic on fishesl. The diversity and eco- 
logical role of microorganisms and micro- 
Invertebrates In freshwater sediments have 
been reviewed". 

Insects with an aquatic larval phase but a 
winged adult phase are often restricted to 
particular river basins (even If adults disperse 
widely, they may not find suitable habitat], but 
in general are much less restricted in this way 
than entirely aquatic species. A relatively 
large number of species, particularly of 
crustaceans, occupy temporary pools and 
have a stage that is desiccation-resistant and 
can undergo long-range passive dispersal 





Ephemeroptera 


84 


614 


224 


2 250 


Odonata 


302 


415 


127 


4 875 


Plecoptera 


196 


578 


387 


2140 


Orthoptera' 


- 


ca20 


- 


ca20 


Blattodea' 


- 





- 


calO 


Hemlptera 


236 


404 


129181]" 


3 200 


Megaloptera 


26 


43 


6 


300 


Neuroptera 


58 


6 


9 


calOO 


Coleoptera 


730 


1655 


1072 


5 000 


Hymenoptera" 


- 


55 


74 


calOO 


Diptera 


1300 


5 547 


4 050 


>20 000 


Trichoptera 


478 


1340 


895 


7 000 


Lepldoptera 


- 


- 


5 


calOO 



between drainage basins; some such species 
are thus widely distributed. 

Insects 

As on land. Insects (phylum Mandibulata] are 
as far as is known by far the most diverse 
group of organisms In Inland waters. The 
true number of aquatic Insects remains 
unknown; data for three relatively well 
known areas (Europe, Australia and North 
America] and extrapolations for possible 
global totals are Included in Table 7.A. In 
contrast to terrestrial faunas, where beetles 
(order Coleoptera] are the most diverse, flies 
and their relatives (order Diptera] appear to 
be by far the most abundant group In inland 
water habitats, although also one of the less 
fully known. 

In terms of life histories, there are two 
main groups of aquatic insects: those In which 
the adult stage and the active Immature 
stages are passed in water (in some cases 
with a terrestrial pupal stage]; and those in 
which, afte'- a nymphal or larval stage in 
water, the adult stage is spent on land or In 
the air The great majority of Diptera, and 
therefore most aquatic insects, form part of 
the latter group. Included amongst their 
number are several of enormous economic 
significance to humans, of which the most 
significant are almost certainly mosquitoes of 
the genus Anopheles, Intermediary hosts of 
the malaria parasite. 

Most aquatic insects are benthic, living In 
or on the bottom; a small number are 
planktonic and live suspended In the water 
column. Around half of the aquatic Hemlptera 
and a few other insects and non-Insect 
invertebrates live on the water surface 
(epipleuston]. 

Fishes 

Around 40 percent of known fish species 
occur In freshwater: almost exactly 10 000 
species are confined to freshwater, and a 
further 1 100 or so occur in freshwater but are 
not confined to It (Table 7.5]. These last 
Include species, such as many salmonlds, 
that grow in the seas but ascend rivers to 
spawn (anadromous], and others, such as 
eels, that grow in inland waters but spawn at 



Inland water biodiversity 173 



sea (catadromousl. Freshwater fishes are 
taxonomically diverse, although not as diverse 
as marine ones. Thirty-tour of the 57 or so 
extant orders of fishes have at least one 
strictly freshwater species, while a further 
two, the sawfishes (Pristiophoriformesl and 



tarpons (Elopiformes) have species that occur 
in freshwater but are not confined to it". 
This compares with 38 orders that have at 
least one marine species. 

Of the orders of fishes with freshwater 
species, ten are entirely freshwater and 



Table 7.5 

Fish diversity In Inland 

waters, by order 



Source: Nelson (differs in detail from 
tfie later taxonomy of Eschmeyer 1. 



■flll!^"""" 




ToT^^ 


■^JffiF 


^^0^^ 


^To^t 


^^oTt 


Approximate Approximate 


[strictly freshwater 




families 


genera 


species 


strictly 


species 


% strictly 


% using 


orders in boldl 










freshwater 
species 


using 
freshwater 


freshwater 


freshwater 


Petromyzontiformes 


Lampreys 


1 


6 


41 


32 


41 


78 


100 


Carcfiarhiniformes 


Ground stiarl<s 


7 


47 


208 


1 


8 





4 


Pristiophoriformes 


Sawfishes 


1 


2 


5 





1 





20 


Rajiformes 


Rays 


12 


62 


456 


24 


28 


5 


6 


Ceratodontiformes 


Australian lungfish 


1 


1 


1 


1 


1 


100 


100 


Lepidoslreniformes 


Lungfishes 


2 


2 


5 


5 


5 


100 


100 


Acipensiformes 


Sturgeons 


2 


6 


26 


14 


26 


54 


100 


Amiiformes 


Bowfin 


1 


1 


1 


1 


1 


100 


100 


Anguiliformes 


Eels 


15 


141 


738 


6 


26 


1 


4 


Atheriniformes 


Silversides 


8 


47 


285 


146 


171 


51 


60 


Batrachoidiformes 


Toadfishes 


1 


19 


69 


5 


6 


7 


9 


Beloniformes 


Needlefisfies, sauries. 


















flyingfishes, halfbeaks 


5 


38 


191 


51 


56 


27 


29 


Characlformes 


Characins 


10 


237 


1343 


1343 


1343 


100 


100 


Clupeitormes 


Herrings and anchovies 


5 


83 


357 


72 


80 


20 


22 


Cypriniformes 


Carp, minnows, loaches 


5 


279 


2 662 


2 662 


2 662 


100 


100 


Cyprinodontiformes 


Rivulines, killifishes, pupfishes, 


















poeclliids, goodeids 


8 


88 


807 


794 


805 


98 


100 


Elopiformes 


Ladyfishesand tarpons 


2 


2 


8 





7 





88 


Esociformes 


Pikes and mudminnows 


2 


4 


10 


10 


10 


100 


100 


Gadiformes 


Cods, hakes, rattails 


12 


85 


482 


1 


2 








Gasterosteiformes 


Pipefishes, sticklebacks, sandeels, etc. 


11 


71 


257 


19 


41 


7 


16 


Gonortiynchiformes 


Milkfish and beaked sandfishes 


k 


7 


35 


28 


29 


80 


83 


Gymnotlformes 


Knifefishes 


6 


23 


62 


62 


62 


100 


100 


Muglliformes 


Mullets 


1 


17 


66 


1 


7 


2 


11 


Ophidiiformes 


Pearlflshes, cusk-eels, brotulas 


5 


92 


355 


5 


6 


1 


2 


Osmeriformes 


Smelts 


13 


74 


236 


42 


71 


18 


30 


Osteoglossiformes 


Bonytongues 


6 


29 


217 


217 


217 


100 


100 


Perciformes 


Perches, basses, sunfishes, whitings, etc. 


U8 


1496 


9 293 


1922 


2815 


21 


30 


Percopslformes 


Trout-perches, pirate perch, cavefishes 


3 


6 


9 


9 


9 


100 


100 


Pleuronectiformes 


Plaice, flounders, soles 


11 


123 


570 


6 


20 


1 


4 


Polypteriformes 


Bichirs 


1 


2 


10 


10 


10 


100 


100 


Salmoniformes 


Salmonids 


1 


11 


66 


45 


66 


63 


100 


Scorpaeniformes 


Gurnards, scorpionfishes, velvetfishes, etc. 


25 


266 


1271 


52 


62 


4 


5 


Semionotiformes 


Gars 


1 


2 


7 


6 


7 


86 


100 


Siluriformes 


Catfishes 


34 


412 


2 405 


2 280 


2 287 


95 


95 


Synbranchiformes 


Swamp-eels 


3 


12 


87 


84 


87 


97 


100 


Tetraodontiformes 


Triggerfishes, puffers, boxfishes. 


















fllefishes, molas 


9 


100 


339 


12 


20 


4 


6 

■1 



174 WORLD ATLAS OF BIODIVERSITY 



Virtually all reptiles of 
inland waters return to 
land, at least to nest. 



anottier five are very largely so (witfi more than 
80 percent of their known species in fresh- 
watersl. A further 13 are very largely marine 
with a small proportion of freshwater species 
l<10 percent] while the remainder have 
significant numbers of both marine and 
freshwater species. Over 80 percent of fresh- 
water species are confined to just four orders: 
the carps and their relatives ICypriniformesI; 
the characins iCharaciniformesl; the catfishes 
(Siluriformesl; and the perches and their 
relatives (Perciformesl. The first three of these 
are wholly or almost entirely freshwater, while 
the last, the largest order of fishes with nearly 
AO percent of known species, is unusual in 
having significant numbers of both marine and 
freshwater species. 

Amphibians 

The great majority of the 5 000 or so living 
amphibian species have aquatic larval stages 
and, as none is known to occur in sea water, 
all these are dependent on inland waters of 
various kinds for continued survival of 
populations. In some cases such water bodies 
may be temporary pools or puddles, or water 
in the leaf axils of plants. Relatively few 
species are fully aquatic (Table 7.6). Although 
the number of fully aquatic species in each of 
the three extant orders is roughly similar 
lea 20-30 in each], these represent very 
different proportions of each order, being less 
than 1 percent of anurans, around 5 percent 
of caudate amphibians and more than 
10 percent of caecilians. 

Aquatic caudate amphibians are neotenic, 
that IS retain features of the larval stage, most 
notably external gills. In addition to the fully 
aquatic amphibians (several of which can 
survive for short periods in damp conditions 
out of water], many other species may lead 
largely aquatic lives or may, as in the case of 
the Mexican axolotl Ambyostoma mexicanum, 
have completely aquatic neotenic populations. 

Reptiles 

Very few completely aquatic inland water 
reptiles are known. The three file-snakes in 
the family Acrochordidae are live-bearing and 
may pass their entire lives in water, often in 
coastal and estuanne areas as well as 



freshwaters. Virtually all other reptiles of 
inland waters are egg-laying and return to 
land at least to nest; most also spend a 
proportion of their time on land, often basking 
on banks or logs. The two most aquatic orders 
are the Crocodilia and the Chelonia. All the 
22 or so extant species of the former are 




predominantly aquatic and occur in fresh- 
waters, although one or two may also be found 
in marine areas. Of the latter, some two thirds 
of the 250 or so extant species are largely or 
predominantly aquatic, and a further 30 or so 
species may be considered amphibious. 

Aside from the file-snakes and the homal- 
opsine snakes, a number of other snake 
species are at least semi-aquatic. These in- 
clude several genera of natricine snakes, 
including the North American Nerodia and the 
Asiatic Sinonatrix, as well as the anacondas 
Eunectes [family Boidae] and the water cobra 
Boulengeria annulata (family Elapidae]. 
Amongst lizards, no wholly aquatic species is 
known. However, many can swim proficiently, 
often using water as a means of escape from 
predators, and a number are semi-aquatic. 
These last include several Australian and 
Old World monitors Varanus spp. [family 
Varanidae], water dragons Hydrosaurus and 
Physignathus [Agamidae], the crocodile lizard 
Shinisaurus from south China [Xenosauridael, 
the Bornean earless monitor Lanthanotus 
[Lanthanotidae], and a number of New World 
teiids [Teiidael. 



Inland water biodiversity 175 



^*K: 



^ Class and order 


Family 


No. of freshwater species 


AMPHIBIA 








Caudata 


Amphiumidae 


Congo eels 


3 




Cryptobranchldae 


Giant salamanders and hellbenders 


3 




Plellnodontidae 


Lungless salamanders 


4' 


^ 


Proteidae 


Mudpuppies and olm 


6 


B 


Sirenidae 


Sirens 


3 


^nura 


Pseudidae 


Paradox frogs 


3 




PIpidae 


Clawed frogs and pipid toads 


28 


Gymnophiona 


Typhlonectidae 


Typhlonectid caecilians 


19" 


REPTILIA 








Chelonia 


Carettochelidae 


Pig-nosed soft-shelled turtle 


1 




Trionychidae 


Soft-shelled turtles 


23 




Platysternidae 


Big-headed turtles 


1 


•', 


Chelydrldae 


Snapping turtles 


2 


K 


Dermatemydidae 


Central American river turtle 


1 


■ 


Chelldae 


Austro-american side-necl<ed turtles 


37 


■ 


Kinosternidae 


Mud and musl< turtles 


12 


■: 


Pelomedusidae 


Side-necl<ed turtles 


22 


^ 


Emydidae 


Pond and river turtles 


6i 


Crocodilia 


Alllgatoridae 


Caimans and alligators 


8 




Crocodylidae 


Crocodiles 


12 




Gavialidae 


Gharialand false gharial 


2 


Squamata 


Acrochordidae 


File-snal(es 


3 




Colubridae 


Colubrid snal<e5 


iO'' 


AVES 








Anseriformes 


Anseranatidae 


Magpie goose 


1 




Dendrocygnidae 


Whistling-ducks 


9 




Anatidae 


Ducl(s, geese and swans 


W 


Gruiformes 


Heliornitiiidae 


Limpkin andsungrebes 


3 




Rallidae 


Rails, gallinules and coots 


ir 


Ciconiiformes 


Jacanidae 


Jacanas 


8 




Laridae 


Gulls, terns, skuas, auks, skimmers 


22 1131' 




Podicipedidae 


Grebes 


21 




Anhingidae 


Anhingas 


4 




Phalacrocoracidae 


Cormorants and shags 


5121' 




Phoenicopteridae 


Flamingos 


5 




Pelecanidae 


Pelicans and shoebill 


6 




Gaviidae 


Divers 


5 


Passeriformes 


Cinclidae 


Dippers 


5 


MAMMALIA 








Monolremala 


Ornithorhynchidae 


Platypus 


1 


Didelphiomorpha 


Didelpliidae 


Opossums 


1 


Insectivora 


Tenrecidae 


Tenrecs and otter shrews 


4 




Soricidae 


Shrews 


3 




Talpidae 


Moles and desmans 


2 


Rodentia 


Castoridae 


Beavers 


2 




Muridae 


Voles and mice 


32 




Hydrochaeridae 


Capybara 


1 


[ Cetacea 


Platanlstidae 


River dolphins 


5 


Sirenia 


TrIchechidae 


Manatees 


3 


Carnivora 


Mustelidae 


Mustelids, otters 


13 




Viverridae 


Viverrids 


3 




Phocidae 


Earless seals 


2 


Artiodactyla 


Hippopotamidae 


Hippopotamus 


1 



Table 7.6 

Tetrapod diversity in 

inland waters 

Notes: Entirely freshwater 
families in bold. Taxonomy 
based on the same vertebrate 
sources cited in Table 2.1. 
Bird groups here differ to 
some extent from Table 7.9 
which uses a more traditional 
arrangement of bird higher 
taxa. 

i Genera Leurognathus, 

Haideotnton. Typhlomolge 

only 

11 Sometimes included in 

the Caecilidae. 

ill Subfamily Homalopsinae. 

iv Genus Fulicula Icoots] 

only 

V Figures in parentheses 

indicate those species of 

the total that breed largely 

or entirely inland also 

included as seabirds in 

Table 6.6. 



176 WORLD ATLAS OF BIODIVERSITY 



Map 7.2 

Major areas of diversity 

of inland water fish 

This map represents an 
informal synthesis of 
documented expert opinion 
on globally important areas 
for freshwater fish diversity, 
taking into account species 
richness and endemism. 
Two categories are shown: 
discrete areas and systems 
known to be of high 
diversity, and areas where 
diversity is globally 
important but less 
concentrated. 

Note: For numbered 
locations see Appendix 6. 

Source: Comprled with the help of 
members of lUCN/SSC specialist groups 
and other ichthyologists; fjrsl published 
inWCMc'. 



Fish diversity 

^^^H Key areas 

Other important areas 

Birds 

Unlike mammals, there are no wholly aquatic 
birds, because all species lay eggs that 
cannot survive prolonged immersion in water; 
however, a much higher proportion of bird 
than mammal species is associated with 
inland water ecosystems. As with other 
tetrapod groups, it is impossible to separate 
rigidly inland water species from primarily 
terrestrial forms or from seabirds. Table 7.6 
includes bird species that are highly adapted 
to aquatic ecosystems and largely or ex- 
clusively inhabit inland waters, rather than 
marine or coastal areas. Nearly 60 percent of 
the roughly 250 species belong to one family - 
the Anatidae - all of whose members are 
more or less associated with aquatic habitats, 
although some [such as most goose species! 
feed very largely on land. It is noteworthy 



?■!' 
W 



103 9^ 

O 101 96 



99 
97 



95 



100~-:98< 



104 t , 



123-1 



124 



N^131; 



116 




that, among the Passeriformes or perching- 
birds - by far the most species-rich order 
(accounting for over half of all bird species) - 
only one small family, the dippers (Cinclidael. 
can be considered truly aquatic in habits. 

In addition to these, there are a large 
number of wading bird species associated 
with inland or coastal wetland and littoral 
habitats. These include all or most members 
of the following families: Anhimidae (scream- 
ers - three species); Eurypygidae (sunbittern - 
one species); Gruidae (cranes - 15 species); 
Rallidae (rails, gallinules and coots - 142 
species); Scolopacidae (sandpipers and their 
relatives - 88 species); Rostratulidae (painted- 
snipe - two species); Charadriidae (plovers 
and their relatives - 89 species); Ardeidae 
(herons, bitterns and egrets - 65 species); 
Scopidae (hammerkop - one species); 



Inland water biodiversity 177 



"OX 




65 ,73 39 ?'^' 



41/ 



^42 



Ttiresl<iornitlnidae (ibises and spoonbills - 3A 
species!; and Ciconiidae (storks - 26 species). 
There are also a considerably smaller number 
of non-wading birds that feed largely on fishes 
and other aquatic animals and are adapted to 
diving or surface-snatching. Among these are 
many kingfishers (families Alcedinidae, 
Dacelonidae and Cerylidael, the fish-owls 
[Ketupa and Scotopelia spp., family Strigidael, 
fish-eagles iHaiiaeetus and Ichthyophagus 
spp., family Accipitridae) and a few other 
raptors. 

Mammals 

Wholly aquatic inland water mammals are 
confined to two orders, Cetacea and Sirenia. 
In the former, four of the five species of the 
family Platanistidae (the river dolphins! are 
confined to river systems and the fifth occurs 



in estuarine and coastal waters. A number of 
other cetaceans may enter the lower reaches 
of river systems but all are predominantly 
marine. Two of the four living species of 
Sirenia - the Amazonian manatee Trichechus 
inunguis and the West African manatee 
T. senegalensis - are wholly or very largely 
confined to freshwaters, and a third, the 
Caribbean manatee T. manatus, is found in 
both inland and marine waters. 

Amongst other groups, a number of 
species may lead more or less aquatic lives 
but all these are effectively amphibious, in that 
at the very least they produce young on land. 
For most of these, a predominantly aquatic life 
is evident from direct observations, but for 
some (chiefly the Muridae! it is inferred from 
morphological adaptations. In direct contrast 
to terrestrial systems, where the majority of 



^«- 



178 WORLD ATLAS OF BIODIVERSITY 



mammal species are herbivores, a high 
proportion of these amphibious species 
laround 50 percent! are carnivores, with only 
eight true herbivores and most of the aquatic 
murids believed to be omnivores (though 
predominantly insectivorous or piscivorousl. It 
is also notew/orthy that at least four of these 
species - the two beavers Castor spp., the 
hippopotamus Hippopotamus amphibius and 
the capybara Hydrochaeris hydrochaeris - 
feed very largely or entirely on land plants. In 
addition to the species included in Table 7.6, a 
number of other mammals are largely or 
wholly confined to wetland habitats (marshes, 
floodplains and swampsl. These include three 
African antelopes - the Nile lechwe Kobus 
megaceros. red lechwe Kobus teche and 
sitatunga Tragelaphus spekei - and the 
South American marsh deer Blastocerus 
dichotomus. 




Freshwater is essential to 
human surA/ival. 



HUMAN USE OF AND IMPACT ON INLAND 
WATERS 

Freshwater - as precipitation, groundwater or 
in inland water ecosystems - is essential for 
human survival, chiefly because humans 
must drink and also because it is needed, in 
far greater quantity, to produce food. It also 
has a wide range of subsidiary uses - for 
transport, industrial production, cleaning, 
waste disposal, generation of hydroelectric 
power, recreation, esthetic purposes and in 
the form of inland water ecosystems as sites 
for the production of food. 

Many of these demands conflict with each 
other, so that for example the use of water for 



disposal of noxious wastes is incompatible with 
the provision of safe drinking water Moreover, 
while the amount of freshwater available is 
limited, demands on it continue to grow 
steadily as the global human population con- 
tinues to expand. This problem is exacerbated 
by the fact that freshwater Is unevenly 
distributed around the world, so that it is often 
not available where and when needed, nor in 
the appropriate amounts, nor with the 
necessary quality The two last are particularly 
important for the maintenance of freshwater 
biodiversity. Freshwater systems are therefore 
under growing pressure, as flow patterns are 
disrupted and the load of waste substances 
increases. Inevitably, per capita shares of 
water for human use are decreasing and water 
stress is becoming more widespread". 

Agriculture consumes around 70 percent 
of all water withdrawn from the world's riv- 
ers, lakes and groundwater^". In places, more 
than half the water diverted or pumped for 
irrigation does not actually reach the crop, 
and problems of waterlogging and salln- 
ization (deposition in soil of salts left by 
evaporation of pumped groundwater! are 
increasing. However, irrigated agriculture 
produces nearly AO percent of world food and 
other agricultural commodities on only 17 
percent of the total agricultural land area, and 
is thus disproportionately important to global 
food security-". 

The principal use of freshwater species, 
not considering the properties of aquatic sys- 
tems themselves. Is as food. Subsidiary uses 
Include the aquarium trade, materials for 
medicinal or ornamental use, and as fertilizer 
Inland water fishery production has two com- 
ponents: capture fisheries and aquaculture, 
although as discussed below the distinction 
between the two is becoming increasingly 
blurred. For many human communities, par- 
ticularly in countries less developed indus- 
trially, capture fisheries provide a major 
portion of the diet. Although it appears to be 
under-reported, inland water production has 
usually been regarded as far less important 
than marine fisheries and. with few excep- 
tions where countries have access to both 
marine and inland aquatic resources, repor- 
ted yield from inland waters is a small 



Inland water biodiversity 179 



^*t^ 



fraction of marine yield. Even m landlocl<ed 
countries, the recorded inland harvest is often 
[o\N both in absolute size and in relation to 
consumption of meat and other agricultural 
produce. 

INLAND WATER FISHERIES 
Capture fisheries and aquaculture 

Globally, the reported inland w/ater capture 
fishery for 1999 amounted to 8.2 million 
metric tons, w/ith 19.8 million metric tons of 
aquaculture production recorded''; over 85 
percent of the former and about 98 percent of 
the latter comprised finfishes, with virtually 
all the remainder being freshwater crus- 
taceans and mollusks"". The crustaceans are 
mainly crayfishes and freshwater shrimps, 
both exploited for food, and most of the 
mollusks are bivalve, taken tor pearls and for 
food. These reported totals compare with 
reported marine capture fisheries of some 84 
million metric tons, and marine and brackish 
water aquaculture animal production of 
around 13 million metric tons (see Chapter 6|. 

The reported global inland water capture 
fishery has increased slowly in the period 
1984-99, by nearly 2 percent per year, al- 
though this masks considerable regional 
variation, with declines in some areas (e.g. 
Europe and North America) and more marked 
increases elsewhere (notably Asia]". Reported 
inland aquaculture production has been rising 
at a higher rate, and was well over twice the 
reported production of inland capture fisheries 
in 1999 (Figure 7.11. A major proportion of 
global inland aquaculture is produced by 
countries in Asia. China alone reportedly gen- 
erates more than one quarter of the global 
total (Table 7.71, and has been responsible for 
most of the recent increase in this sector In 
this particular instance the dividing line 
between aquaculture and capture fisheries is 
indistinct; no husbandry is involved beyond 
release of hatchery stock, and the fishery 
operates as a capture fishery''. 

However, national statistics do not ad- 
equately reflect the actual magnitude, location 
or importance of inland fisheries. The repor- 
ted inland capture production is certainly a 
gross underestimate because much of the 
catch is made far from recognized landing 



30 



25 



c20 



15 



10 



Aquaculture 




1994 



1995 



1996 



1997 



1998 



1999 



places where catches are monitored, and is 
consumed directly by fishers or marketed 
locally without ever being reported. The evi- 
dence suggests that actual capture fisheries 
catch may be twice or conceivably even three 
times the reported total, i.e. around 15-23 
million metric tons per year". Because a far 
higher proportion of inland fisheries than 
marine fisheries harvest is apparently used 
directly for human consumption (rather than 
production of oils and meals, often used for 
livestock feed), and because discards are 
believed to be negligible, it has been argued by 
some that the provision of foodfish from inland 
waters is not that much less than that from 
recorded marine catch". 

Inland water capture fisheries, particularly 
in countries less developed industrially, 
certainly provide a staple part of the diet for 
many human communities. This is the case in 
West Africa generally, locally in East Africa, 



Figure 7.1 

Reported global inland 

fisheries production 



Table 7.7 

Major inland fishery 

countries 

Note. The top ten producing 
countries in 1998. 





Production (metric tons) 


% of world total 


China 


2 280 000 


28.5 


India 


650 000 


8.1 


Bangladesh 


538 000 


6.7 


Indonesia 


315 000 


3.9 


Tanzania 


300 000 


3.7 


Russia 


271 000 


3.4 


Egypt 


253 000 


3.2 


Uganda 


220 000 


2.8 


Thailand 


191 000 


2.4 


Brazil 


180 000 


2.3 



180 WORLD ATLAS OF BIODIVERSITY 



Map 7.3 

Major areas of diversity of 

Inland water moUusks 

This map illustrates the 
location of areas regarded 
as globally important for 
diversity in the bivalve and 
gastropod moUusks of 
inland waters, taking into 
account species richness 
and local endemism. 

Note: For numbered 
locations see Appendix 6. 

Source: Compiled using information and 
expertise provided by ttie lUCN/SSC 
Mollusc Specialist Group; tirst published 
inWCMC'. 



MoUusk diversity 

^^^H Innportant areas 



and in parts of Asia and Amazonia. In some 
landlocked countries inland fisheries are of 
crucial importance, providing more than 
50 percent of animal protein consumed by 
humans In Zambia" and nearly 75 percent 
In Malawi''. In the low-income food-deficit 
countries fish protein may be particularly 
important In times of food scarcity. 

It is impossible at the global level to carry 
out any meaningful analysis of the relative 
contribution of different species or species 
groups to inland capture fisheries because of 
the inadequacy of reporting. In FAO IFood and 
Agriculture Organization of the United Nations! 
statistics, by far the largest group recorded Is 
'freshwater fishes not elsewhere included', 
that is those that are completely unclassified 
other than being identified as finfishes. These 
make up just under half of all reported landing 






il07 



^ 



;^H' 



■f 



134 



by weight with a further 15 percent consisting 
of moUusks and crustaceans similarly classi- 
fied. The majority of the remaining catch Is 
classified Into broad species groups (e.g. 
cyprlnlds, characlns, siluroldsl, with only three 
individual fish species having annual reported 
global landings of more than 100 000 metric 
tons. These are the Nile perch Lates niloticus 
lea 330 000 metric tons reported in 1997), Nile 
tllapia Oreochromis niloticus (226 000 metric 
tons in 19971 and the common carp Cypnnus 
carpio (100 000 metric tons in 19971. 

It Is, however, evident that the importance 
of different species of freshwater finfishes 
varies considerably between different areas. 
In terms of food security for local subsistence 
or mixed market/subsistence communities, 
particularly In the tropics, there is an increas- 
ing amount of evidence that the diversity of 



Inland water biodiversity 181 



yfP^^^^ 




X. 



v£) 



85 



83 -''1^ ;!|^,89 y, 



-.r 



.'iS 




species harvested is in itself a major factor in 
ensuring a continuous food supply. Many of 
the species that contribute to these fisheries 
are often small and would be considered 
'trash' fishes in orthodox fisheries, Vi^ith much 
lower market value than larger (often non- 
nativel species that might therefore be 
considered for introduction. However, these 
small species are easy to preserve and keep 
under local conditions and moreover are 
eaten whole, providing a valuable source of 
calcium and other minerals. Larger species, 
such as the introduced Ni'ie perch in Lake 
Victoria, cannot be easily preserved locally 
and are in any case not eaten whole, leading 
to a danger of calcium deficiency. Fisheries 
for such species tend to become indus- 
trialized or semi-industrialized, producing 
fish products for commercial high-value 



markets, often for export. While these may 
improve the balance of payments for the 
countries concerned, they may ultimately 
worsen the nutritional status of local people. 
Additionally, there are some indications that 
fish populations in mixed species fisheries are 
more stable over time, less susceptible to 
boom and bust' than those based on a small 
number of often introduced species. 

It is difficult rigorously to assess the 
condition of inland fish stocks because they 
appear able to respond rapidly to changing 
environmental conditions. However, there is a 
consensus that, regionally, most stocks are 
fully exploited and in some cases over- 
exploited. Exploitation has become more 
efficient because of new technologies, and 
developing infrastructure has allowed easier 
access to freshwater resources. Some stocks. 



182 WORLD ATLAS OF BIODIVERSITY 



Map 7.-4 

Major areas of diversity of 
selected inland water 
crustacean groups 

This map represents a 
preliminary assessment of 
areas believed to support 
high diversity among three 
of the several major 
crustacean groups that 
occur in inland waters, 
taking into account species 
richness and local 
endemism 

Note: For numbered 
locations see Appendix 6. 

Source: Compiled using information and 
expertise provided by the lUCN/SSC 
Inland Water Crustacean Specialist 
Group; first published in WCt-tcV 




especially in nver fisheries, appear to be in 
decline, but this is seemingly a result mainly 
of anthropogenic changes to the freshwater 
environment. 

In many parts of the world fishing has high 
recreational value, as well as being a means 
of food gathering. Locally, notably in the 
Amazon basin and in parts of Southeast Asia, 
capture for the ornamental fish trade may be 
an important source of income with potential 
impact on wild populations. Increasingly it is 
becoming difficult to distinguish between 
truly wild fish stocks and those that are 
artificially managed or enhanced in some way. 

Other harvested species 

Other exploited animal groups in inland 
waters are far less important globally than 
finfishes, but may still be highly significant. 







•N). 



112 




Apart from crustaceans and mollusks, men- 
tioned above, these include: frogs (chiefly 
family Ranidael, exploited for food; crocodil- 
ians, hunted mainly for leather; freshwater 
chelonians, taken for food and to a lesser 
extent for medicinal purposes, particularly in 
eastern Asia; waterfowl which are hunted for 
recreation and for food; fur-bearing mam- 
mals, such as beavers Castor spp., otters 
(subfamily Lutrinael and muskrats {Ondatra 
zibetfiicusand Neofiber atleni], taken for their 
skins; and manatees (family Trichechidael, 
taken mostly for food although also used non- 
consumptively on a small scale for biological 
control of weeds. 

Rice is the principal cultivated wetland 
plant of global importance to food security. 
Most of the relatively few plants associated 
with inland waters that are heavily exploited in 



Inland water biodiversity 183 



.^' 




the wild state are also marginal or wetland 
species. Some species le.g. Aponogeton, in 
Madagascar) are collected for use as orna- 
mentals; reeds are used as building materials 
le.g. thatch); and some are collected for 
food or as medicines le.g. Spirulina algael. 
Rhizomes, tubers and seeds [rarely leaves! 
of aquatic and wetland plants are used as a 
food source, mainly in less-developed regions 
where they can be important to food security 
in times of shortage, but globally they make 
a relatively minor contribution to human 
nutrition, fvlost important are some forms of 
edible aroid lAraceae), notably some cultivars 
of Colocasia Itarol and the giant swamp taro 
Cyrtosperma chamissonis that grow in flood- 
ed conditions and are important food crops in 
the Caribbean. West Africa and the Pacific 
islands. Conservation and collection of wild 



forms of these is considered a high priority. 
Sago palms Metroxylon spp. in Southeast 
Asia and the Pacific and watercress Rorippa 
nasturtium-aquaticum in Europe are other 
examples of cultivated aquatic plants, the wild 
relatives of which merit conservation. Aquatic 
plants have been widely used for medicinal 
purposes, documented for at least two 
millennia, but such use appears at present to 
be minor and probably of real significance in 
few areas. However, interest in ornamental or 
aquarium water plants is widespread and of 
some economic importance. 

OTHER MAJOR IMPACTS 

Physical alteration and destruction of habitat 

Destruction of inland water ecosystems is 
most simply effected by the removal of water 
Although humans have always made use of 



184 WORLD ATLAS OF BIODIVERSITY 



freshwater systems, the last 200 years 
(spanning the Industrial Revolution, the growth 
of cities, the spread of high-input agriculture! 
have brought about transformations on an 
unprecedented scale. The global rate of water 
withdrawal rose steeply at the start of the 20th 
century, and further after mid-century. Major 
changes in the distribution of water have 
resulted mainly from withdrawals for Irrigation 
and secondarily from domestic and industrial 
use. It has been estimated that humans use 26 
percent of the total evapotranspiratlon from 




Aral Sea" 

Until the mld-20th century the Aral Sea was the world's fourth largest inland water body 
(after the Caspian Sea, Lake Superior and Lal<e Vlctorial. Located within a catchment area of 
some 1 .9 million l<m' extending over six countries, the Aral is fed by two major rivers, the 
Amu Darya, rising in the Pamir, and the Syr Darya, rising in the Tien Shan. Starting in the 
19605, excess water withdrawal from these rivers, primarily for cotton irrigation, has 
severely affected the Aral Sea. Its area had reduced from more than 65 000 km' to about 
28 500 km' In 1998, with volume falling by 75 percent, water level falling by around 20 
meters (ml, and salinity greatly Increasing. In consequence, problems with drinking water 
quality and availability and with dustborne pollutants, have severely affected the health 
status of the human population; the commercial fishery has collapsed; waterlogging and 
salmlzation have degraded agricultural lands; and the deltaic marshlands of the two feeder 
rivers have largely been replaced by sandy drylands. 

Mesopotamia"" 

Serial satellite images confirm a loss of around 90 percent of the lakes and marshlands in 
the lower fvlesopotamlan wetlands during the last three decades. The only significant 
permanent marshland remaining is in the Al-Hawlzeh region. The large number of dams 
now present on upstream parts of the Tigris-Euphrates system may have contributed to this 
loss, but it appears to be pnmarlly the result of major hydrologlcal engineering works In 
southern Iraq, notably the completion of the major outfall drain (or 'third river'] which diverts 
water to the head of the Gulf This loss has placed further pressure on the t^la'dan (l»1arsh 
Arabs!, now largely displaced within Iraq or in refugee camps In Iran. Recent Information Is 
scarce, but biodiversity In the region will inevitably have been affected, probably including the 
endemic form of smooth-coated otter iLutrogale perspiciUata, an otherwise oriental species, 
assessed as globally threatened!. 

Azraq oasis'' 

Groundwater extraction for urban needs in Jordan rose from about 2 million m' in 1979 to 
about 25 million m' in 1993, with an additional 25 million m' per year used for agricultural 
Irrigation. The important Azraq wetlands natural reserve, formerly extending over some 
12 000 hectares, and a vital staging site for bird migrants, now supplies around one quarter 
of Amman's water needs, and as a consequence has lost most of its marshland and migrant 
bird populations. 



land surfaces and 54 percent of the accessible 
runoff". Unregulated withdrawal can lead to the 
wholesale destruction of inland water eco- 
systems, as has occurred with the Aral Sea In 
central Asia (Box 7.2!. Similarly, many wetlands 
have been completely destroyed by drainage, 
often for conversion to agriculture. Other fac- 
tors can modify or destroy particular habitats 
within Inland water ecosystems. For example, 
canalization, usually undertaken to improve 
navigability, generally destroys riparian (shore- 
line! habitats while flood-control systems 
drastically alter regimes In floodplalns". 

Dams and reservoirs 

Dams, particularly large dams, have a major 
impact on the rivers on which they are 
built. They affect flow regimes, often dram- 
atically, destroy large areas of existing habitat 
(while at the same time creating new ones! 
and can catastrophlcally disrupt the life cycles 
of species that migrate up and down rivers". 
Large dams are unevenly distributed across 
the world's major catchments, with a partic- 
ularly high concentration in North America, 
especially within the contiguous states of 
the United States, where at least eight catch- 
ments have In them more than 100 large 
dams each. In contrast, small and medium- 
sized dams, which may cumulatively have as 
substantial an Impact, are concentrated In 
eastern Asia, particularly China"'. Dams may 
be primarily for the generation of hydro- 
electric power, or to create reservoirs for the 
storage of water, or both. The size of a dam is 
not necessarily directly related to the area or 
volume of the Impoundment created or to its 
downstream impact. 

Pollution and water quality 

Assessment of anthropogenic changes in 
water quality Is not always easy as such 
changes are Invariably superimposed on 
natural background variations. Historically, a 
similar sequence of water quality issues has 
became apparent In both Europe and North 
America during rapid socioeconomic develop- 
ment over the past 150 years. Problems of 
fecal and other organic pollution were evi- 
dent In the mld-19th century, followed by 
salinization, metal pollution and eutrophi- 



Inland water biodiversity 185 



^^\ 



cation in tlie first lialf of tfie 20tfi century, with 
radioactive waste, nitrates and other organic 
micropoUutants, and acid rain most promin- 
ent in recent decades. Newly industrializing 
countries are likely to face these problems 
over a much more compressed period, and 
typically without the capacity to monitor and 
analyze water quality, or manage water use 
appropriately^l 

Different kinds of pollutants appear to 
affect different classes of water system to 
differing extent^'. With regard to quality for 
human use, contamination by pathogens of 
fecal origin is the major problem in river 
systems, and eutrophication probably the 
most widespread problem affecting lake and 
reservoir waters" "". 

Acid deposition through precipitation has 
been recognized as a regional transboundary 
phenomenon since the 1960s. Industrial 
emissions of sulfur and nitrogen oxides ISO2, 
NOJ, mainly a result of fossil fuel combustion, 
are the principal source of acid rain. Most 
evidence of acid rain and its effects relates to 
North America and Europe, but emission rates 
are rising steeply in rapidly industrializing 
countries elsewhere. Acid rain in one country 
may be a consequence of compounds released 
into the atmosphere by industry in another 
country hundreds of kilometers distant. The 
geology, soil and vegetation of drainage basins 
strongly influence the acidification process. 
Acid rain has been shown to decrease species 
diversity in lakes and streams but has not 
been implicated in any recorded species 
extinction or any major species decline. It has 
not yet been shown to be a significant issue in 
tropical freshwaters, where global freshwater 
diversity is concentrated'. 

Sedimentation 

Removal or extension of forest cover, or any 
anthropogenic interference with soils and 
landcover (e.g. agriculture, urbanization, road 
construction, mining], modifies the rate of 
runoff from catchment slopes and also the 
density of particles carried in the drainage 
system. All moving waters carry some mass 
of suspended material, and there is consider- 
able natural variation in this in space and 
time, but logging can increase sediment load 



by up to 100 percent for a short period, and 
20-50 percent over the longer term. Sediment 
reaching lakes will be deposited and in effect 
enter long-term storage; depending on water 
velocity, sediment in rivers will settle out on 
floodplains or other parts of the course, or be 
carried into the coastal marine environment. 
Increased sedimentation can have several 
effects on aquatic biodiversity: deposition can 
radically change the physical environment of 
species restricted to particular conditions of 
depth, light penetration and velocity; it is a 
major carrier of heavy metals, organic pollu- 
tants, pathogens and nutrients; and it can 
interfere mechanically with respiration 
In gill-breathing organisms'. The endemic 




cichlid fishes of the African Great Lakes rely 
on complex visual signals in breeding, and 
reduced water clarity because of sedimen- 
tation is suspected to be affecting their 
breeding success. 

Introduced species 

Unplanned or poorly planned introduction of 
non-native species and genetic stocks is a 
major threat to freshwater biodiversity. Such 
introductions can have negative or positive 
effects on fishery production; it is a reasonable 
assumption that all successful introductions 
will have an impact on existing population size 
and community structure, and many changes 
are likely to be undesirable''. The incomplete 



Although some factories 
have introduced cleaner 
production procedures, 
industrial emissions are 
still the principal source 
of acid ram. 



18« WORLD ATLAS OF BIODIVERSITY 



1 000 




Decade total 



_ 


■ ■ 




•^^ 


■^ 


m 


Wk 












1 


^" 


^" 


1 






1^ 


1 


1 


^^ 








i 










CD 


Cfl 


in 


yi 


Ul 


Ul 


Ul 


in 


in 


in 


in 


in 


in 


in 


in 


ID 


O 


CD 


CD 


CD 


CD 


CD 


CD 


CD 


o 


CD 


o 


CD 






CO 


Ln 


-o 






CT- 


O 




CNJ 


n 


-<f 


Ln 


o 


tr^ 


CO 




CO 


CO 


CO 


CO 


00 


o~ 


CN 


cr- 


Cr- 


o^ 


o- 


CN 


c>- 


CN 



Figure 7.2 
Inland water fish 
introductions 

Source: Welcomme . 

Figure 7.3 

Fresliwater population 
trends 

Note: A simplified 
representation of tfie average 
population change in a 
sample of 194 inland water 
species, see text. 



information available suggests tfiat althougfi a 
significant number of fish introductions took 
place during the 19th century, the three dec- 
ades starting with the 1950s were particularly 
important (Figure 7.21. A classic example of 
the effect of introduced species is the impact 
of the Nile perch Lafes niloticus on the 
haplochromine cichlids of Lake Victoria 
discussed further below. 

Several species of aquatic plant, in 
particular free-floating species that are able 
to spread rapidly by vegetative growth (most 
notoriously the South American water 
hyacinth Eichhornia crassipes], but also other 
forms, have dispersed widely over the globe 
and become major pest species. They block 




1970 



1975 



1980 



1985 



1990 



1995 



1999 



drainage channels, sluices and hydroelectric 
installations, impede boat traffic and hinder 
fishing. In recent decades the question of how 
best to control or eradicate pest species has 
been the foremost issue in conservation and 
management of aquatic plants'. 

THE CURRENT STATUS OF INLAND WATER 
BIODIVERSITY 

As with marine species, assessment of the 
status of wholly aquatic inland water species 
is hampered by difficulties of direct obser- 
vation. However, because these species also 
in general have far more restricted ranges 
than marine species, it is easier to infer their 
status from assessment of habitat condition 
and from sampling efforts. Amphibious or 
surface-dwelling species may be relatively 
easier to monitor. Where such species are of 
economic importance - as for example with 
those European and North American water- 
fowl that play a role in the recreational 
hunting industry - they may be among the 
best monitored of all wild species. 

Threatened and extinct species 

In the few cases where elements of inland 
water faunas - usually fishes - have been 
studied in any detail, it has generally been 
found that more species than suspected are 
threatened or cannot be re-recorded'°"". 
Among the 20 or so countries where the entire 
inland water fish fauna has been evaluated, 
an average of 17 percent of the species are 
regarded as globally threatened (categories 
critically endangered', endangered' or 
vulnerable' in the lUCN (World Conservation 
Union) threat categorization system"! (Table 
7. 81 A far larger proportion is likely to be in 
local decline, although not in danger of global 
extinction. The proportion of inland water 
chelonians that is believed threatened is even 
higher: 99 such species were categorized as 
threatened in 2000, equivalent to about 60 
percent of the number of inland water chelon- 
ians listed in Table 7.6. 

Amongst mammals and birds the pro- 
portions are considerably lower, probably 
because many semi-aquatic species are able 
to disperse from one inland water body to 
another relatively easily. Nevertheless, some- 



Inland water biodiversity 187 



wtiat more species ttian average for the 
groups as a wtiole are regarded as threatened. 
Table 7.9 shoves the category and taxonomic 
distribution of threatened inland water 
vertebrates. 

Two groups of species, the Lake Victoria 
cichlid fishes and the Mobile Bay drainage 
gastropod moUusks, serve as exemplary case 
studies illustrating the major threats faced by 
inland water biota worldwide. 

Lake Victoria, the largest tropical lake in 
the world, provides a classic example of the 
potential negative impacts of species intro- 
ductions. Until some 30 years ago, when the 
large top predator, the Nile perch La(es 
niioticus, was introduced, the lake supported 
an exceptional 'species flock' of more than 300 
species of haplochromine cichlid fishes, as 
well as smaller numbers from other families. 
The cichlids are of enormous interest in the 
study of evolutionary biology. Not all the 
species have yet been formally described; 
many of these are known among aquarists and 
others only by informal common names. At 
least half of the native species are believed 
to be extinct or so severely depleted that too 
few individuals exist for the species to be 
harvested or recorded by scientists. 

Predation by the Nile perch is believed to 
be the major cause of this decline, but 
important additional factors include increas- 
ing pollution and sediment load, excess fishing 
pressure, and possible competition from intro- 
duced tilapiine cichlids. The lake itself has 
now become depleted of oxygen, and a shrimp 
tolerant of oxygen-poor waters provides a 
major food source for the Nile perch. In recent 
years the Nile perch and one of the introduced 
tilapiines have formed the basis of a high- 
yielding fishery and an important national and 
export trade. However, it is thought unlikely 
that such high yields will be maintained 
because of continued overfishing and the 
suspected instability of the already highly 
disturbed lake ecosystem'. 

Dam construction is the prime cause of 
extinction in the gastropod fauna of the Mobile 
Bay drainage in Alabama, United States. 
Historically, the freshwater snail fauna of 
Mobile Bay basin was probably the most 
diverse in the world, followed by that of the 



Mekong River Nine families and about 118 
species were known at the turn of the century 
to occur in the Mobile Bay drainage. Several 
genera and many species were endemic, 
particularly in the Pleuroceridae. Recent 
surveys suggest at least 38 species are extinct 
(32 percenti; decline in species richness 
ranges between 33 percent and 84 percent in 
the main river systems. The richest fauna was 
in the Coosa River and this system has 
undergone the greatest decline (from 82 to 30 
species]. Almost all the snail species 
presumed extinct were members of the 
Pleuroceridae and grazed on plants growing 
on rocks in shallow oxygen-rich riffle and 
shoal zones. The system has 33 major hydro- 
electric dams and many smaller impound- 
ments, as well as locks and flood control 
structures. A combination of siUation behind 
dams and submergence of shallow water 
shoals has removed the snails' former 
habitat. Where habitat remains it has dimin- 
ished in area and become fragmented". 

The inland water living planet index 

An impression of the overall trend in a large 
sample of species for which indicators of 



Table 7.8 

Numbers of threatened 
freshwater fishes in 
selected countries 

Notes: These are the 20 
countries whose fish faunas 
have been evaluated 
completely, or nearly so, and 
which have the greatest 
number of globally 
threatened freshwater fish 
species. The estimates of 
total fish species present are 
all approximations. 

Source: Total species estimates from 
UNEP-WCMC database; threatenetj 
species (Jata from online Red List 
fitfpV/www redlist.org [accessed 
Marcfi 20021 



I^B 


Total species 


Threatened species 


% threatened 


USA 


822 


120 


15 


Mexico 


384 


82 


21 


Australia 


216 


27 


13 


South Africa 


94 


24 


26 


Croatia 


64 


22 


34 


Turkey 
Greece 


174 
98 


22 
19 


13 
19 


Madagascar 
Canada 


41 
177 


13 
12 


32 

7 


Papua New Guinea 
Romania 


195 
87 


11 
11 


6 
13 


Italy 
Bulgaria 


45 
72 


11 
11 


24 
15 


Hungary 

Spain 

Moldova 


79 
50 
82 


10 
10 
9 


13 
20 
11 


Portugal 
Sri Lanka 


28 
90 


9 
9 


32 
10 


Slovakia 


62 


9 


15 


Japan 


150 


9 


6 



188 WORLD ATLAS OF BIODIVERSITY 



Table 7.9 

Taxonomic distribution 
and status of threatened 
inland water vertebrates 

Notes: Family selection 
based on Table 7 6. 
Numbers refer to species 
(not subspecies and 
geographic populations] 
recorded in the Red List 
database as occurring in 
freshwater; only the birds 
and mammals have been 
comprehensively assessed 
lor species at risk. 

i Emydidae here includes 
batagurme turtles 
sometimes treated as a 
separate family (several of 
the species are primarily 
terrestrial not aquaticl. 

Source: Status categories from 2000 
Red List database, www.redlistorg 
iaccessed February 2002! 



Phylum and 


Family 


Common name 


Critically 


Endangered' 


'Vulnerable- 1 


class or order 




endangered' 




J 


Craniata - fishes 












Petromyzontiformes 




Lampreys 




1 


2 


Carchariniformes 




Ground stiarks 


1 


1 




Prisliformes 




Sawfish 


2 


3 




Myliobatiformes 




Rays 




5 


2 


Acipenseriformes 




Sturgeons 


6 


10 


8 


Alheriniformes 




Silversides 


6 


5 


31 


Beloniformes 




Needlefishes, sauries, etc. 


2 


3 


8 


Characiformes 




Characins 




1 


1 


Clupeiformes 




Herrings and anchovies 




2 


3 


Cypriniformes 




Carp, minnow, loaches 


42 


37 


117 


Cyprinodontiformes 




Rivulines, ((illifish, etc. 


18 


20 


26 


Gasterostiformes 




Sticl<lebacks 


1 






Ophidiiformes 




Pearlfishes, etc. 






6 


Osteoglossiformes 




Bonylongues 




1 




Perciformes 




Perches, etc. 


51 


25 


104 


Percopsiformes 




Trout-perches, cavefishes 


1 




3 


Salmoniformes 




Salmonids 


8 


8 


22 


Scorpaeniformes 




Gurnards, scorpionfishes, etc. 


2 




4 


Silunformes 




Catfishes 


8 


7 


22 


Synbranchiformes 




Swamp eels 




1 


j^_ 


Syngnathiformes 




Pipefishes, seahorses, etc. 


1 




M 


Craniata - Amphibia 










m 


Caudate 


Cryptobranchidae 


Giant salamanders and hellbenders 




1^^ 




Proteidae 


Ivtudpuppies and olm 






1 


Anura 


Pipidae 


Clawed frogs and pipid toads 






1 


Craniata - Reptilia 












Chelonia 


Carettochelidae 


Pig-nosed soft-shelled turtle 






1 




Chelidae 


Austro-american side-necked turtles 3 


4 


- 7 




Chelydridae 


Snapping turtles 






1 




Dermatemydidae 


Central American river turtle 




1 






Emydidae' 


Pond and river turtles 


13 


22 


16 



population change are available can be 
derived from the WWF living planet index. 
This method is designed to represent the 
change in the average species' in the sample 
from one five-year interval to the next, 
starting in 1970. The inland waters sample 
represents 194 species of mammals, birds, 
reptiles and fishes, and the index suggests a 
significant declining trend over the last three 
decades of the 20th century (Figure 7.31". 

The sample includes a large number of 
wetland and water margin species in addition 
to truly aquatic forms. The declining trend in 
the inland water sample is a little steeper 
than the equivalent marine index, and sub- 
stantially steeper than the terrestrial index. 



Assessment of the status of inland water 
ecosystems 

Indicators of habitat condition in river 
catchments 

While the living planet index methodology 
provides an indication of global trends in inland 
water biodiversity, an alternative approach 
aims to assess the overall condition of inland 
water ecosystems. In one approach to a global 
assessment' two high-order indicators of likely 
habitat condition in different river catchments 
were combined. First, a wilderness measure 
for each river catchment was calculated using 
the wilderness index methodology developed 
by the Australian Heritage Commission"" 
(see Chapter 4 and particularly Ivlap 4.51. This 



Inland water biodiversity 189 



■\V 



E Phylum and 


Family 


Common name 


'Critically 


'Endangered' 


'Vulnerable' 


■ class or order 






endangered' 








Kinosternidae 


Mud and musk turtles 






4 




Pelomedusidae 


Side-necked turtles 




2 


6 




Trionychidae 


Soft-shelled turtles 


4 


5 


6 


Crocodilia 


Alligatondae 


Caimans and alligators 


1 








Crocodylidae 


Crocodiles 


3 


2 


3 




Gavialidae 


Ghanal and false gharial 




1 




Craniata - Aves 












Anseriformes 


Dendrocygnidae 


Whistling-ducks 






1 




Anatidae 


Ducks, swans and geese 


5 


7 


12 


Charadriiformes 


Charadriidae 
Laridae 
Rhynchopldae 
Scolopacidae 


Plovers, etc. 

Gulls, terns, skua, auks 

Skimmers 

Curlews, etc. 


1 


1 
1 


2 
1 
1 


Ciconiiformes 


Ardeidae 


Herons, egrets 




3 


5 




Ciconiidae 


Storks 




3 


2 




Threskiornithidae 


Ibis, spoonbill 


2 


2 






Phoenicopteridae 


Flamingos 






2 


Gruiformes 


Heliornithidae 


Limpkin and sungrebes 






1 




Rallidae 


Rails, gallinules and coots 


1 


7 


6 


Pelicaniformes 


Pelecanidae 


Pelicans and shoebill 






1 


Passeriformes 


Cinciidae 


Dippers 






1 


Podicipediformes 


Podlcipedidae 


Grebes 


2 




2 


Craniata - Mammalia 












Artiodactyla 


HIppopotamldae 


Hippopotamus 






1 


Carnivora 


Mustelidae 
Phocidae 


Mustellds, otters 
Earless seals 




2 

1 


3 

1 


Cetacea 


Platanistidae 


River dolphins 


1 


2 


1 


Insectivora 


Soricidae 


Shrews 


2 


1 






Talpidae 


Moles and desmans 






2 




Tenrecidae 


Tenrecs and otter shrews 




4 




Rodentia 


Murldae 


Mice, voles, etc. 


1 


5 


3 


Sirenia 


Trichechldae 


Manatees 






3 



Figure 7.4 

River basin richness and 

vulnerability 

Notes: Eacti symbol 
represents a river basin 
scored on fish family 
richness and basin 
vulnerability: the darker 
symbols score high on both 
counts and may be regarded 
as high priority. See Map 7,5 
and Table 7,10. 

Source: WCMC' 



provides a measure of the spatial extent of 
human impacts in the land area of the 
catchment as a whole and does not directly 
reflect the condition of riverine ecosystems. 
Secondly, a national water resource vulnera- 
bility index IWRVIl" developed on the basis of 
three water resource stress indices (reliability, 
use-to-resource and coping capacity) was 
resolved at catchment level Iby measuring that 
proportion of each catchment that lies within 
any given country and weighting this proportion 
by the national WRVIl. This provides an indirect 
measure of vulnerability tor each catchment. 
By normalizing and combining these two 
measures, a single value representing vulner- 
ability or stress level for each major river 



30 



20 



S 10 



-10 



-20 
-0 

























4 


t • • 

•• 


• 














rf> 


r • 


%t 


» 










1 

1 














1 






.8 -0 


6 -0 


4 -0 


2 ( 


) 


2 


4 


6 0.8 



Overall vulnerability 



190 WORLD ATLAS OF BIODIVERSITY 



Map 7.5 

Priority river basins 

A possible scheme for 
prioritizing river basins at 
global level Major 
catchment basins are 
assessed for fish diversity 
lat family level) and for 
vulnerability (a combined 
indication of disturbance 
and potential water stress]. 
Systems with both high 
diversity and vulnerability 
are proposed as high 
priority for investment and 
management action; those 
with low diversity and little 
disturbance are lower 
priority. See text and source. 

Source: WCMC'. 



Table 7.10 

Thirty high-priority river 

basins 



Notes: These are 30 river 
basins that support high 
biodiversity (assessed as fish 
family richness! and are most 
vulnerable to future 
pressures (have a low 
wilderness score, and are 
high on the water resource 
vulnerability index). See text 
for further explanation and 
Figure 7.4. Basins are listed 
in alphabetical sequence. 



Priority 




High 

tuledium 

Low 



catchment can be calculated. Overall, the 
pattern mapped agrees well with what might 
intuitively be expected, with few evident 
anomalies. Globally, the most stressed catch- 
ments are to be found in South Asia (the Indian 
subcontinent), the Middle East and western 
and northcentral Europe. The least stressed 



^^^^^^^^^^I^^I^^^^^^H 


Ca 


Irrawaddy 


Nile 


Senegal 


Cauvery 


Krishna 


Pahang 


Sittang 


Ctiao Phraya 


Ma 


Parana 


Song Hong (Red) 


Gambia 


Magdalena 


Parnaiba 


Tapti 


Ganges- 


Mahanadi 


Penner 


Tembesi-Hari 


Brahmaputra 


Mekong 


Peral< 


Uruguay 


Godavari 


Narmada 


Salween 


Volta 


Indus 


Niger 


Sao Francisco 





are those in the northwestern part of North 
America. Further refinements of this analy- 
sis would involve applying water resource 
vulnerability measures for individual catch- 
ments" rather than to countries and incor- 
porating measures of direct impacts on inland 
waters, in particular water quality and the 
number and kind of dams. 

The number of freshwater fish families 
present in each basin can be calculated from 
the family density surface (Map 7.1) in order to 
provide an indication of biodiversity value. 
Plotting family number against vulnerability 
allows the basins with high diversity and high 
vulnerability to be identified, and these can 
reasonably be regarded as global priorities 
for management intervention designed to 
minimize biodiversity loss (Figure 1 A, Table 
7.10, Map 7.5). 



Inland water biodiversity 191 




Habitat condition in lakes 
One semi-quantitative study attempted to 
evaluate the ctianging condition of freshwater 
lakes during the last three decades of the 20th 
century^'. A baseline was provided by Project 
Aqua, a project initiated by the Societas 
Internationalls Limnologlae In 1959, which 
collated and later published" Information 
provided by national and regional specialists in 
relation to more than 600 water bodies. Many 
of these lakes were treated in later Infor- 
mation sources relating to the 1980s and 
1990s", and In some 93 cases it was possible 
to make an assessment that, although 
Imprecise, is likely to be indicative of changing 
conditions. Each lake was scored according to 
whether Its condition appeared to have 
deteriorated (or Impacts Increased], or to have 
Improved, or whether no change (or no new 



information) was reported. Improvement was 
reported In a very small number of lakes, but 
the overwhelming trend was for a deterior- 
ation In conditions (see Figure 7.5|. 



Figure 7.5 

Changes in condition of a 
sample of freshwater lakes 
between 1950s and 1980s 

Note: Number following area 
name is number of lakes In 
sample. 

Source: various sources 



Better 



Not known 



Worse 



100 
% 
80 

60 

20 






Africa 
(201 



Asia 
(241 



Central & South Australia 
America (91 (3| 



Europe 
(371 



TOTAL 
(931 



192 WORLD ATLAS OF BIODIVERSITY 



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conservation, preservation and management, pp. 129-169. Chapman and Hall, New York 
and London. 

41 Stiassny, M. 1996. An overview of freshwater biodiversity with some lessons learned from 
African fishes. Fisheries 2'\{9]: 7-13. 

42 Reinthal, P.N. and Stiassny, M.L.J. 1991. The freshwater fishes of Madagascar: A study of 
an endangered fauna with recommendations for a conservation strategy. Conservation 
Biology 5[2]: 231-243. 

43 Kirchhofer, A. and Hefti, D. (eds) 1996. Conservation of endangered freshwater fish in 
Europe. Advances in Life Sciences. Birkhauser Verlag, Basel. 



194 WORLD ATLAS OF BIODIVERSITY 



44 HiUon-Taylor. C. Icompiler] 2000. 2000 lUCN Red List of threatened species. lUCN-the 
World Conservation Union, Gland and Cambridge. Also available online at 
http://www.redlist.org/ (accessed April 20021. 

45 Bogan, A.E., Pierson, J.M. and Hartfield, P. 1995. Decline in the freshwater gastropod 
fauna in the Mobile Bay basin. In: LaRoe, E.T. et al. (edsl. Our living resources: A report to 
the nation on the distribution, abundance, and health of U.S. plants, animals, and 
ecosystems, pp. 240-252. US Department of the Interior, National Biological Service, 
Washington DC. 

46 Loh, J. led.l 2000. Living planet report 2000. WWF - World Wide Fund for Nature, Gland. 

47 Lesslie, R., in litt., 30 May 1998. 

48 Lesslie, R. and Maslen, M., 1995. National wilderness inventory, handbool< and 
procedures, content and usage. 2nd edition. Commonwealth Government Printer, 
Canberra. 

49 Raskin, P. et al. 1997. Water futures: Assessment of long-range patterns and problems. 
Background Report No. 3 of Comprehensive Assessment of the Freshwater Resources of 
the World. Stockholm Environment Institute. 

50 Alcamo, J., Henrichs, T. and Rosch, T. 2000. World water in 2025. Global modeling and 
scenario analysis for the World Commission on Water for the 21st Century. Kassel World 
Water Series. Report No. 2. University of Kassel, Centre for Environmental Systems 
Research. 

51 Loh, J. etal. 1998. Living planet report 1998. WWF-World Wide Fund for Nature, Gland. 

52 Luther, H. and Rzoska, J. 1971. Project Aqua: A source book of inland waters proposed for 
conservation. IBP Handbook No. 21. lUCN Occasional Paper No. 2. International Biological 
Programme. Blackwell Scientific Publications, Oxford and Edinburgh. 

53 For example, Scott, D.A. led.) 1989. A directory of Asian wetlands. lUCN, Gland and 
Cambridge; Scott, D.A. and Carbonell, M. 1986. A directory of neotropical wetlands. lUCN, 
Gland and Cambridge; International Lake Environment Committee (ILEC), World lakes 
database, available online at http://www.ilec.or.jp/database/database.html 

(accessed March 2GQ2I. 

54 Anon 1978. Centre for Natural Resources, Energy and Transport of the Department of 
Economic and Social Affairs, United Nations. Register of International Rivers. Water 
Supply and Management!: 1-58. (Special issue). Pergamon Press. 

55 Shiklomanov, I. A. 1998. World water resources at the beginning of the XXlst century. 
Report to UNESCO (Division of Water Sciences). Available online in summary form at 
http://webworld.unesco.org/water/ihp/db/shiklomanov/summary/html/summary.html 
(accessed March 2002). 

56 Martens, K., Goddeeris, B. and Coulter, G. (eds) 1994. Speciation in ancient lakes. Archiv 
fur l-lydrobiologie. Ergebnisse der Limnologie 44. 

57 Hutchinson, G.E. 1993. A treatise on limnology. Vol. IV. The zoobenthos. John Wiley and 
Sons, Inc., New York. 

58 Eschmeyer, W.N. ef al. 1 998. A catalog of the species of fishes. Vols 1 -3. California 
Academy of Sciences, San Francisco. Available online in searchable format at 
http://www.calacademy.org/research/ichthyology/catalog/fishcatsearch.html [accessed 
January 20011. 

59 Welcomme, R.L. (compiler) 1988. International introductions of inland aquatic species. FAO 
Fisheries Technical Paper No. 294. Food and Agriculture Organization, Rome. 

60 Berra, TM. 2001. Freshwater fish distribution. Academic Press, San Diego and London. 

61 Johnson, T.C. et al. 1996. Late Pleistocene desiccation of Lake Victoria and rapid evolution 
of cichlid fishes. Soence 273: 1091-1093. 

62 Snoeks, J. 2000. How well known is the ichthyofauna of the large East African Lakes? 
Advances in Ecological Research 31 : 17-38. 



1 



Global biodiversity: responding to change 



8 



Global biodiversity: 
responding to change 



CONCERN OVER THE RATE OF CHANGE IN THE BIOSPHERE, particularly In populations 
of larger animals and In the naturalness of landscapes, dates primarily from the turn 
of the 20th century. Although some early measures were taken, prompted by 
pioneering conservation organizations, concerted International effort did not develop until 
mid-century. Since this period, actions have tended to focus on conservation of Individual 
species or of large areas of habitat, as national parks and other protected areas. 

During the 1970s several International agreements aimed at conserving v/ild species and 
habitats were agreed and entered Into force. During the 1980s the word 'biodiversity' was 
coined, and a new paradigm formulated, aiming to Integrate biodiversity conservation with 
sustainable human development. In 1992 the pivotal Earth Summit was held In Rio, 
culminating In agreement on the Convention on Biological Diversity, and Agenda 21 - a plan 
of action for sustainable human development. 

After a decade of planning and Implementation, the status of some species and the 
condition of some ecosystems have Improved. In some areas, this has been achieved by 
restoration of degraded ecosystems, an approach likely to be of Increasing Importance In the 
future. However, given future trends In human population growth and development. It seems 
probable that pressures on biodiversity will continue to Intensify In coming decades. In many 
parts of the world, the most significant challenge to conservation will be to minimize losses 
of biodiversity while improving the livelihood of human populations, particularly those 
experiencing severe poverty. 



BIODIVERSITY CHANGE 

Change is a donninant theme in the biosphere; 
species have diversified and become extinct 
throughout the history of life, and habitats too 
have expanded and declined, along w/ith 
changes in community composition, climate 
and landforms. However, the recent rate of 
change in global biodiversity appears higher 
than that prevailing over most periods of 
geological time. Earlier chapters have indi- 
cated how in well-assessed groups of 
organisms, such as birds and mammals, a 
significant proportion of species now appears 
to be threatened with global extinction. 
Countless other species exist in reduced 
numbers and as fragmented populations, 
many of which are threatened with extinction 
at national or more local scale. The rate of 
extinction over the past few centuries, so far 



as this can be reliably estimated, appears to 
be much higher than the average background 
rate estimated from the fossil record. 
Whatever proximate causes may be impli- 
cated, the increasing numbers and material 
aspirations of the human species, the in- 
creasing burden of waste and continuing 
ineguitles in the distribution of wealth and 
resources, together appear to drive most con- 
temporary biodiversity loss. 

Despite abundant evidence of change in 
biodiversity, often Involving radical modifi- 
cation of landcover or water bodies, much 
of the relevant information on the status of 
species or populations is qualitative or anec- 
dotal in nature. At the level of individual 
species, a qualitative assessment of trends In 
numbers or range may be adequate evidence 
of the need for management intervention, or a 



196 WORLD ATLAS OF BIODIVERSITY 



Concern about the decline 
of species became the 
focus of sustained 
international attention only 
in the last 30 years. 



guide to its effectiveness, but it remains diffi- 
cult to develop a comprehensive and quantit- 
ative view of global species trends. The 
relatively few large-scale population monitor- 
ing programs that exist have tended to 
concentrate on marine fishery stocks, on par- 
ticular groups le.g. farmland birdsl or the 
larger threatened species of animals. The res- 
ulting series of data are not always directly 
comparable, and often not amenable to pres- 
entation in a manner appropriate for assisting 
global and regional planning. Much recent 
discussion' has focused on the design of 
biodiversity indicators that could serve this 
purpose, but few operational systems yet exist. 
This chapter first provides an overview of 
current approaches to conservation, focusing 




on species, areas and ecosystems, and the 
particular role of protected areas, as well as 
the potential for ecological restoration. The 
international dimension is then considered, 
with specific reference to multilateral environ- 
mental agreements and conventions relating 
to biodiversity. Finally, changes in global 
biodiversity that might be anticipated in future 
are explored through description of recently 
developed scenarios. Such changes highlight 
the challenges facing current and future 
efforts aimed at the conservation and sus- 
tainable use of global biodiversity. 



RESPONSES TO BIODIVERSITY CHANGE 

Although concern about the decline of species 
and loss of undisturbed habitats arose within 
developed industrialized societies around the 
start of the 20th century, they became subject 
to sustained international attention only during 
the 1970s. In general, major sectors that used 
biodiversity, such as forestry and fisheries, 
failed to incorporate consideration of bio- 
diversity into their planning and regulations. 
The issue was widely perceived as marginal to 
other concerns, and conservation of nature 
was commonly seen as an impediment to 
human development. 

The past three decades have been marked 
by the emergence of concerted responses to 
the crisis in biodiversity at all levels from the 
local to the global. Civil society, largely in the 
form of a hugely diverse and increasingly 
sophisticated network of non-governmental 
organizations INGOsI, has undoubtedly been 
the driving force for most of this change (see 
Box 8.11. Response to pressure from civil 
society can be seen at both governmental and 
intergovernmental levels. 

Much of the progress that has been made in 
recent decades can be attributed to the vision 
articulated in the World Conservation Strategy 
by the International Union for Conservation of 
Nature and Natural Resources, the United 
Nations Environment Programme and the 
World Wildlife Fund In 19801 This document 
helped set the conservation agenda during the 
period leading up to the UN Conference on 
Environment and Development (the Earth 
Summit] in Rio de Janeiro in 1992. It em- 
phasized conservation for development, and 
embraced the notion of the sustainable use of 
natural resources as a means of achieving 
this. Maintenance of biological diversity as part 
of a functioning biosphere was presented as a 
fundamental prerequisite of sustainable hu- 
man development, not an impediment to It. On 
the ground, this was manifested in increasing 
numbers of integrated conservation and de- 
velopment projects In developing countries. 
The Earth Summit was undoubtedly the major 
environmental milestone of the 1990s, and 
from it emerged the Convention on Biological 
Diversity, which entered Into force in 
December 1993. 



Global biodiversity: responding to change 



197 



At the start of the 21st century concerns 
about loss of species and habitats have 
intensified. Additional issues have emerged, 
such as climate change, with its inevitable 
effects on coastal, montane and other eco- 
systems, and the ability of humans directly to 
modify gene structure and expression in living 
organisms, with debate over the risks involved 
when genetically modified organisms are re- 
leased into the biosphere. 

Protecting species 

Maintaining biodiversity requires that viable 
populations of diverse organisms are main- 
tained in viable ecosystems. This may involve 
activities carried out on site or off site. While 
the latter lex situ] has an important role, as 
with the seed banks and germplasm collec- 
tions for agricultural plants, it can only have 
very limited application. The former {in situ] is 
essential for the vast majority of organisms, 
and ecosystem conservation selt-evidently 
depends entirely on maintaining environ- 
ments in which communities of organisms 
can interact and evolve. 

Because the vast majority of the worlds 
biodiversity exists within the territorial boun- 
daries established by nations, most conser- 
vation action is carried out within the policy 
and legal systems established by national 
governments lor in a few instances, by 
regional or provincial governments). A wide 
range of national measures exists, varying to 
some extent from country to country depen- 
ding on the social, political and economic 
environment. Despite this variety, most such 
measures involve regulation of the taking, 
possession and trade in a set of species, 
typically named and listed in legislation. The 
protection of wild fauna has generally been 
given much more attention than the pro- 
tection of wild flora, but most modern 
examples of species-specific legislation cover 
both flora and fauna, as exemplified by the US 
Endangered Species Act, passed in 1973. 

The species approach has been supported 
at the international level by the lUCN (World 
Conservation Union] Red Data Book program 
of activities and information tools. The focus is 
on documenting and disseminating infor- 
mation on species at risk of extinction, and on 



Box 8.1 Pioneering NGOs in biodiversity conservation 

Some of tfie earliest establisfied non-governmental organizations remain influential today. 
The Sierra Club was founded in the United States in 1392, with a focus on maintaining 
wilderness areas in North America. The Society for the Preservation of the Wild Fauna of the 
Empire - now Fauna and Flora International IFFI) - was founded in 1903 in order to safeguard 
southern Africa's declining large mammal populations and is now worldwide in scope. 
Initiated in 1962, Operation Oryx was a landmark project of FFI, and averted extinction of the 
Arabian oryx by means of a captive breeding and reintroduction program. The International 
Council for Bird Preservation - now BirdLife International IBLII - was founded in 1922. It has 
played a lead role in advancing standards for conservation information and has promoted 
grassroots involvement in conservation through its worldwide membership structure. 

The International Union for the Protection of Nature - now lUCN-The World Conservation 
Union - was created in Switzerland in 1948. Sir Peter Scott became the chair of two of 
lUCN's key commissions: on protected areas and on species survival (SSCI, and later of FFI, 
and in 1966 he initiated the Red Data Book approach to documenting species at risk. The 
World Wildlife Fund (WWFI - now known as WWF-World Wide Fund for Nature outside North 
America - was designed originally to generate public contributions to support lUCN's work, 
and was launched in 1961 with a campaign on black rhino. These two NGOs have been 
driving forces in global policy development. 



detailing the management steps - often in- 
cluding preservation of important areas or 
reduction in exploitation levels - needed to 
reduce that risk. In some cases, targets and 
indicators of progress may also be defined. 

In practice, conservation attention has 
tended to focus on species that are large, 
charismatic and possibly also ecologically 
important or highly threatened, or both. 
The tiger Panthera tigris, Arabian oryx Oryx 
[eucoryx, white rhinoceros Ceratotherium 
simum and the blue whale Balaenoptera 
musculus are familiar mammalian examples. 
Considerable success has been achieved with 
the species named above and the many others 
(mainly animals) subject to similar conser- 
vation efforts. However, the species approach 
could never be extended to cover more than a 
minute fraction of the approximately 300 000 
larger organisms [plants and vertebrates) in 
the biosphere, let alone the other several 
million that probably exist. In fact, the primary 
benefit may be that large organisms, and 
terrestrial ve'^tebrates in particular, generally 
require large areas of suitable habitat, and if 
such areas can be managed to minimize risk, 
other species may be safeguarded. 



198 WORLD ATLAS OF BIODIVERSITY 



Figure 8.1 

Development of the global 
network of protected 
areas 

Source: UNEP-WCMC database, 
maintainted in collaboration with lUCN 
World Commission on Protected Areas 

2 500 



2 000 



1 500 



1 000 



500 



Protecting areas 

The regulation of access to and use of 
particular areas has always been seen as 
complementary to the focus on individual 
species. Protected areas are in many ways the 
most important form of legislative measure 
for the conservation of biodiversity. Whereas 
the initial purpose of many such areas was to 
protect spectacular scenery and provide 
recreational facilities, the concept evolved to 
encompass habitats of threatened species 
and ecosystems rich in biodiversity. 

By the beginning of the 20th century many 
countries had either already established 
protected areas or were contemplating doing 
so. The concept, however, was slow to be put 
into practice and it was not until the 19^0s 
that protected areas were being established 
in any significant number The rate at which 
land was being incorporated into the system 
did not increase markedly until the early 
1960s. The first World Parks Congress held in 
Seattle, in the United States, in 1962 was an 
important stimulus for the increase. This 
meeting signified the emergence of the mod- 
ern protected area network with over 80 
percent of the worlds protected areas being 
established since thenl The cumulative and 
periodic growth of world protected areas are 
plotted in Figure 8.1 (the creation of the 
Greenland National Park in 1974, covering 
some 97 million hectares, and the Great 



BARS: 
Number of sites 

lover 1 000 ha. lUCN categories I to VII 
gazetted over a five-year period 
(left-hand scale! 



LINE: 

Cumulative protected area 

[right-hand scale! 




UOOO 



12 000 



- 10 000 



-8 000 



6 000 



4 000 



2 000 



1970 



1975 



1980 



1985 



1990 



1995 



2000 



Barrier Reef Marine Park in the 1980s, 
extending over around 34 million hectares, 
have had marked individual effects on the 
global totals). The location of protected areas 
in the lUCN/WCPA (World Commission on 
Protected Areas! categories l-VI greater than 
100 000 hectares in extent is shown in 
Map 8.1. 

Many protected areas, particularly those 
where secure funding allows management to 
be maintained, are effective in conserving 
species, habitats and landscapes of value. 
An extensive questionnaire study involving 
93 protected areas in the tropics, in 22 
countries, showed that areas where basic 
management activities are in place tend to 
avoid wholesale land clearance (but often still 
suffer some degree of disturbance, from 
hunting or logging, for example!'. An inte- 
grated measure of their effectiveness is not 
yet available, and it may be that firm manage- 
ment within park boundaries can have the 
effect of increasing pressure on outside land 
that is less controlled; on the other hand, 
many protected areas are badly under- 
resourced, reducing their effectiveness. 

Wise management of areas outside the 
protected areas network, by means of 
planning controls and voluntary agreements 
and by incorporating conservation principles 
in landuse planning, also plays an essential 
role in conservation of biodiversity. Indeed, in 
many countries, management of landuse 
outside the national network of protected 
areas - in agricultural landscapes, for 
example - will play as important a role in the 
maintenance of national biodiversity as will 
the network itself. 

Maintaining ecosystems 

Increasingly in recent years a more holistic 
approach has emerged in which ecosystems 
themselves have become the focus of con- 
servation efforts. Many would argue that the 
primary goal of conservation action is the 
long-term maintenance of ecosystem pro- 
cesses at the global scale and over 
foreseeable human generations. This is 
essentially equivalent to achieving sustain- 
ability in human development and use of 
biodiversity. The Convention on Biological 



Global biodiversity: responding to change 



199 



Diversity adopted 'the ecosystem approach' 
as the guiding framework for actions taken in 
pursuit of its goals and, in elaborating on the 
meaning of this term, stressed the need to 
conserve ecosystem structure and function 
in order to maintain ecosystem services. 
However, making an assessment of the 
organization, vigor and resilience of eco- 
systems - the key components of ecosystem 
health' - is fraught with immense practical 
difficulties. Much conservation activity there- 
fore is guite properly based on a strong form 
of the precautionary principle iBox 8.2), and 
focuses on maintaining so far as possible 
the prominent elements of ecosystems, i.e. 
species and populations and their physical 
environment, in anticipation that the system 
will thus be perpetuated. 



benefit will accrue from ensuring the integrity 
of areas of higher biodiversity value than 
areas of lesser value. 

One early area-based approach identified 
some 12 'megadiversity' countries, which 
between them include a large proportion of 
global biodiversity in selected major groups^ 



F 



Box 8.2 The precautionary principle 

A concise definition of the precautionary principle is provided by Principle 15 of the Rio 
Declaration, made at the 1992 Earth Summit: 

'In order to protect the environment, the precautionary approach shall be widely applied by 
States according to their capabilities. Where there are threats of serious or irreversible 
damage, lack of full scientific certainty shall not be used as a reason for postponing cost- 
effective measures to prevent environmental degradation.' 



Defining priority areas for protection 

Management action aimed at biosphere con- 
servation demands financial resources, but 
these are limited, while human numbers and 
impacts continue to increase. This implies 
that choices must be made between possible 
actions, whether by design or by default. 

Rational and informed decision-making 
should seek to increase the efficiency with 
which conservation funds are used. Several- 
studies have been undertaken that aim to 
identify and sometimes to rank areas of high 
biodiversity value. The approach is generally 
based on the premise that more is better'. 
That is, an area with greater biodiversity value 
is more worth conserving than an area with 
lower value. At its simplest biodiversity may 
be eguated with species number overall or 
per unit area, but other selection criteria 
are possible. Higher priority may be accorded 
to an area with populations of threatened 
species, or one rich in endemic species 
lespecially if there are endemics in several 
different groups), or in species of commercial 
or cultural importance; or one which is par- 
ticularly representative of an ecosystem 
(perhaps one that is widely degraded else- 
where). While the results of such assess- 
ments are usually of considerable biological 
interest, they may also have direct application 
as a basis for choice between different 
courses of action, on the grounds that greater 



Another early study delineated 18 endemic- 
rich botanical 'hotspots', which between them 
included around 20 percent of the world's 
known plant species in less than 1 percent of 
the land surface, and were also undergoing 
rapid habitat conversion". A strength of the 
former approach is that it is focused at country 
level, and it is at this administrative level that 
most conservation action is undertaken; a 
weakness is that by evaluating species rich- 
ness alone it was not able to address unigue- 
ness, and adjacent countries rich in species 
are likely to have many species in common. A 
strength of the latter approach is that it focus- 
ed on areas rich in restricted-range endemic 
species and such areas by definition make a 
large contribution to global biodiversity. 

The country-based approach has been 
extended in a study" using a database of 
estimates of richness and endemism in land 
vertebrates and vascular plants for all coun- 
tries of the world (Appendix 5). Indices 
of overall diversity (weighting richness and 
endemism equally) and diversity adjusted 
for country area have been produced. Within 
the relatively wide margin of error associated 
with species inventory (see Chapter 2), these 
indices can yield a useful view of variation in 
diversity in geopolitical terms or. If the data 
are treated as geographic samples, a view of 
general global variation In diversity (see Map 



200 WORLD ATLAS OF BIODIVERSITY 



Map 8.1 

World protected areas 

An overview of the world's 
surface nominally subject 
to protection and 
appropriate management. 
Tfie location of protected 
areas in lUCN/WCPA 
categories l-VI greater than 
100 000 hectares in area is 
shown, represented by a 
point symbol. Protected 
areas greater than 1 million 
hectares in extent are 
represented by polygons 
instead of point symbols 
whenever boundary data 
are available. 



Source; UNEP-WCMC database (data 
extracted Marcti 20021, maintained in 
collaboration witti lUCN World 
Commission on Protected Areas. 








Protected areas 

^^^H > 1 million hectares 
> 100 000 hectares 



5. A). The hotspots approach has been further 
developed by Conservation International in a 
recent synthesis, and adopted as the basis for 
its conservation planning and action'. Twenty- 
five terrestrial hotspot regions were identi- 
fied, combining high species endemism, 
particularly among plants, and high rates of 
habitat loss, with at least 70 percent of origi- 
nal natural vegetation having been lost. Coral 
reef areas of the world have now also been 
evaluated in this way [see Chapter 6 and Map 
6.1), and freshwater ecosystems are currently 
being evaluated. The term 'hotspof is now 
often applied loosely to any area that has a 
concentration of diversity, whether by a 
measure of simple richness, or endemism or 
number of threatened species, and whether 
undergoing habitat conversion or not. 

Of studies based on biogeographic rather 



^^hh'i^^'^ 



■'U^^i 




than geopolitical areas, the Centres of Plant 
Diversity project'" remains one of the largest. 
It relied heavily on extensive consultation 
among botanists to identity several hundred 
important sites worldwide, defined semi- 
quantitatively on the basis of a general 
combination of richness and endemism (Map 
8.2|. A broadly similar but less structured 
approach, also based on expert knowledge, 
has been taken to identify 43 areas of special 
importance for amphibian diversity, again 
using species richness and endemism as 
criteria" (Map 8.31. A preliminary selection of 
areas of importance for inland water bio- 
diversity has also been made on the basis of 
expert knowledge of richness and endemism 
among fishes, mollusks and crustaceans [see 
Chapter 7, and Maps 7.2-7.-41. 

However, the most systematic and com- 



Global biodiversity: responding to change 






::-^:':c^ 



^. 




%€/^. 







plete global level assessment to date has 
involved bird species. Birds are by far the 
best-known major group of organisms on the 
planet, with a relative wealth of distribution 
and population data available, and global 
analyses by BirdLife International and its 
partners have set a standard yet to be 
matched for other taxonomic groups. The dis- 
tributions of all restricted-range bird species 
(defined as those in which the area encom- 
passing all distribution records is less than 
50 000 km1, amounting to 25 percent of all 
birds, have been mapped in digital format. 
The co-occurrence of restricted-range species 
defines a set of 218 endemic bird areas 
lEBAs)''; these are shown, ranked in three 
categories according to biodiversity impor- 
tance, in Map 8.4. Importance' here takes 
account of the number of restricted-range 




species in the EBA and the number of EBAs in 
which they are present, taxonomic unique- 
ness and EBA area. 

There are a number of limitations to area- 
based methods for establishing priorities 
among possible conservation actions. First, 
there is no single unequivocal way of com- 
paring value in different categories: how 
many vulnerable species is a single critically 
endangered species worth? Or how many 
endemic beetles is a single endemic bird 
worth? Second, information is always incom- 
plete, in that it never covers all taxa at all sites 
of interest. Resolving the former requires 
more or less arbitrary assigning of value. 
Attempts to resolve the latter entail the 
search for indicators, that is groups of species 
or other variables that can act as surrogates 
for wider measures of biodiversity. 



202 WORLD ATLAS OF BIODIVERSITY 



Map 8.2 

Centers of plant diversity 

This map stiows the 
location of the sites and 
areas identified as 
important centers of plant 
diversity at regional and 
global levels, using expert 
knovi/ledge. and mixed 
criteria emphasizing 
species richness and 
endemism. See the three- 
volume source cited below 
tor further details and 
extended documentation 

Source; WWF and IUCn'° 



Plant diversity 

^^H • Areas and centers 



The search for biological indicators of this 
kind has generated a great deal of research 
and discussion. Findings to date have gen- 
erally been equivocal, and depend in part on 
spatial scale. In general, though, it seems that 
at coarse scales there may be quite close 
agreement between different taxa so that, for 
example, many EBAs (which may be up to 
600 000 km- in extenti also hold significant 
numbers of other restricted-range species". 
At this scale, birds may serve as indicators of 
high biodiversity value more generally In 
other instances, and perhaps more generally 
at finer resolution, the relationship appears to 
break down. Studies in areas as disparate as 
North America, South Africa", Cameroon" 
and the British Isles"" indicate that areas 
important for rare species in different groups 
often do not coincide [and these may be 





■ ■^. 



negatively correlated with areas of high 
species richness). Also, richness levels in any 
one group do not necessarily provide an 
indication of species richness in others. 

A slightly different approach that avoids 
some of the limitations faced by methods 
based on species-specific information is to 
identify areas on the basis of their biological 
communities, and on biogeographic criteria, 
with reduced emphasis on richness or 
endemism in particular taxa, and none on 
current habitat condition. The conservation 
organization WWF's 'ecoregions' system, 
mentioned in Chapters 5-7 above, is the 
principal example of this approach, and 
currently covers much of the world's marine 
and inland waters, together with the entire 
land surface. It has been adopted as the basis 
for conservation planning and action by WWF. 



Global biodiversity: responding to change 



'^C^ 




Whilst a number of different groups and 
organizations have explored the use of spatial 
biodiversity information in suggesting con- 
servation priorities, these generally remain 
disconnected and to some extent duplicative". 
Recent developments suggest that a more 
integrated approach is feasible. Firstly, the 
conceptual and methodological framework of 
a more systematic approach to priority setting 
has been elaborated" (see Box 8.3). Secondly, 
sustained effort to map species distribution in 
a globally consistent manner is beginning to 
allovi/ robust analysis of large-scale multi- 
taxon patterns of biodiversity" ". Combining a 
systematic approach with a geographically 
extensive set of spatial biodiversity data 
has the potential to allow priorities to be 
defined in a way that is flexible, transparent, 
and robust. 



Box 8.3 Systematic conservation planning 

WL. 

Key stages in systematic planning for conservation 
by means of adding to or modifying protected area 
systems: 

1. Compile data on the biodiversity of the planning 
region. 

2. Identify conservation goals for the planning 
region. 

3. Review existing conservation areas. 

4. Select additional conservation areas. 

5. Implement consen/ation actions. 

6. Maintain the required values of conservation 
areas. 

Source: Margules and Pressey . 



204 WORLD ATLAS OF BIODIVERSITY 



Map 8.3 

Major areas of amphibian 

diversity 

The location of areas 
identified on the basis of 
expert knowledge as 
globally important for 
amphibian diversity, using 
data on species richness 
and endemism. Areas are 
here shown classified in 
five categories according to 
species richness. See 
source cited below for 
further details. 

Source: Adapted from Duellman 




37- 76 
21 -36 
4-20 



It should, however, be emphasized that 
many priority-setting exercises are likely to 
remain only theoretical. This is because the 
opportunities to establish entirely new terres- 
trial protected areas, or redesign existing 
protected area networks in the light of priorities 
identified, are in general extremely limited. The 
designation and management of protected 
areas, as so much other conservation action, is 
generally driven and constrained far more by 
socio-political and economic considerations 
than it is by conservation biology. 

Management of inland waters 
To a greater extent than is typical for terrestrial 
ecosystems, inland waters are often subject to 
management and use decisions made within 
many different sectors, including agriculture 
and forestry, navigation, public utility supply 








'^^- t 




Ipower, water and waste disposal! and recre- 
ation. Most inland waters or their catchments 
also typically intersect several subnational 
administration units Icounties, provinces, etc.) 
Although it has been recognized for some time 
that the catchment basin is the fundamental 
unit within which management must be 
formulated, it has proved difficult to reconcile 
the many different interests concerned and 
coordinate action. This is particularly problem- 
atic because sources of adverse impacts on 
such systems often originate far distant from 
where they are felt: thus pollutants and other 
inputs that enter the top reaches of a river 
system may have an impact in all downstream 
parts of the system as far as the river mouth 
and beyond. Similarly, water abstraction may 
have little effect in upper reaches, but be a 
serious constraint downstream. 



Global biodiversity: responding to change 205 




-^, 



X.-, 






K>T\ 



>;iv#> 



\ 



t^ 



Biodiversity maintenance and conser- 
vation of inland w/ater capture fisheries lother 
than Sonne lucrative sports fisheries! have 
not ranked highly among these competing 
interests, so that it has been difficult to impose 
catchment-vifide regulations or remedial 
measures for their benefit. This problem is 
exacerbated by the fact that it is often difficult 
to pinpoint a distant source of a problem or to 
unequivocally demonstrate that actions in one 
place are having an adverse effect somewhere 
else (e.g. to convince farmers that application 
of large doses of nitrogenous fertilizer on up- 
stream agricultural land is causing deleter- 
ious eutrophication of estuarine wetlands!. 

Although they cannot solve catchment-wide 
problems, inland water protected areas may 
play a valuable role in safeguarding particular 
sites or populations of species from immediate 



-^ 



threats. Protection may be most effective 
where sites are relatively small and thus 
manageable, and have a relatively low level of 
allochthonous inputs. Wetlands, with their 
often abundant and highly conspicuous avi- 
fauna, have in general received most attention 
in this regard. 

Notable wetland protected areas include 
the Moremi game reserve in the Okavango 
delta (Botswana!, Camargue national reserve 
(France!, Keoladeo (Bharatpur! national park 
(India), Dohana national park (Spain! and 
Everglades national park (United States!. 
Inland water ecosystems are unusual in that 
an international convention is dedicated 
specifically to them: the Convention on 
Wetlands of International Importance espec- 
ially as Waterfowl Habitat (the Ramsar 
Convention, see belowl. 



206 WORLD ATLAS OF BIODIVERSITY 



Map 8.^ 

Endemic bird areas 

More than one quarter 
12 5611 of ttie world's bird 
species, including more 
than 70% of the threatened 
birds, have a range 
restricted to less than 
50 000 km'. Virtually all 
occur within the 218 
endemic bird areas lEBAsI 
defined by BirdLife 
International'-. 
The world's EBAs are 
shown on this map 
categorized 1. 2 or 3 
according to increasing 
biodiversity importance 
(based on the number of 
restricted range species, 
whether shared between 
EBAs, taxonomic 
uniqueness and EBA sizel 

Source: Data provided by BirdLife 
International, and see Stattersfield . 



Category 



BSBJUS 



Transboundary inland waters 

Waters that delineate or cross international 
boundaries present a special class of 
management issue. Such waters and the 
living resources they contain are shared by 
one or more countries, and require positive 
international collaboration for effective use 
and management. 

Available water in any given country within 
an international basin lor other administrative 
unit within a basin more generaltyl can be 
divided into endogenous, i.e. locally generated 
runoff available in national aquifers and 
surface water systems, and exogenous, i.e. 
remotely generated runoff imported in flow 
from upstream. Some countries (e.g. Canada 
and Norway! have an abundance of water 
from endogenous sources; others (e.g. Egypt 
and Iraqi have a small endogenous supply but 




large exogenous volumes (others have small 
supplies from both sources). Use of exogen- 
ous water carries an increasing risk because 
of dependence on sufficient supply from 
upstream countries. 

There are well over 200 major inter- 
national rivers and a host of smaller ones''. 
As demands on inland water resources 
continue to grow through the 21st century, as 
they undoubtedly will, management of these 
and the biological resources they contain will 
also grow ever more challenging. 

Management of marine ecosystems 
Management of the terrestrial environment is 
typically carried out alongside more or less 
severe anthropogenic disturbance. Although 
the particular nature of the marine biosphere 
has to some extent buffered it from the 



Global biodiversity: responding to change 



207 




/ 



impacts of humans, it also imposes its own 
set of difficulties and constraints for rational 
management. Firstly, because almost all of it 
is generally out of sight, impacts are not 
immediately apparent, so that extreme deter- 
ioration may take place before anyone is 
aware of the fact. There may also be less 
incentive to take action than with terrestrial 
ecosystems where such deterioration has a 
direct impact on people. Secondly, with some 
exceptions (such as communal property 
resources on reefs and other inshore areas in 
parts of the South Pacific], living marine 
resources have been widely considered open- 
access resources, particularly those outside 
territorial waters (usually up to 12 nautical 
miles (nml from shore]. There is thus, quite 
simply, an incentive for any given individual 
to exploit a resource as fast and as intensively 



as possible before someone else does. 
Thirdly, the ability of water to transport large 
amounts of dissolved and suspended mater- 
ials, including living organisms, means that it 
is extremely difficult to manage limited areas 
of marine habitat in isolation. 

With the introduction of the exclusive 
economic zone (EEZ) under the United 
Nations Convention on the Law of the Sea 
lUNCLOS], which allows nations control over 
resources in an area up to 200 nm offshore, a 
far greater proportion of the world's seas now 
come within the control of individual nations. 
At present 99 percent of world fisheries catch 
is taken within EEZs. Although this should 
theoretically allow more rational manage- 
ment of marine resources, and more effective 
enforcement of management measures, 
progress in both has in practice been limited 



208 WORLD ATLAS OF BIODIVERSITY 



Map 8.5 

Marine protected areas 

The map shows the location 
of protected areas in lUCN 
categories l-VI that are 
entirely or in part marine, 
with map symbols graded 
according to protected area 
size (including any land 
present!. 

Note; For presentation 
purposes it has been 
necessary to use symbols 
that in most cases at this 
map scale greatly exceed the 
size of the protected area 
represented, giving the 
impression that much more 
of the world's coastal waters 
are protected than 
is in fact the case. 

Source: UNEP-WCMC database Idala 
extracted March 20021. mamtained m 
collaboration with lUCN World 
Commission on Protected Areas. 




to date, as evidenced by the increasing 
proportion of world's fisheries that are over- 
exploited. This is because fisheries manage- 
ment regimes are frequently subject to 
political pressure, so that in many countries 
quotas are habitually set higher than those 
recommended by fisheries biologists, and 
also because the active enforcement of regu- 
lations is difficult and expensive. Many 
countries lack the resources or the political 
will, or both, to enforce such regulations 
adequately. 

It is also increasingly apparent that individ- 
ual marine resources cannot be effectively 
managed in isolation from each other 
Complex interactions between populations of 
different organisms, when combined with 
perturbations in the environment and vari- 
ations in human impact le.g. changes in 




fisheries technology or fishing effort! create 
responses that may be far from intuitively 
predictable. Recognizing that the under- 
standing of such responses will require 
modeling and management of large-scale 
ecosystem processes, a number of large 
marine ecosystem (LMEl units have been 
identified-, based on the world's coastal and 
continental shelf waters, which are regarded 
as central to such analysis. Over 95 percent of 
the usable annual global biomass yield of 
fishes and other living marine resources is 
produced within 64 identified LMEs, nearly all 
of which lie within and immediately adjacent 
to the boundaries of EEZs of coastal nations. 
Many LMEs include the coastal waters of 
more than one state. In these cases, it will be 
effectively impossible for individual nations to 
assess whether their use of marine resources 



Global biodiversity: responding to change 



209 



t'i^^ 




¥'^-^_^- 



-rJ 



.vj; 



A f^' 



'-.-TC-v ;^ 



"'^^ XW^' 



w 



i. 






\K^--'' 




is sustainable in isolation from neighboring 
nations. Coordination between states in mon- 
itoring and resource management will thus 
become increasingly necessary as the press- 
ures placed on these areas increase. 

A critical need in monitoring marine 
ecosystems is the development of consistent 
long-term databases for understanding 
between-year changes and multi-year trends 
in biomass yields. For example, marked 
alterations in fish abundances were observed 
during the late 19605 when there was intense 
fishing within the northeast US continental 
shelf LME. The biomass of economically 
important finfish species le.g. cod, haddock, 
flounders! declined by approximately 50 
percent, and this was followed by increases 
in the biomass of lower-valued small elasmo- 
branchs Idogfish and skates). Management of 



marine fisheries will need to take these kinds 
of species dominance shifts into account in 
the development of strategies for long-term, 
economic sustainability of the fisheries". 
Monitoring the changing states of LMEs has 
received considerable attention, with several 
now being assessed and managed from a 
more holistic ecosystem perspective*- ■'. 

Marine protected areas 

The long-term management of LMEs is highly 
complex and there is an urgent need for 
smaller-scale and more immediate app- 
roaches. As with terrestrial ecosystems, the 
establishment of protected areas in marine 
ecosystems has been viewed as a major 
contribution to maintenance of biodiversity. A 
1995 review' identified just over 1 300 marine 
protected areas in existence at that time. 



210 WORLD ATLAS OF BIODIVERSITY 



Map 8.6 

International protected 

area agreements 

The map shows the location 
of protected areas managed 
under the Ramsar 
Convention on Wetlands, 
under the World Heritage 
Convention or as a 
biosphere reserve within 
the UNESCO Man and the 
Biosphere Programme. 

Source: UNEP-WCMC database Idata 
extracted March 20021. maintained in 
collaboration with lUCN World 
Commission on Protected Areas, 



^^' 



World protected areas 
Ramsar site 
World Heritage site 
■ MAB biosphere reserve 



ranging in size from 1 hectare to 2UM million 
hectares Ithe Great Barrier Reef Marine Park) 
IMap 8.5). Effective management and control of 
marine protected areas is problematic, partic- 
ularly if, as is often the case, they are in areas of 
intensive and potentially conflicting resource 
use. As noted above, marine ecosystems are 
also in general more difficult to protect than 
terrestrial ones from allochthonous inputs (i.e. 
those originating elsewhere). Although a no- 
catch regime can be effective in small marine 
reserves, in general it has been found that 
large, carefully zoned, multiple-use areas are 
more practical and effective than small re- 
serves. Sanctuaries or strict reserves may still 
be required for critical habitat areas such as 
nutrient sources, areas of high biological diver- 
sity, nesting sites of threatened species or to 
protect breeding stocks of important fishes" ". 










■^-. 









Reversing change: restoration and 
reintroduction 

Increasing recognition of widespread environ- 
mental degradation has led to a growth of 
interest in both the science and practice 
of ecological restoration. The main aim of 
such restoration is to reestablish the key 
characteristics of an ecosystem, such as 
composition, structure and function, which 
were present before degradation took place. It 
has been suggested that ecological restor- 
ation IS a crucial complement to the estab- 
lishment of protected areas for safeguarding 
biodiversity'', and it is widely expected that 
restoration will become a central activity in 
environmental management in the future. 
Such efforts are being supported by develop- 
ment of national and international policies. 
For example, the UN Convention on Biological 



Global biodiversity: responding to change 






'Jp- 



-^ 




• w^f 



Diversity, Article 8f, states ttiat parties should 
'rehabilitate and restore degraded eco- 
systems and promote the recovery of threat- 
ened species, through the development and 
implementation of plans or other manage- 
ment strategies'. 

A large number of restoration projects 
have now been initiated in different parts of 
the Vi^orld, focusing on a variety of different 
ecosystem types, including grasslands, wet- 
lands and forests. Although a number of 
national governments are active in ecological 
restoration, sometimes on a very large scale 
(notably in North America], most projects are 
being undertaken by NGOs, often as grass- 
roots or community-based initiatives. For 
example, together with a variety of local 
partners, the Forests for Life program of 
WWF/IUCN is implementing restoration 






J" 




programs in areas such as the Lower Mekong, 
New Caledonia, the Mediterranean, India and 
the Carpathians. WWF/IUCN is increasingly 
recognizing the critical importance of de- 
veloping plans for restoration at the land- 
scape scale, and the need to provide benefits 
to local communities as well as to biodiversity 
Experience of restoration projects to date 
has highlighted how difficult such ecological 
rehabilitation can be in practice. Although 
many degraded ecosystems display an ability 
to recover through natural processes if the 
causes of degradation are removed, in many 
areas the extent of degradation has been so 
severe that greater management intervention 
is required for restoration to be effective. For 
example, severely deforested areas may 
require large-scale tree planting in order for 
forest ecosystems to reestablish on a partic- 



212 WORLD ATLAS OF BIODIVERSITY 



ular site. Restoration projects may also be 
difficult to manage or monitor, as it is often 
fiard to define with precision what the 
structure, composition or function of a given 
ecosystem was prior to degradation, partic- 
ularly in areas where degradation occurred a 
long time ago. Another key challenge to 




Restoration projects have 
been initiated in different 
parts of the world, focusing 
on a variety of ecosystem 
types, including grasslands, 
wetlands and forests. 



restoration projects is the high cost involved: 
for example, a plan to restore the Florida 
Everglades has recently been launched, at a 
total cost of US$7.8 billion^'. 

Efforts at restoring degraded ecosystems 
can be complemented by programs focusing 
on the reintroduction or reestablishment of 
species that have become extinct within a 
particular area. For such reintroductions to be 
successful, thorough knowledge of a species 
and its habitat requirements are needed, in 
addition to a clear understanding of the 
original causes of extinction. In some cases, 
such as large vertebrate predators, there may 
be considerable public antipathy to reintro- 
duction being attempted. However, there have 
been some notable examples of successful 
reintroductions, such as the Arabian oryx 
[Oryx leucoryx] to Oman", the white-tailed 
eagle [Haliaeetus albicilla] to Scotland, and 
the tvlexican gray wolf ICan/s /upusfaa/(ey/l and 
California condor [Gymnogyps californianus] 
to parts of the United States. Such examples 
provide important lessons for successful 
reintroductions and, together with habitat 



restoration projects, illustrate how positive 
action can contribute to reversing the trends of 
biodiversity loss. 

THE INTERNATIONAL DIMENSION 

National boundaries do not enclose all the 
world's biological diversity: the high seas, the 
deep seabed and Antarctica all contain natur- 
al resources, some of great interest or 
economic importance. (Management of bio- 
diversity in such areas can, by definition, only 
be achieved by means of international meas- 
ures. Similarly, areas or communities of 
particular interest may be crossed by national 
boundaries, and in such cases international 
cooperation is essential for conservation 
measures to be planned and implemented 
effectively. 

More generally, it is important to develop 
policy and planning at the global level to place 
national efforts in a broader context. In a 
hypothetical example, an individual country 
might devote more effort to conserving 
species that are rare or peripheral at national 
level, but widespread elsewhere, than to 
more common, nationally endemic species. 
Although the former may be regarded as 
national priorities, the latter may be more 
important to global biodiversity. Regardless of 
differences between national and global 
priorities, an international forum is needed to 
develop conservation science, to provide 
exposure for diverse opinions and an oppor- 
tunity for NGOs and other bodies to comment 
on policy, and to formalize agreements that 
can guide the way individual countries manage 
their environments. Such opportunities are 
offered by the international agreements that 
have recently been developed. 

International agreements 
A multilateral treaty is an international 
agreement concluded between three or more 
states and governed by international law (see 
Box 8.AI. Existing international treaties that 
deal entirely or in part with biological diversity 
have evolved in an uncoordinated manner. 
Despite this, and the consequent gaps and 
duplications in overall coverage, a handful 
of such treaties whose text was agreed 
during the 1970s have come to exert a 



Global biodiversity: responding to change 



powerful influence on the conservation and 
management of elements of biodiversity. 
Among the most notable are the 1971 
Convention on Wetlands of International 
Importance especially as Waterfowl Habitat 
iRamsarl; the 1972 Convention Concerning 
the Protection of the World Cultural and 
Natural Heritage (World Heritage); the 1973 
Convention on International Trade in 
Endangered Species of Wild Fauna and Flora 
(CITES); and the 1979 Convention on the 
Conservation of Migratory Species of Wild 
Animals ICMS). The 1982 UN Convention on 
the Law of the Sea (UNCLOS). which entered 
into force In 1994, has strong potential for 
enhancing marine and coastal conservation. 

The names of these major treaties indicate 
their sectoral focus and, even if the many 
Important regional and species-related trea- 
ties are also considered, it is clear that the 
total obligations explicit in existing treaties 
fall short of the demands of an adequately 
comprehensive system. The Convention on 
Biological Diversity (CBD), agreed at the 1992 
Earth Summit In Rio. attempted to meet many 
of these demands. It was the first treaty 
planned to concentrate specifically on the 
conservation and use of global biodiversity. Its 
text establishes the conservation and use of 
biological resources as a matter of common 
Interest to all. It has as Its objectives the con- 
servation of biological diversity, the sustain- 
able use of biological resources and the 
equitable sharing of benefits arising from the 
use of genetic resources, recognizing that a 
careful balance must be maintained between 
these If biological resources are to be used 
wisely. It not only acknowledges the control 
of individual states over their biological 
resources, but it also states their respon- 
sibility for protecting them and using them 
sustalnably. 

The United Nations Framework Conven- 
tion on Climate Change lUNFCCCl was also 
agreed at the Earth Summit and Is relevant to 
biodiversity management. The CBD, the 
UNFCCC and the United Nations Convention 
to Combat Desertification (UNCCD) (which 
arose from the Summit but was not agreed 
until 199i) are sometimes termed the 'Rio 
conventions'. 



Those with a particular focus on bio- 
diversity (CBD, CITES, CMS, Ramsar, World 
Heritage) are informally termed 'the bio- 
diverslty-related conventions'. They each 
Impose more or less rigorous reporting 
requirements on parties to them, and also 
generate a significant demand for information 
from their parties and others. Meeting these 
demands can place a substantial burden on 
governments, particularly those with limited 
resources, and work is proceeding on 
harmonizing information management among 
the treaties. 

In addition to the Rio conventions, the Rio 
Declaration (a set of guiding principles), and a 
comprehensive plan of action - Agenda 21 - 
were also agreed and adopted by more than 
178 governments at the Earth Summit. 
Agenda 21 Is designed to support sustainable 

Box 8.i Negotiating a multilateral treaty 



Text 

Negotiation of tfie text of tfie treaty can require many years and numerous meetings. It Is 
concluded by the adoption of the text, typically when all the participating states reach 
agreement over the wording. Adoption of a treaty does not by itself create any obligations. 

Consent 

A treaty does not come into force until two or more states consent to be bound by it. 
The expression of such consent is usually by signature, notification, acceptance, approval or 
accession or by other means where so agreed. Signature followed by ratification is the most 
frequent means of expressing consent. Signature refers to the signature of the diplomats 
negotiating the treaty and Is often synonymous with the adoption of the treaty. Ratification is 
the need for approval of the treaty by the head of state or the legislature. Accession is the 
normal way that states which did not participate In the negotiations become parties to the 
treaty and has the same effect as signature and ratification combined. 

Entry into force 

This final stage usually occurs when all the negotiating states have expressed their consent 
to be bound by the treaty; the date may be delayed to provide time for parties to adapt 
themselves to its requirements. When a large number of states participate In the drafting of 
a large multilateral treaty it often enters into force once a specified number have ratified. 



human development and stewardship of the 
environment, and to be implemented at a 
range of scales - globally, nationally and 
locally - by organizations within the United 
Nations system, governments and other 
major groups. 



2U WORLD ATLAS OF BIODIVERSITY 



Table 8.1 

Major global conventions 

relevant to biodiversity 

maintenance 

Notes: Conventions are 
listed in order of entry into 
force. Year' is date of 
agreement, Entry' is year in 
Vi/hicfi agreement entered 
into force, 'Parties' is 
number of party states as 
indicated at eacti agreement 
website in Marcti 2002. 



^^^^^^^^^^^F 


Scope ^^^^^^^^^^^^^^^^^H 


Convention on Wetlands of International Importance 


All aspects of wetland conservation and wise use. 


especially as Waterfowl Habitat 


Parties are required to list at least one w/etland of 




international importance for special management and 


(Convention on Wetlands or Ramsar Convention! 


protection. 


Year Entry Parties 




1971 1975 131 


Convention Concerning the Protection of the World 


To define and conserve the world's fieritage, by dravKing 


Cultural and Natural Heritage 


up a list of sites whose outstanding values should be 




preserved for all humanity, and to ensure their pro- 


(World Heritage Convention! 


tection through a closer cooperation among nations. 




Sites may be of importance as cultural heritage or 


Year Entry Parties 


natural heritage or both. 


1972 1975 167 


Convention on International Trade in Endangered 


Aims to prevent species being threatened with 


Species of Wild Fauna and Flora 


extinction because of international trade. Parties act by 




banning commercial international trade in an agreed 


[CITES! 


list of endangered species (Appendix-l listed species! 




and by regulating and monitoring trade in others that 


Year Entry Parties 


might become endangered or whose trade needs to be 


1973 1975 154 


regulated to ensure control over trade in Appendix-l 




species |Appendix-ll listed species!. 


Convention on the Conservation of Migratory Species 


Aims to protect migratory species and their habitats. 


of Wild Animals 


Parties cooperate in research relating to migratory 




species and provide immediate protection for species 


ICMS or Bonn Convention! 


listed in Appendix 1 of the convention. For those species 




listed in Appendix 11, parties are required to endeavor to 


Year Entry Parties 


conclude range state' agreements on their con- 
servation and management, a number of which have 


1979 1983 79 




been concluded. 


Convention on Biological Diversity 


The major international agreement on biodiversity, the 




CBD sets out a framework within which parties 


ICBD! 


undertal<e to carry out national and international 




measures aiming to: conserve biodiversity, make 


Year Entry Parties 


sustainable the use of its components, and share 


1992 1993 183 


equitably the benefits derived from the use of genetic 




resources The Conference of the Parties has so far met 




five times. The Cartagena Protocol on Biosafety was 




adopted in 2000 but is not yet in force. 



Global biodiversity: responding to change 



Ronve 



:onvention 



United Nations Frameworl< Convention on Climate 
Change 



lUNFCCCI 



Year 



Entry 



Parties 



1992 



1994 



179 



United Nations Convention on the Law of the Sea 

lUNCLOSI 



Year 



Entry 



Parties 



1982 



1994 



138 



United Nations Convention to Combat Desertification 
in those countries experiencing serious drought 
and/or desertification, particularly In Africa 



lUNCCD - Desertification Convention] 
Year Entry 



Parties 



1994 



1996 



179 



Scope 

Aims to stabilize greenliouse gas concentrations in tlie 
atmosphere at safe levels. Parties are required to make 
an inventory of their sources and sinks of greenhouse 
gases and to formulate policies and measures to 
mitigate and/or adapt to the effect of climate change. 
Developed country parties were required to reduce their 
emissions of greenhouse gases to their 1990 level by 
the year 2000. The Kyoto Protocol establishes further 
reduction commitments for developed country parties. 

Contains a comprehensive codification of the principles 
and rules relating to the seas. UNCLOS establishes 
rights and obligations relating to navigation, the 
conservation and use of marine resources, and the 
protection of the marine environment 



Aims to ensure improved management of dryland 
ecosystems and use of development aid. National action 
programs (NAPsI will address the underlying causes of 
desertification and drought and seek to identify 
preventative or remedial measures. Subreglonal and 
regional action programs ISRAPs, RAPsI will be devel- 
oped, particularly when transboundary resources such 
as lakes and rivers are involved. 



Agreement for the Implementation of the Provisions of The objective is to ensure the long-term conservation 



the UN Convention on the Lavif of the Sea relating to 
the Conservation and Management of Straddling Fish 
Stocks and Highly Migratory Fish Stoclts 



and sustainable use of straddling and highly migratory 
fish stocks. Emphasizes the precautionary approach, 
protection of marine biodiversity and the sustainable 
use of fisheries resources. 



(Straddling Fish Stocks Agreement] 



Year 



Entry 



Parties 



1995 



2001 



30 



International protected area systems 

Two international conventions and one inter- 
national program include provision for desig- 
nation of sites internationally important for 
biodiversity conservation. These are the World 
Heritage Convention, the Ramsar (Wetlands] 
Convention, and the UNESCO Man and the 
Biosphere (MABI Programme. The location of 



protected areas managed under these 
agreements is shown in Map 8.6. 

Ramsar Convention 

The Convention on Wetlands of International 
Importance especially as Waterfowl Habitat 
was signed in Ramsar (Iran] in 1971, and 
provides a framework for international 



214 WORLD ATLAS OF BIODIVERSITY 



The Ramsar Convention 
provides a framework for 
International cooperation 
for tfie conservation of 
wetland fiabitats. 



cooperation for tfie conservation of wetland 
habitats. It places general obligations on con- 
tracting party states relating to the 
conservation of wetlands throughout their 
territories, with special obligations pertaining 
to those wetlands which have been added 
to the List of Wetlands of International 
Importance. Each state party is obliged to list 
at least one site. Wetlands are defined by the 
convention as: areas of marsh, fen, peatland 
or water, whether natural or artificial, per- 
manent or temporary, with water that is static 
or flowing, fresh, brackish or salt, including 
areas of marine waters, the depth of which at 
low tide does not exceed 6 meters. There are 
currently (March 20021 131 contracting 
parties to the convention; 1 1^8 wetlands have 
been designated for Inclusion in the List of 
Wetlands of International Importance, cover- 
ing more than 96 million hectares. 

World Heritage Convention 

The Convention Concerning the Protection of 
the World Cultural and Natural Heritage was 
adopted in Paris in 1972, and provides for the 
designation of areas of 'outstanding universal 
value' as World Heritage sites, with the prin- 
cipal aim of fostering International co- 
operation In safeguarding these important 




areas. Sites must be nominated by the signa- 
tory nation responsible and are evaluated for 
their world heritage quality before being 
listed by the international World Heritage 
Committee. Of the 721 sites distributed 
among 12^ countries that are currently listed, 
144 cover natural heritage and 23 sites are 
mixed cultural and natural. 

Article 2 of the World Heritage Convention 
considers as natural heritage: natural fea- 
tures consisting of physical and biological 
formations or groups of such formations 
which are of outstanding universal value from 
the esthetic or scientific point of view; 
geological or physiographlcal formations and 
precisely delineated areas which constitute 
the habitat of threatened species of animals 
and plants of outstanding universal value 
from the point of view of science or con- 
servation; and natural sites or precisely 
delineated areas of outstanding universal 
value from the point of view of science, 
conservation or natural beauty. Criteria for 
inclusion in the list are published by the 
United Nations Educational, Scientific and 
Cultural Organization lUNESCOl. 

Biosptiere reserves 

The establishment of biosphere reserves is 
not covered by a specific convention, but is 
part of an international scientific program, 
the UNESCO Man and the Biosphere (MABI 
Programme. The objectives of the network of 
biosphere reserves, and the characteristics 
which biosphere reserves might display, are 
Identified in the Action Plan for Biosphere 
Reserves. There are currently 411 biosphere 
reserves, spread over 94 countries. 

Biosphere reserves differ from the pre- 
ceding types of site in that they are not 
exclusively designated to protect unique 
areas or important wetlands, but for a range 
of objectives which include research, mon- 
itoring, training and demonstration, as well 
as conservation. In most cases the human 
component Is vital to the functioning of the 
biosphere reserve, which does not nec- 
essarily hold for either World Heritage or 
Ramsar sites. Some biosphere reserves 
coincide spatially with Ramsar or World 
Heritage sites. 



Global biodiversity: responding to change 



217 



POSSIBLE FUTURES 

Previous chapters have introduced some of 
the impacts on the biosphere of human 
expansion and development, and this chapter 
has outlined some of the general ways in 
which humans have attempted to manage 
the extent or severity of these impacts on 
biodiversity. These approaches reflect current 
conditions of climate and human development 
and the present policy environment, but is 
there a way to incorporate possible future 
conditions within the planning process? 

Scenarios 

Scenarios are increasingly being used to inform 
policy development and implementation, by 
illustrating the possible outcome of current 
trends, and by highlighting the implications of 
different policy decisions. The development of 
scenarios has become an important tool by 
which scientists communicate the results of 
their research to decision-makers, as well as 
constituting a significant research endeavor 
in its own right. An example is provided by the 
Intergovernmental Panel on Climate Change 
[IPCCI, which provides scientific, technical 
and socioeconomic advice to the world 
community on the specific issue of climate 
change. Climate change scenarios have been 
developed by the IPCC through workshops 
and meetings involving experts in modeling, 
climate impact assessment and emissions 
scenarios. These scenarios have formed the 
basis of the decisions made by parties to 
the UN Framework Convention on Climate 
Change, which provides the overall policy 
framework for addressing the climate 
change issue. 

Few attempts have been made to develop 
scenarios of biodiversity change. One recent 
approach^" involved consultation between 
ecological specialists from a number of 
different regions, who developed global scen- 
arios for ten terrestrial biomes for the year 
2100, based on global scenarios of changes in 
environment and landuse. Five key deter- 
minants of changes in biodiversity were 
identified: landuse change, atmospheric car- 
bon dioxide concentration, nitrogen depo- 
sition and acid rain, climate and the introduc- 
tion of exotic species into an ecosystem. The 



expected changes in these drivers were then 
considered for different ecosystem types 
Ibiomesl, and the relative impact of the 
different drivers on biodiversity was also 
estimated. Three scenarios of future bio- 
diversity were developed for each biome. 
based on various assumptions about the 
interactions between the drivers affecting 
biodiversity change. 

The results of this investigation suggest 
that when all biomes are considered together, 
landuse change is the driver expected to have 
the largest impact on biodiversity at the global 
scale, with climate change ranking second in 
importance. Considering the impact of all 
drivers together, Mediterranean ecosystems 
appear to be most at risk, with grassland and 
savannah ecosystems also being at relatively 
high risk. 

To further illustrate possible future trends 
in global biodiversity, examples of two alter- 
native approaches to scenario development 
are described below. Both these approaches 
were developed as contributions to the Global 
Environment Outlook IGEOl produced by the 
United Nations Environment Programme 
lUNEPl, in collaboration with a wide range of 
partners. The latest report of the GEO process 
[GEO-31 provides further details of these 
scenarios, and the methods employed in their 
development". 

RIVM IMAGE scenario evaluation 

For GEO-3, UNEP developed scenarios through 
a highly participative process involving GEO 
collaborating centers and other partners 
throughout the world, allowing particular 
issues identified at regional scale to be incor- 
porated. The scenarios involved development 
of narratives or storylines, respectively termed 
Markets First, Policy First, Security First and 
Sustainabitity First. These each describe 
possible futures based on different inter- 
pretations of prevailing global driving forces. 

A number of models and analytical tools 
were used to develop the scenarios, to help 
guantify the dynamics described qualitatively 
in the narratives, and to compare impacts 
across regions. These included PoleStar, 
developed at SEI-Boston: IMAGE, developed 
at RIVM in the Netherlands; WaterGAP, 



218 WORLD ATLAS OF BIODIVERSITY 



Figure 8.2 

Possible future scenarios 
from GEO 3, evaluated 
witfi RIVM IMAGE: 
pressures on remaining 
natural areas in 2002 (topi 
and changes in pressure 
between 2002 and 2032 
under the Markets First 
scenario (bottom) 

Notes: The pressures 
selected are those believed to 
have a major influence on 
biodiversity and for which 
data are available. These 
include habitat loss, 
population density, primary 
energy use, temperature 
change, and restoration time 
for agricultural land and 
deforested areas. The loviier 
map shows the relative 
increase or decrease in 
pressure between 2002 and 
2032 {difference divided by 
the pressure in year 20021. 
No change means less than 
10% change over the 
scenario period; small 
increase or decrease means 
between 10 and 50% change, 
substantial increase or 
decrease means 50 to 100% 
change: and strong increase 
means more than doubling. 

Source; RIVM IMAGE; Bnnk"*; Bnnk"; 
Brink e( at.". 










Low pressure 
Very high pressure 



.■it '-•''. ,. ■■> 













y 



■■ Domesticated area 
I I Ice and polar area 







y 



^H Substantial decrease in pressure 

Small decrease in pressure 

No change in pressure 

Small increase in pressure 

Substantial increase in pressure 
^H Strong increase in pressure 

developed by CESR at Kassel in Germany; 
and the AIM model developed at NIES and 
Kyoto University in Japan. PoleStar offers a 
simple modeling approach to explore how 
assumptions affect social and environmental 
performance, and incorporates scenarios 
developed by the Global Scenario Group'^ For 



From domesticated to natural area 
From natural to domesticated area 
Remains domesticated 



I I Ice and polar area/no data 

the GEO-3 exercise, the scenario assumptions 
were applied to different regions individually, 
and were then used to initiate and inform the 
scenario design process. PoleStar was also 
used to test the plausibility of storylines 
identified through the participatory process in 
most of the regions. 



Global biodiversity: responding to change 



219 



The GEO-3 scenarios look 30 years ahead. 
Markets First \s a world in which market-driven 
developments converge on the values and 
expectations that prevail in industrialized 
countries. Policy First \s a world in which con- 
certed action on environment and social issues 
occurs through incremental policy adjust- 
ments. Security First is a world of fragmen- 
tation, where inequality and conflict prevail, 
brought about by socioeconomic and environ- 
mental stresses. Sustainability First is a world 
in which a new development paradigm emerges 
in response to the challenge of sustainability, 
supported by new values and institutions. 

In order to assess the impact on bio- 
diversity of such diverging trends, IMAGE was 
used to assess changes in the natural capital 
index. This index is designed as a measure of 
biodiversity in natural ecosystems and agri- 
cultural land, and is calculated as the product 
of remaining area and its quality. The current 
remaining natural area is taken as a percent- 
age of the total land area. Changes over time 
are caused by the conversion of natural 
ecosystems into agricultural and built-up 
areas, and vice versa. Ideally, ecosystem 
quality should incorporate measures of the 
abundance of characteristic species relative to 
a low-impacted baseline state, but such data 
are not generally available at global scale, and 
ecosystem quality is therefore approximated 
by means of four pressure factors that are 
assumed to have a major influence on bio- 
diversity and for which data are available. For 
each pressure factor, a preliminary range is 
defined from a review of relevant literature, 
from no effect to complete deterioration of the 
habitat if the maximum value is exceeded. The 
four selected pressure factors are population 
density (min-max: 10-150 persons per km-); 
primary energy use 10.05-100 petajoules per 
kml; rate of temperature change 10.2-2.0°C in 
a 20-year period); and a restoration time for 
abandoned agricultural land and deforested 
zones in reconversion towards natural, low- 
impacted ecosystems Imin-max: 100-0 years 
after conversion or clear-cutting). The proxy 
for ecosystem quality is a reversed function of 
these pressures, calculated as a percentage of 
the low-impacted baseline state. The higher 
the pressure, the lower the quality. Finally, the 



percentages for habitat area and quality are 
multiplied, resulting in a pressure-based 
natural capital index. The calculations have 
been carried out on a detailed latitude- 
longitude grid, before aggregation to sub- 
regions and regions. 

The maps Isee Figure 8.2) illustrate the 
combined effect of decreasing area and qual- 
ity of habitat. The Markets F/'rsf scenario sug- 
gests a strong decrease of habitat quality and 
quantity in most regions, but in some regions 
agricultural land is taken out of production and 
may therefore revegetate naturally through the 
process of succession. However, in biodiversity 
terms this reconverted land must be con- 
sidered of low quality during initial decades. 

GLOBIO 

GLOBIO (Global methodology for mapping 
human impacts on the biosphere! was 
launched by UNEP in 2001 as a simple global 
model to help visualize the growing impacts 
of infrastructural development on biodiversity 
and ecosystem resources". GLOBIO provides 
an assessment of the probability of such 
impacts occurring by defining buffer zones 
around infrastructure. The extent of impact 
zones varies with the type of human activity 
and density of infrastructure, region, vege- 
tation type, climate and sensitivity of species 
and ecosystems, and is based on extensive 
literature surveys of scientific studies assess- 
ing environmental impacts resulting from 
human development. 

By linking impacts in different ecosystems 
and regions with satellite imagery lAVHRR 
data from 1992-93 on 1 km- resolution in the 
Global Land Cover Characterization database 
version 2.0), available resources and infra- 
structure, overviews of the cumulative im- 
pacts of continuous development can be 
derived. Scenarios for impacts on biodiversity 
are based upon data describing existing 
infrastructure, historic growth rates of infra- 
structure, availability of petroleum and min- 
eral reserves, vegetation cover, population 
density, distance to coast and projected 
development. The outcome is a simple over- 
view of the current and possible future 
cumulative impacts of infrastructural develop- 
ment on biodiversity, assuming continued 



220 WORLD ATLAS OF BIODIVERSITY 



Figure 8.3 

Possible future scenarios, 

GLOBIO: the pressures on 

the environnnent in 2002 

(topi and in 2032 under 

the Markets First scenario 

(bottom) 

Notes; GLOBIO provides 
projections of human impacts 
based on the development of 
infrastructure. With an 
economy dominated by 
market forces, biodiversity 
will be affected by 
infrastructural development 
on over 72% of the land area 
by 2032, according to GLOBIO 
projections. 

Source: GLOBIO-'. GRID Arendal. 
UNEP. 





High impact 
Medium-high impact 
Lo»/-medium impact 



Semi-deserts and deserts 

Wetlands 

Forests 



Grasslands/savannahs 

Croplands 

Water 



growth in demand for natural resources and 
the associated infrastructure development. 

GLOBIO defined four different degrees of 
human impact: 

• Human populations and urban areas (high 
impact). This includes all areas that are 
within 0.5 km of a road, or within a few 
hundred meters of any development, inclu- 
ding urban areas. 

• Converted land (medium-high impact). This 
includes areas converted to agricultural 
lands, plantations, and with a high density of 
infrastructure (i.e. generally within 0.3-2 km 
of any infrastructure). 

• Areas under conversion and fragmentation 



Imedium-moderate impact). This includes 
areas within 2-10 km of infrastructure. 
• Relatively intact (low impact). These are 
areas that still retain their natural state to a 
large extent, i.e. more than 10 km from any 
infrastructure. 

GLOBIO scenarios are based on two assum- 
ptions: that stable gross domestic product 
(GDP) or an increase in GDP will require 
further development in infrastructure, and 
that impacts of such developments on bio- 
diversity will decrease with distance from the 
infrastructure. The scenario illustrated here 
[Markets First, Figure 8.3) assumes a 3-^ 
percent annual increase in economic growth. 



Global biodiversity: responding to c ti a n g e 



Because of differences in tilstory, population 
density, coastal distance, landcover, availa- 
bility of mineral and petroleum resources 
[included as part of the model) and current 
economic capacity, ttie rate of change is not 
the same in all parts of the world. The 
implication of this scenario is that if further 
expansion is not controlled, increasingly 
severe losses of biodiversity will occur as a 
result of continuing human development. 
GLOBIO projections suggest that increased 
industrial exploration for oil, gas and minerals 
will accelerate road construction, draining of 
rivers for hydro-power and irrigation, and 
increased immigration, logging, and conver- 
sion of land to plantations and farmland, 
resulting in heavy losses of biodiversity in 
many areas. During the last 150 years, 
humans have directly impacted and altered 
close to Ul percent of the global land area. 
Under the scenario presented here, bio- 
diversity will be threatened in almost 72 
percent of the land area by 2032. Losses of 
biodiversity are likely to be particularly severe 
in Southeast Asia, the Congo basin and parts 
of the Amazon. As much as 48 percent of these 
areas will become converted to agricultural 
land, plantations and urban areas, compared 
with 22 percent today, suggesting wide 
depletions of biodiversity. 

Conclusions 

In considering the results of such scenario- 
building exercises, it is important to recognize 



that the future is very uncertain. None of 
the approaches described above attempts to 
provide accurate predictions of what is likely 
to happen in the future; rather, the aim is 
to illustrate what could occur if current 
trends continue. Although differing in the 
methodologies adopted and the results 
obtained, each of these approaches suggests 
that human activities will continue to have 
a major impact on biodiversity in coming 
decades. It is clear that those areas charac- 
terized by the highest numbers of species, 
such as tropical forests and Mediterranean 
ecosystems, are at particularly high risk of 
suffering major losses of biodiversity. In 
contrast, northern temperate areas appear 
less likely to experience such severe impacts, 
although significant losses of biodiversity may 
still be anticipated. The challenge for future 
efforts at scenario development will be to 
identify those policy decisions that could make 
the greatest contribution to the conservation 
of biodiversity while securing the economic 
benefits of sustainable development. With 
widespread declines in biodiversity likely to 
continue, given current trends in economic 
development and population growth, it seems 
probable that such decisions will at best 
contribute to slowing the rate of biodiversity 
loss. Significantly, none of the scenario 
approaches suggests much scope for the 
ecological recovery or rehabilitation of eco- 
systems that have already been degraded, 
except in localized areas. 



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8 WCMC 1998. Development of a national biodiversity index. Version 2. Unpublished report 
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11 Duellman, W.E. (ed.l 1999. Patterns of distribution of amphibians: A global perspective. 
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U Prendergast, J.R. et al. 1998. Biodiversity inventories, indicator taxa and effects of habitat 
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16 Prendergast, JR. and Eversham, B. 1997. Species richness covariance in higher taxa: 
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17 Mace, G. et al. 2000. it's time to work together and stop duplicating conservation 
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18 Margules. C.R. and Pressey, R.L. 2000. Systematic conservation planning. Nature U05: 
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20 Brooks, T. et al. 2001 . Towards a blueprint for conservation in Africa. BioScience 51 : 
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21 Wolf, A.T. et al. 1 999. International river basins of the world. International Journal of Water 
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24 Sherman, K. and Busch. D.A. 1995. Assessment and monitoring of large marine 
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27 Dobson. A.P., Bradshaw, A.D. and Baker, A.J. M. 1997. Hopes for the future: Restoration 
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28 WRI 2000. World resources 2000-2001: People and ecosystems. The fraying web of life. 
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Global biodiversity: responding to change 



223 



29 Spalton, A. 1990. Recent developments in the relntroduction of the Arabian oryx [Oryx 
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30 Sala, O.E. ef at. 2000. Global biodiversity scenarios for the year 2100. Science 287: 
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A 



APPENDIX 1 225 



APPENDIX 1: 

THE PHYLA OF LIVING ORGANISMS 

The phyla are listed in alphabetical sequence 
within each higher group, as are the 'kingdoms' 
within the Eukarya. The symbols associated with 
each phylum name indicate whether the species 
occur in marine, inland water or terrestrial 
habitats. Where more than one symbol is shown, 
this does not mean that species are equally 
distributed between them. In some cases, the text 
notes the principal habitat. For parasitic forms 
the symbol refers to the host habitat. 

Where possible, an estimate has been given of 
the number of described extant species in each 
eukaryote phylum. Because of the vagaries of 
taxonomy and lack of a consolidated catalogue of 
such species Isee Chapter 21, even this number is 
subject to considerable uncertainty in many 
cases; '?' indicates no estimate available. 

An attempt has also been made to gauge 
whether the given number represents a low, 
medium or high proportion of the possible total 
number of species in each phylum (i.e. including 
currently undescribed species). In rough terms, a 
low proportion is taken to indicate that the total 



diversity of the phylum may be an order of 
magnitude lor morel higher than the number of 
currently described species. A high proportion 
indicates that well over half the total number of 
species has almost certainly already been 
described. Generally, these estimates reflect the 
likelihood that microscopic, aquatic Especially 
marine! and parasitic groups are less thoroughly 
sampled than most macroscopic terrestrial forms. 
Because applying the species concept to 
prokaryotes is so problematic, no figures for 
species diversity are given for Archaea or 
Bacteria: it can be safely assumed that in almost 
all these phyla the proportion of existing diversity 
characterized to date is low. 



Source: Margulis, L. and Schwartz. K V. 1V98. Five kingdoms. An 
Illustrated guide to the ptiyla of lite on earth. 3rd edition. W.H. 
Freeman and Company, New York. 



T Marine 

♦ Freshwater 

• Terrestrial 



Crenarchaeota T ♦ 


Bacteria adapted to hot, acidic sulfur-rich 


Thermoacidophils 


environments often found in hot springs and around 




submarine vents. Pyrotobus grows at temperatures of 


Size Microscopic 


1 13°C. Sulfolobus tolerates temperatures up to 90°C 


Nutrition Chemoautotrophs or heterotrophs 


and may die if the temperature drops below 55°C; it 


l^odeoflife Free-living 


also tolerates highly acid conditions (pH of less than 




1, or stronger than concentrated sulfuric acidl. 


Euryarchaeota T ♦ • 


swamps, bogs and estuary sediments; many others 


Methanogensand halophils 


live in the guts of herbivorous animals, from termites 




to cows. They are cnemoautotrophs that obtain 


Size lulicroscopic 


energy by reducing COj (carbon dioxidel and oxidizing 


Nutrition Methanogens are chemoautotrophs 


H2 IhydrogenI to produce CH^ (methanel and H^O 


halophils are photosynthetic 


(waterl. They are responsible for production of most 


Mode of life Free-living or symbiotic, inhabiting the 


natural gas and for liberation of organic carbon from 


intestines of animals 


sediments into the atmosphere where it can be 


Euryarchaeota share similanties m ribosomal RNA 


reused, involving around 2 billion tons of methane 


sequence but consist of two very different groups. 


annually Halophils are aerobes that live in extremely 


f/lethanogens cannot tolerate oxygen lare obligate 


salty or highly alkaline environments, such as soda 


anaerobesi and free-living forms tend to occur in 


lakes, where tliey may be visible as a pink scum. 



ARCHAEA 



226 WORLD ATLAS OF BIODIVERSITY 



BACTERIA 



Actinobacteria ♦ • 

Actinomycetes. actinomycota and their relatives 

Size Microscopic 

Nutrition Helerotroptiic 

Mode of life Some free-living; some symbionts 



A large and diverse group of heterotrophic 
unicellular rod-shaped bacteria Icoryneforms), and 
filamentous, multicelled bacteria lactinomycetesl 
originally regarded as fungi. Some form pathogenic 
lesions on skin, while others are found in leaf litter; 
some of the latter can break down cellulose. Frankia 
15 a nitrogen-fixmg symbiont in plants. Streptomyces 
produces streptomycin and other antibiotics. 



Aphragmabacteria 



Size Microscopic 

Nutrition Heterotrophic 

Mode of life All symbionts, some parasitic 



? ? • Very small bacteria, lacking a cellwall, widespread 

in insect, plant and vertebrate tissues. Normally 
benign, but pathogenic in some conditions, and 
responsible for some forms of pneumonia and tick- 
borne diseases le.g. Ehrlichia]. Eight named genera 
to date. 



Chlorobia T ♦ • 

Anoxygenic green sulfur bacteria 

Size Microscopic 

Nutrition Photosynthetic 
Mode of life Mainly free-living; some symbiotic with 
other bacteria 



Phototrophic obligate anaerobes, inhabiting sunlit 
sulfide-nch habitats, particularly anaerobic muds. 
Some are tolerant of extremely high or low 
temperatures and salinities. Most use hydrogen 
sulfide or sodium sulfide in photosynthesis, instead 
of water, releasing sulfur instead of oxygen. Others 
form symbiotic associations with oxygen-respiring 
heterotrophic bacteria. 



Chloroflexa 

Green nonsulphur phototrophs 

Size Microscopic 

Nutrition Photosynthetic 
Mode of life Free-living 



♦ 9 Anaerobic filament-forming bacteria known from 

sulfur-rich habitats such as hot springs. Whilst 
these forms are typically photosynthetic, 
Chlorollexus can also grow heterotrophically in 
the dark. Three genera named to date. 



Cyanobacteria ▼ ♦ • 

Blue-green bacteria and chloroxybacteria 

Size Microscopic but relatively large 

Nutrition Photosynthetic 
Mode of life Free-living 



Photosynthesizmg bacteria, present in a great variety 
of habitats Until recently called 'blue-green 
algaeand considered to be plants. These bacteria 
dominated the landscape in the Proterozoic eon 
between 2 400 and 545 million years ago. 
Prochlorococcus occurs at the base of the photic zone 
throughout the world's oceans and may be one of the 
commonest bacteria. Many fix atmospheric nitrogen. 
Form reef-like stromatolites in some shallow-water 
marine environments. Around 1 000 named genera. 



Deinococci 

Heat- or radiation-resistant bacteria 

Size Microscopic 

Nutrition Heterotrophic 
Mode of life Free-living 



Spherical, heterotrophic, obligate or facultative 
aerobic bacteria highly resistant to heat [Thermus] or 
radiation iDe/nococcusl. Most metabolize sugars. 
Thermus aqualicus, isolated from hot springs in 
Yellowstone National Park, United States, is the 
source of Tag polymerase used in the polymerase 
chain reaction technigue. 



APPENDIX 1 227 



Endospora T ♦ • 

Endospore-forming and related bacteria 

Size Microscopic 

Nutrition Heterotrophic 

Mode of life Many synnbionts and parasites 



A very large, important and varied group of 
tieterotrophic bacteria, some obligate anaerobes, 
others facultative or obligate aerobes. Most form 
endospores Ipropagules within the parent cell 
resistant to heat and desiccation!. Some can break 
down lignin and cellulose, others are fermenters, 
breaking down sugars to produce compounds such 
as lactic acid and ethanol. Some, such as 
Streptococcus, are associated with infections. 



Pirellulae ♦ 

Protemaceous-walled bacteria and their relatives 

Size Microscopic 

Nutrition Heterotrophic 
Mode of life Mostly aquatic in freshwaters; some 
symbionts. some parasitic 



Diverse bacteria with proteinaceous cell walls; 
mostly obligate aerobic heterotrophs living in 
freshwaters. Chlamydia is parasitic, inhabiting 
animal cells and with apparently no independent 
means of producing energy C. psiHao causes 
psittacosis; C. tracliomatis causes trachoma 
blindness. 



Proteobacteria 

Purple bacteria 



▼ ♦• 



Size Microscopic 

Nutrition Include virtually all nutritional modes 

known 
Mode of life Major parasites and symbionts. some 

free-living aquatic forms 
An enormous and extremely varied group of bacteria 
including many disease-causing forms leg. 
Salmonella, a cause of food poisoning, and 
Neisseria, which causes gonorrheal and symbionts 



such as Escherichia coli. Proteobacteria show a 
great range of physical structure and metabolic 
activity; the group includes heterotrophs, 
chemotrophs. chemoheterotrophs, 
chemolithoautotrophs, photoautotrophs, 
photoheterotrophs, methylotrophs, hydrogen- 
oxidizers and sulfide-oxidizers. Many are facultative 
aerobes, respiring oxygen when this is available but 
able to survive by respiring, for example, nitrogen 
IN2I or sulfate ISO4' I when not. Responsible for a 
significant proportion of atmospheric nitrogen 
fixation. 



Saprospirae 
Fermenting gliders 



T4« 



Size Microscopic 

Nutrition Heterotrophic 

Mode of life Free-living, or symbiotic in animals 



One group includes anaerobic fermenters restricted 
to anoxic environments. Some free-living; some le.g. 
Bacteroides] inhabit intestinal tract of vertebrates, 
including humans, in enormous numbers. A second 
group includes aerobic forms inhabiting decaying 
vegetation and other organic-rich environments. 



Spirochaetae 

Spirochaetes 



T*« 



Size Microscopic 

Nutrition Heterotrophic 

Mode of life Free-living or symbiotic; some parasitic 



Spiral-shaped bacteria occurring in marine and 
freshwater habitats, including deep muddy 
sediments, and in animals, where many are major 
parasites or symbionts. Some respire gaseous 
oxygen, others are poisoned by it. Treponema 
pallidum causes syphilis and yaws, and Leptospira 
causes leptospirosis Twelve genera named to date. 



TJiermotogae 

Thermophilic fermenters 

Size Microscopic 

Nutrition Heterotrophic 
Mode of life Free-living 



▼ ♦ • Recently discovered obligate anaerobic bacteria 

known from submarine hot vents, terrestrial hot 
springs and subterranean oil reservoirs. Highly heat 
tolerant, living at temperatures of 50°C to 80°C. 
Ferment sugar and other organic compounds. 



BACTERIA 



228 WORLD ATLAS OF BIODIVERSITY 



EUKARYA: ANIMALIA 



Acanthocephala T ♦ • 


Adult individuals anchor themselves to the gut wall 


Thorny-headed worms 


of vertebrates. Infection generally occurs after an 


No. species described Over 1 000 


intermediate invertebrate host is ingested. Thorny- 


Proportion of group known Low/moderate 


headed worms appear to alter host behavior so as to 


Size Between 1 mm and 1 m in length 


increase probability of host being ingested by 


Nutrition Heterotrophic 


predator, and so transfer parasite to further host. 


Mode of life Parasitic worms that lack a free-liuing stage 


Humans are seldom parasitized. 


Annelida ▼ ♦ • 


A large phylum including polychaetes 19 000 


Annelids 


species], oligochaetes 16 0001 and leeches 15001. 


No. species described ca 16 000 


Most are active predators and scavengers. 


Proportion of group known Low/moderate 


Polychaetes include free-living and tube-dwelling 


Size From 0,5 mm to 3 m 


marine species, mainly benthic but some pelagic. 


Nutrition Heterotrophic 


Oligochaetes occur in freshwater, estuaries and 


Mode of life Segmented worms, mostly free-living m 


deep sea, but are most numerous on land where 


soils and sediments; some parasitic 


earthworms are very important to soil structure. 




Leeches are mainly free-living predators of 




vertebrates and invertebrates in freshwaters or 




water film on land; formerly more widely used 




for medicinal purposes IH/rudo medianalisl. 


Brachiopoda ▼ 


Cosmopolitan. Present between the intertidal zone 


Lampshells 


and 4 000 m depth. Usually occur cemented to surface 


No. species described ca 350 


by stalk Ipediclel; some species free-living on or in 


Proportion of group known Low/moderate 


marine sediment. Unlike moUusks, brachiopods have 


Size From 2 mm to 10 cm 


a lophophore, a specialized surface for gas exchange 


Nutrition Heterotrophic 


and food collection, and have dorso-ventral symmetry 


t^ode of life Benthic, mainly sessile, marine animals 


instead of lateral symmetry. Previously very diverse. 




especially during the Paleozoic era Isee Chapter 3), 




Some 30 000 extinct species have been described, and 




Lingula, with fossils from 400 million years ago, may 




be the oldest genus with living species. 


Bryozoa ▼ ♦ 


Marine bryozoans mainly intertidal, but also on 


Ectoprocts 


seafloor to considerable depths About 50 freshwater 


No. species described ca k 000 


species known, with jelly-like colonies on plant 


Proportion of group known Low 


surfaces in slow streams. Large colonies 10.5 m in 


Size Individuals mainly microscopic; 


diameter! derived from the asexual budding of zooids 


colonies to 0,5 m diameter 


Hess than 1 mm in length! may contain several 


Nutrition Heterotrophic 


million individuals. Marine forms contribute to reef 


Mode of life Sessile, colony-forming filter-feeders. 


diversity 


mainly marine 




Cephalochordata T 


Lancelets occur in estuary sediments and shallow 


Lancelets 


sandy seafloors. and live with the head protruding 


No. species described 23 


in order to screen out small plankton and organic 


Proportion of group known Moderate/high 


materials. They make up a small phylum of 


Size From 5 cm to 15 cm 


chordates with a cartilaginous rod dorsal to the 


Nutrition Heterotrophic 


gut InotochordI, a dorsal hollow nerve cord, and 


f«1ode of life Free-living, filter-feeding marine animals 


persistent gill slits in the pharynx, but without an 




internal bony skeleton or cerebral ganglion. They 




are the closest living relatives of vertebrate 




animals. Used as human food in some areas. 



APPENDIX 1 229 



Chaetognatha ▼ 


Common plankton in open seas, especially abundant 


Arrow worms 


in warm seas down to 200 m. Detect prey, mainly 


No. species described ca 7C 


copepods, by vibration sensors, and can inject 


Proportion of group known Low 


neurotoxins. Important to marine fisheries as a 


Size From 5 to 15 cm 


source of food for fishes. 


Nutrition Heterotrophic 




Mode of life Worm-lil<e, planktonic marine predators 




Cfielicerata T ♦ • 


segmented bodies, chitinous exoskeleton, jointed 


Chelicerates 


appendages! with insects and crustaceans. Most 


No. species described ca 75 000 


chelicerates are in Arachnida (more than 75 000 


Proportion of group known Low 


speciesi; others are horseshoe crabs IMerostomatal 


Size Macroscopic: the largest species of 


and sea spiders jPycnogonidaj, Arachnids are 


Pycnogonida have a leg span of almost 


ubiquitous on land, with a few freshwater species; the 


80 cm 


group includes ticks and mites, some of considerable 


Nutrition Heterotrophic 


importance as vectors of disease in humans and 


Mode of life Generally free-living and in most habitats 


livestock Merostomata include Limulus, superficially 


A very large and diverse arthropod phylum 


unchanged since the Silurian (see Chapter 31, 


characterized by claws Ichelicerae) on the anterior 


Pycnogonids range from the shallows to the deep 


pair of appendages, and sharing other features le.g. 


ocean 16 800 ml and from pole to pole. 


Cnidaria T ♦ 


A diverse phylum of radially symmetrical animals. 


Cnidarians, hydras 


including sea anemones, jellyfishes and corals. 


No. species described ca 9 000 


Specialized stinging cells called cnidoblasts are 


Proportion of group known Moderate 


diagnostic. Largest individuals le.g. lion's mane 


Size Mainly macroscopic 


Cyanea] may have tentacles many meters long. Reef- 


Nutrition Heterotrophic, mostly carnivorous; reef- 


building corals are of major importance in clear- 


building corals contain photosynthetic 


water coastal shallows in tropics and subtropics. 


symbionts 


Coral reefs often highly diverse and of great economic 


Mode of life Aquatic, almost all marine; colonial and 


value, Photosynthetic symbionts IdinomastigotesI of 


solitary, free-swimming and sedentary 


reef corals occur in polyp tissue at density up to 5 


forms known 


million/cm'; these require sunlight and limit reef 




growth to upper part of the photic zone. Non-reef 




coral without symbionts range down to 3 000 m. 


Craniata T ♦ • 


Agnatha in contrast to other vertebrates, all of which 


Craniates or vertebrates 


possess jaws IGnathostomatal. About half of all 


No. species described ca 52 500 


described vertebrate species are fishes: the 


Proportion of group known High 


Chondrichthyes (sharks and raysl, Osteichthyes (bony 


Size From about 1 cm to 35 m 


fishesi, the lungfishes and coelacanths. These make 


Nutrition Heterotrophic 


up around 25 000 species in lotal. The tetrapods (the 


Mode of life Free-living species present in most 


four-limbed non-fish vertebratesi include amphibians, 


habitat types 


reptiles, birds and mammals. Although not so 


This group contains the vertebrates; a very large and 


versatile as bacteria, vertebrates between them 


very diverse phylum of chordates. all of which, unlike 


extend from the air above the highest mountain to 


the acraniate chordates lurochordates. 


abyssal ocean depths, from sand desert to tropical 


cephalochordatesi, have a brain enclosed within a 


forest, and from hot springs to polar ice and subzero 


skull Icraniuml. The majority have a bony internal 


waters. The vertebrates include the most familiar 


skeleton. A small group of mainly marine species 


animals, and, with mollusks and crustaceans, most of 


(lampreys and hagfishl lack jaws and are grouped in 


those of direct nutritional importance to humans. 



EUKARYA: AN I MALI A 



230 WORLD ATLAS OF BIODIVERSITY 



EUKARYA: ANIMALIA 



Crustacea T ♦ • 


water fleas, woodlice, etc. Size ranges between 0.25 


Crustaceans 


mm [Alonella] and 2.8 m [Macrocheira claw span]. 


No. species described ca 40 000 


They are the predominant arthropods in most 


Proportion of group known Low/moderate 


freshwaters and widespread in all marine habitats. 


Size Mostly macroscopic 


from pelagic waters to ocean depths at 5 000 m, and 


Nutrition Heterotrophic 


in moist terrestrial situations. Numerous parasitic 


Mode of life Mostly free-living in aquatic and humid 


and commensal forms exist; pentastomids 


terrestrial habitats, some parasitic 


sometimes parasitize humans. Crustaceans sucti as 


A very large and very diverse arthropod phylum 


krill \Euph3U5ia superba] form key components of 


distinguished by having two pairs of antennae. 


the marine food web. There are numerous important 


Species occur in virtually all habitats. Includes crabs, 


crustacean fisheries. 


crayfish, prawns, barnacles, copepods. brine shrimp. 




Ctenophora T 


A small phylum of translucent, soft-bodied predators. 


Comb jellies 


Widespread in marine waters and possibly the most 


No. species described ca 100 


abundant planktonic animals between 400 and 700 


Proportion of group known Low 


m depth. Their fragility makes them difficult to 


Size Typically around 1 cm; largest up to 2 m 


collect and study. 


in length 




Nutrition Heterotrophic 




Mode of life Free-swimnning marine organisms 




Ectiinodermata ▼ 


Invertebrates with five-part radial symmetry, an 


Echinoderms 


internal calcium carbonate skeleton, and a water 


No. species described ca 7 000 


vascular system. Includes starfish, sea urchins, sea 


Proportion of group known Moderate 


cucumbers and others. Mostly benthic in intertidal or 


Size A few centimeters to near 2 m 


subtidal habitats. Sea lilies extend to 10 000 m, and 


Nutrition Heterotrophic 


sea cucumbers in places make up nearly the entire 


Mode of life Mostly free-living, benthic marine species 


animal biomass at these abyssal depths. Viviparous 




forms exist. Some, such as dried sea cucumbers 




ItrepangI, used as human food. 


Echiura T 


Echiurans live in U-shaped burrows in marine 


Spoon worms 


sediments, rock crevices and mangrove, with some 


No. species described ca UO 


forms extending to abyssal depths around 10 000 m. 


Proportion of group known Low/moderate 


Echiurans have a flexible proboscis which may 


Size From a few millimeters to 40 cm 


extend to 1.5 m from the bulbous, unsegmented 


Nutrition Heterotrophic 


body Cilia move food items down the proboscis to 


Mode of life Exclusively free-living marine organisms 


the mouth. 


Entoprocta T 


Vl/idely distributed in shallow coastal waters. 


Entoprocts 


One freshwater species known. Colonies 


No. species described ca 150 


permanently attached by stalks, horizontal stolons 


Proportion of group known Low 


and basal discs to solid substrate, algae or other 


Size Very small macroscopic animals, up to 1 cm 


animals. Often form conspicuous mat-like growth on 


Nutrition Heterotrophic 


seaweed and rocks. Filter-feeders, consuming 


Mode of life Mostly sessile, colonial marine 


diatoms, desmids, other plankton and detritus. 


organisms 


Loxosometla is free-living, and moves by 




somersaulting basal disc over tentacles. 



APPENDIX 1 231 



— ^y 



Gastrotricha 

Gastrotrichs 

No. species described ca 400 

Proportion of group known Low 
Size Average length 0.5 mm 

Nutrition Heterotrophic 
Mode of life Free-livmg, worm-lil<e animals, 
mainly marine 



▼ ♦ The ventral side is ciliated and often glued to 

substrate; exposed surfaces bear bristles or scales. 
Most occur in subtidal or intertidal sediments where 
form part of the marine meiobenthos: freshwater 
forms most abundant in small still waters. Important 
scavengers of dead bacteria and plankton. 



Gnathostomulids 

Jaw worms 

No. species described ca 80 

Proportion of group known Low 

Size Average length around 1.5 mm 

Nutrition Heterotrophic 

Mode of life Free-living marine worms 



A small phylum of translucent, benthic. worms 
capable of surviving in sediments very low in oxygen 
and high m hydrogen sulfide. Graze on baclena, 
proloctists and fungi in marine sediments. Have 
been found at several hundred meters depth. 
Population densities may exceed 6 000 per liter of 
sediment, outnumbering nematodes. 



Hemichordata 

Acorn worms 

No. species described ca 90 

Proportion of group known Low/moderate 

Size Adults between 2.5 and 250 cm 

Nutrition Heterotrophic 

Mode of life Sedentary, benthic marine species 



A small phylum of soft-bodied, benthic marine 
worm-like animals. Adults mostly sedentary and live 
burrowed in soft sediment of shallow seas, or in 
secreted tubes. Sexual and asexual reproduction 
occurs; colonies may be formed by budding. 
Hemichordates were previously classified as 
chordates. and resemble them in having ciliated 
gill slits in pharynx. 



Kinorhyncha 

Kinorhynchs 

No. species described ca 150 

Proportion of group known Low/moderate 

Size Up to 1 mm in length 

Nutrition Heterotrophic 

Mode of life Free-living marine animals 



Kinorhynchs are cosmopolitan in muddy bottom 
habitats, including estuaries and the intertidal zone, 
and to a depth of approximately 5 000 m. Some 
species are commensal with hydrozoans, bryozoans 
and sponges. 



Loricifera 

Loriciferans 

No. species described ca 100 

Proportion of group known Low 

Size Microscopic 

Nutrition Heterotrophic 

Mode of life Benthic marine species 



Widespread, probably cosmopolitan part of 
mlerstitial fauna. Life history incompletely known. 
Adults are sedentary on sand or graveL sometimes 
ectoparasites; the larvae are believed to be free- 
living and mobile. Protective plates cover the 
abdomen, into which the neck and head with 
mouth cone can be retracted. 



Mandibulata T ♦ • 

Mandibulates 

No. species described ca 950 000 

Proportion of group known Low 

Size From near microscopic to many 

centimeters 
Nutrition Heterotrophic 
Mode of life Most are free-living terrestrial species 
An exceptionally large and diverse arthropod phylum 
distinguished by a pair of crushing mandibles. 
Jointed chilinous exoskeleton and a single pair of 



antennae. Includes insects IHexapodal. centipedes 
and millipedes iMyriapodal. symphyla and 
pauropods. The largest group of animals, Hexapoda. 
contains some 950 000 described species and may 
number in millions. Insects range up to 30 cm length 
(giant stick insect Pharnacia serratipes]. Social 
insects such as termites, ants, some bees and 
wasps can form large colonies. Many insects are 
important crop pests or disease vectors; others are 
beneficial becarse of crop pollination or pest control. 



EUKARYA: ANIMALIA 



232 WORLD ATLAS OF BIODIVERSITY 



EUKARYA: ANIMALIA 



Mollusca T ♦ • 


deserts. Includes snails, slugs, mussels, chitons. 


Mollusks 


octopods, squid and others. Most species are free- 


No. species described At least 70 000 and 


living, although some are parasitic or commensals. 


possibly more than 


Many, e.g. bivalves, are sedentary as adults. Size 


100 000 


reaches maximum, approaching 20 m, in the giant 


Proportion of group known Moderate 


squid Architeuthis. Many important as food source. 


Size Range from near microscopic to 


Some forms act as intermediate hosts to parasites 


several meters 


(e.g. Sch/5(osom3| that cause serious human 


Nutrition Heterotrophic 


disease; other species can cause significant damage 


Mode of life Mainly free-living species present in 


to crops and constructions le.g. Dre/ssenal. Venom 


most habitat types; some parasitic 


of some marine gastropods is of medical interest. 


A large and highly diverse phylum, with species 


Monoplacophorans, the most primitive mollusks. 


occurring in benthic and pelagic marine viiaters, in 


were first seen alive in the 1970s but abundant in 


freshvioters of all kinds, and on land, from forests to 


Paleozoic Isee Chapter 31. 


Nematoda T ♦ • 


Possibly the most abundant animals living on 


Nematodes 


Earth, found in virtually all habitats and in many 


No. species described ca 20 000 


other organisms. Free-living forms are key to 


Proportion of group known Low 


decomposition and nutrient cycling. Many species 


Size From 0.1 mm to 9 m in length 


are important parasites of plants and animals. 


Nutrition Heterotrophic 


including humans le.g. filariasisl. Nematodes have 


Mode of life Free-living or parasitic worm-like 


provided important research animals in genetics and 


animals 


cell differentiation. 


Nematomorpha T ♦ • 


A small phylum of leathery, unsegmented, worm-like 


Nematomorphs 


animals. Occur widely in aquatic or moist terrestrial 


No. species described ca 240 


habitats. Eggs hatch into minute motile larvae which 


Proportion of group known Low 


enter host and metamorphose into immature worms. 


Size Ranging from 10 to 70 cm in length 


These burst out, killing the host, when near water or 


Nutrition Heterotrophic 


during ram. Hosts include annelids and arthropods. 


Mode of life Adults are free-living and usually 


Rarely found in humans, where appear non- 


aquatic; all are endoparasitic at some 


pathogenic. 


stage 




Nemertina T ♦ • 


Characterized by the slender anterior proboscis, 


Ribbon worms 


used for predation. defense and locomotion. 


No. species described ca 900 


Abundant in the intertidal zone; some forms are 


Proportion of group known Low/moderate 


pelagic. Freshwater and terrestrial forms are known. 


Size Macroscopic, from 0.5 mm to 30 meters 


Some species are symbionts or parasites; recorded 


in length, mostly small 


from echmoderms, nemertmes, annelids, mollusks 


Nutrition Heterotrophic 


and flatworms. May affect host reproduction. 


Mode of life Mostly free-living, predatory marine 


Malacobdella is a filter-feeder, inhabiting the mantle 


worms 


cavity of clams. 


Onychophora • 


Require high humidity levels to counter water 


Velvet worms 


loss through thin chitinous cuticle. Many occur 


No. species described ca 100 


in forest; some in caves. Alt carnivorous. Walk 


Proportion of group known Low 


slowly on U-43 pairs of stumpy legs. Two geographic 


Size From 1 to 20 cm 


groups exist; one mainly warm northern hemisphere; 


Nutrition Heterotrophic 


the other southern hemisphere. Many tropical 


Mode of life A small phylum of free-living, 


species are viviparous. 


terrestrial, worm-like animals 





APPENDIX 1 233 



Orthonectida T 


Recorded from echinoderms, nemertines, 


Orthonectida 


annelids, mollusks and flatworms. Less benign 


No. species described ca 2G 


than rhombozoans; may affect host reproduction. 


Proportion of group known Low 




Size Microscopic 




Nutrition Heterotrophic 




Mode of life Worm-lil(e internal parasites or 




symbionts of marine invertebrates 




Ptioronida T 


Most inhabit leathery chitinous tubes encrusted with 


Phoronids 


sand or shell fragments, some burrow in mollusk 


No. species described U 


shells or rock. From coastal shallows to 400 m 


Proportion of group known Low/moderate 


depth. Filter-feed on plankton and detritus. 


Size From 1 mm to 50 cm 


Cosmopolitan but not abundant; half the known 


Nutrition Heterotrophic 


species occur on Pacific coast of North America. 


Mode of life Sedentary filter-feeding marine worms 




Placozoa T 


Discovered in a seawater aquarium in 1883, and 


Trichoplax 


since reported in shallow marine water and marine 


No. species described 1 


research stations, Tnchoptax adhaerens is the least 


Proportion of group known High 


complex of all living animals, consisting of a few 


Size Up to 1 mm 


thousand cells but no distinct tissues, little is known 


Nutrition Heterotrophic 


of its life history. 


Mode of life Very small marine animal 




Platytielmintties T ♦ • 


Flatworms, flukes and tapeworms. Four classes. 


Flatworms 


Free-living soil flatworms most abundant in tropics; 


No. species described ca 20 000 


aquatic forms mainly temperate. Some can survive 


Proportion of group known Low/moderate 


in environments low in oxygen by oxidizing hydrogen 


Size Often a few millimeters; tapeworms to 


sulfide. Parasitic forms include flukes such as 


30 m in length 


Schistosoma, the cause of schistosomiasis, and 


Nutrition Heterotrophic 


tapeworms, obligate parasites of vertebrate gut- 


Mode of life Free-living or symbionts, many 


Many flatworm parasites have complex life cycle 


parasitic; found in freshwater, marine 


with infective larvae and intermediate hosts. 


and terrestrial environments 




Pogonophora ▼ 


shallow polar seas or Ithe vestimentiferansi around 


Beard worms 


hot submarine vents with a high hydrogen sulfide 


No. species described More than 1 20 


and methane content. Greatest diversity in the 


Proportion of group known Low 


western Pacific, Adult pogonophorans have no gut 


Size From 10 cm to 2 m in length 


and probably absorb nutrients directly from tentacles. 


Nutrition Heterotrophic 


Vent-living forms derive nutrients and energy from 


Mode of life Sessile, benthic marine worms 


the oxidation of hydrogen sulfide through symbiotic 


Pogonophora live in fixed upright chitin tubes secreted 


chemoautotrophic bacteria, which can occur at 


in sediments, shell or decaying wood on the ocean 


densities of 1 billion per gram of body tissue. 


floor Most abundant in cold, deep waters. 




Porifera T ♦ 


The vast ma]onty of sponges are marine and about 


Sponges 


100 species are freshwater Filter-feeders lone 


No. species described 5 000-10 000 


Mediterranean form passively captures crustaceans 


Proportion of group known Moderate 


and digests them externallyl. Many include 


Size Macroscopic, some to 2 m in height 


pholosynthetic symbionts, e.g. cyanobacteria. and 


Nutrition Heterotrophic 


brown, red or green algae. Sponges have simple 


Mode of life Sedentary aquatic animals 


structure with no tissues or organs and are generally 




supported by calcareous or silicaceous spicules or 




fibrous, proteinaceous matrix. 



EUKARYA: ANIMALIA 



234 WORLD ATLAS OF BIODIVERSITY 



EUKARYA: AN I MALI A 



Priapulida ▼ 


Found in sand or mud. from intertidal pools to 


Pnapulids 


abyssal depths, and from tropical waters to the 


No. species described 17 


Antarctic, Approximately half of the described 


Proportion of group known Low 


species are part of the marine meiobenthos li.e. 


Size Range between 0.5 mm and 30 cm 


small bottom- or sediment-living species between 


Nutrition Heterotrophic 


about 0.5 and 1 mm). 


Mode of life Exclusively marine, free-living, worm 




like animals 




Rhombozoa ▼ 


Mostly found in the kidneys of squid and octopus m 


Rhombozoans 


temperate waters. 


No. species described ca 70 




Proportion of group known Low 




Size Up lo 5 mm 




Nutrition Heterotrophic 




Mode of life Worm-like internal parasites or 




symbionts of benthic cephalopod 




mollusks 




Rotifera ▼ ♦ • 


Mainly freshwater, also in moist habitats on land; 


Rotifers 


about 50 species occur in benthic and pelagic 


No. species described ca 2 000 


marine habitats. Rotifers are the most abundant 


Proportion of group known Low 


and cosmopolitan of the freshwater zooplankton. 


Size Mostly microscopic, some to 2 mm 


Mostly free-living. Many live on other invertebrate 


Nutrition Heterotrophic 


organisms; many are endoparasites of invertebrates. 


Mode of life Mostly free-swimming in freshwaters 


Most free-living rotifers reproduce parthenogenetically. 


Sipuncula ▼ 


Benthic species, mainly in shallow, warm marine 


Peanut worms 


habitats; most abundant on rocky shores, but also 


No. species described ca150 


present in polar regions and down to 7 000 m in the 


Proportion of group known Low 


abyssal ocean. Ingest diatoms and other protoctists 


Size A few millimeters to 0.5 m in length 


or organic debris. Used locally as human food in the 


Nutrition Heterotrophic 


Indo-Pacific and China. 


Mode of life Exclusively marine, worm-like. 




burrowing or crevice-dwelling 




organisms 




Tardigrada T ♦ • 


Widely distributed from pole to pole. All are aquatic; 


Water bears 


land species live in the water film on mosses, forest 


No. species described ca 750 


litter and other habitats. A few marine species. Move 


Proportion of group known Low 


on four pairs of stumpy legs. Mainly ingest liquid 


Size Ivlainly microscopic 


food obtained by piercing protoctists, animals or 


Nutrition Heterotrophic 


plants The Mesotardigrada Igenus Thermozodiuml 


Mode of life Free-living animals, mostly in moist 


inhabit hot springs. Can survive extreme desiccation 


terrestrial and freshwater habitats 


with low metabolism or m encysted form, and when 




dormant Icryptobioticl may be tolerant of 




exceptionally high or low temperatures, approaching 




absolute zero. 



APPENDIX 1 235 



Urochordata ▼ 

Sea squirts 

No. species described ca 1 400 

Proportion of group l<nown Low/moderate 

Size From 1 mm to 2 cm 

Nutrition Heterotroptiic 

Mode of life Small, marine, filter-feeding animals 



Adults may be either benthic and sedentary Iclass 
Ascidiacae, tunicatesl or pelagic and free-swimmmg 
Iclass Larvaceal; Thaliacea or salps also free- 
swimming. All are ciliary filter-feeders, Urochordata 
have a dorsal hollow nerve cord, a cartilaginous rod 
dorsal to the gut InolochordI and gill slits in the 
pharynx at some stage, these being features of 
chordates. 



Asconnycota T ♦ • 

Ascomycotes 

No. species described ca 30 000 

Proportion of group l<nown Low 

Size Mainly microscopic but reproductive 

body up to several centimeters 
Nutrition Heterotrophic 
Mode of life Mainly terrestrial, in soil and leaf litter, 

or on and in other organisms 
A large diverse phylum distinguished from other 
fungi by the microscopic, sac-like, spore-producing 
reproductive structure lascusL Includes yeasts, 
morels, truffles, blue-green molds and lichens. 
Thread-like hyphae form network Imyceliuml 



through substrate. Many are Iree-living; many 
are parasitic. Several hundred forms occur in 
freshwaters. More than 10 000 are the heterotrophic 
components of lichens, which are joint organisms 
formed by ascomycotes with either photosynthetic 
green algae or cyanobacteria. As with other fungi, 
ascomycotes fulfil a key ecological role in breaking 
down organic material and transferring inorganic 
nutrients and water from soil to plants. Some are 
important in food preparation (e.g. baker's yeastl. 
Many cause important diseases of plants and 
animals, including humans, while others are 
sources of key medicinal substances such as 
penicillin and similar antibiotics. 



Basidiomycota ? ♦ • 

Basidomycotes 

No. species described ca 22 250 

Proportion of group known Low/moderate 
Size Generally microscopic or somewhat 

larger, but hyphae extend considerable 

distance, and fruiting bodies to 10 cm 

or more 
Nutrition Heterotrophic 
Mode of life Mainly terrestrial, in soil and leaf litter, 

or on trees and other organisms 
A large phylum including typical mushroonns, 
puffballs, stinkhorns, and rusts and smuts (some of 
which cause economically important plant diseases!. 



All are characterized by microscopic, club-shaped, 
spore-producing reproductive structures, typically 
borne in great numbers on a basidiocarp - the 
familiar mushroom. Largest known mushroom 
specimen grew to U6 cm wide and 54 cm high. 
Many basiomycotes form mycorrhizas with trees 
and shrubs; as with zygomycotes. the fungi 
move basic nutrients from soil to plant, and 
plant carbohydrates move into the fungus. Many 
ascomycotes fulfil a key ecological role in breaking 
down organic material, and transferring inorganic 
nutrients and water from soil to plants. Several 
species are valued wild food items. A few freshwater 
forms are known. 



Zygomycota ? ♦ • 

Zygomycotes 

No. species described ca 1 100 

Proportion of group known Low 

Size Generally microscopic or a little larger, 

but hyphae may extend considerable 

distance through soil 
Nutrition Heterotrophic 
Mode of life Mainly terrestrial in soil and leaf litter 



Most are saprobic Isaprophyticl, secreting digestive 
enzymes into organic material and absorbing 
nutrients released. Many are parasites on 
protoctists, small animals, plants or other fungi. 
Zygomycotes include about 100 species that form 
mycorrhizal associations, in which fungal partner 
contacts or enters plant roots and assists inflow of 
nutrients and absorbs organic plant substances in 
exchange. Most vascular plants probably have such 
relationships with fungi and these may be critical in 
nutrient-poor soils. A few occur in freshwaters. 



EUKARYA: ANIMALIA 



EUKARYA: FUNGI 



236 WORLD ATLAS OF BIODIVERSITY 



^r 



EUKARYA: PLANTAE 



Anthocerophyta 

Horned liverworts 

No. species described ca 100 

Proportion of group l<nown Moderate 

Size Low-growing plants 

Nutrition Ptiotosynttietic 

Mode of life Terrestrial, in moist tiabitats 



A small group of non-vascular plants of moist 
tiabitats, typically on woodland floor or water 
margins. Present worldwide in temperate and 
tropical regions. Among first colonists of bare 
substrates, including rocks. Some species have 
associated nitrogen-fixing cyanobacteria. 



Anthophyta ▼ ♦ • 

Flowering plants, angiosperms 

No. species described ca 270 ODD 

Proportion of group known High 

Size From less than 1 mm in length IWolffia 

angusta] to more than 100 m 

|fuca/yp(us regnansi 
Nutrition Photosynthetic 
Mode of life Flowering plants occurring in most 

habitat types 
An extremely diverse, geographically cosmopolitan, 
phylum of vascular seed plants, distinguished by 
flowers, and fruits that enclose the fertilized seeds. 
The great majority of species are terrestrial, in 
virtually all habitat types. Many occur in or around 



lakes, rivers and wetlands, and seagrasses occur 
subtidally in shallow marine waters. Two main 
groups are distinguished, according to whether 
the germinating seed has one or two seed leaves; 
Monocotyledones and Dicotyledones. Monocots 
include palms, lilies and the economically vital 
grasses; most monocots are herbaceous and 
woody forms lack special tissue that secondarily 
adds width to the trunk. Dicots form the larger 
group. Success of anthophytes appears linked 
to coevolution with animals, m particular with 
specialized modes of pollination and seed dispersal. 
All major food and medicinal plants, and hardwood 
timber trees, are found in this ohylum. 



Bryophyta ♦ # 

Mosses 

No. species described ca 10 000 

Proportion of group known Moderate/high 

Size Low-growing plants 

Nutrition Photosynthetic 

Mode of life Terrestrial, mainly in moist habitats and 

wetlands 
A large phylum of non-vascular plants li.e. lacking 
specialized xylem and phloem transport tissue! 



consisting of mosses and the genus Takakia. Often 
conspicuous in cold or cool temperate habitats, 
particularly tundra, where mosses are the dominant 
plants, and also in heathland, bogs, woodland, 
waterlogged areas and freshwater margins. Most 
diverse in moist tropical habitats. Many mosses well 
adapted to withstand desiccation; some occur in 
warm and regions. Peat moss [Sphagnum] 
contributes to the development of new soils. 



Coniferophyta # 

Conifers 

No. species described 630 

Proportion of group known High 

Size Shrubs or large trees, up to 100 m in 

height 
Nutrition Photosynthetic 
Mode of life Terrestrial 

Conifers are cone-bearing gymnospermous vascular 
plants, with needle-shape leaves, and are mostly 



evergreen trees. They form extensive forests at high 
latitudes in the northern hemisphere, and also occur 
more locally, often on and mountains; also common 
in the tropics and in temperate southern forests, 
where Araucana is widespread. In Sequoiadendron 
and Sequoia, conifers include the largest living plants. 
Many species in mountainous and northern areas 
have characteristic symbiotic mycorrhizal fungi. 
Conifers provide timber, paper pulp and ornamental 
plants, and some have food or medicinal value 



Cycadophyta t 

Cycads 

No. species described U5 

Proportion of group known High 

Size From shrubs to small trees of 18 m in 

height 
Nutrition Photosynthetic 
Mode of life Terrestrial 

A small phylum of seed-bearing, often palm-like, 
vascular plants restricted to the tropics and 



subtropics, where are present in a range of habitats, 
from moist forest to deserts and coastal mangroves. 
Diverse in the Cretaceous. Cycads are gymnosperms, 
I.e. seeds do not become enclosed in a fruit. Many 
are insect pollinated, often by beetles. All species 
have symbiotic, nitrogen-fixing cyanobacteria. 
Cycads provide a variety of materials, including 
thatch, food, medicines and ornamental plants. 
Cycad starch for bread requires special treatment to 
destroy potentially fatal toxins. 



APPENDIX 1 237 



Fillcinophyta 4 • 

Ferns 

No. species described ca 12 000 

Proportion of group known High 

Size From a few centimeters to 25 m 

Nutrition Ptiotosynttietic 

Mode of life Terrestrial; a few in freshwater 

A diverse phylum of vascular plants, the most 

species-rich group of plants lacking seeds. 

widespread from cold temperate areas to the tropics. 



Mainly in moist areas, such as forest floor and 
stream margins; species diversity highest In tropics, 
where many forms are epiphytic and some species 
grow as trees to 25 m in height. The aguatic Azolla. a 
very small floating fern, has symbiotic, nitrogen- 
fixing cyanobacterla. Several food, medicinal and 
other products are derived from ferns. Ferns, 
especially tree ferns, were diverse and very abundant 
in Devonian and Carboniferous times. 



Ginkgophyta 

Ginkgo 

No. species described 1 

Proportion of group known High 
Size To 30 m in height 

Nutrition Photosynthetic 
("lode of life Terrestrial 



A vascular seed-bearing tree, characterized by a fan- 
shaped leaf with bifurcating veins and a fleshy 
exposed ovule, now restricted as a wild species to 
steep forest in southern China. A wide diversity of 
ginkgophytes, of which Ginkgo biloba is the only 
survivor, existed during the Mesozoic, Now widely 
planted for ornamental purposes. It is a gymnosperm. 
I.e. seed not enclosed in a fruit. Leaf extract used as a 
traditional food and medicine in East Asia. 



Gnetophyta 

Gnetophytes 

No. species described ca 70 

Proportion of group known High 

Size Small trees, shrubs or vines 

Nutrition Photosynthetic 

Mode of life Terrestrial 



A small phylum of vascular seed plants, distinguished 
from other gymnosperms by having vessels for water 
transport similar to those of flowering plants. The 
three living genera differ greatly from each other 
Some plants of the genus Gnetum in tropical moist 
forest grow to 7 m in height; some Welwitschia, a 
unique, low-growing, cone-bearing plant of southwest 
African deserts, may be 2 000 years old. 



EUKARYA: PLANTAE 



Hepatophyta 

Liverworts 

No. species described ca 6 000 

Proportion of group known Ivloderale 

Size Low-growing plants 

Nutrition Photosynthetic 

Mode of life Terrestrial in moist habitats 



♦ • Non-vascular plants typically found in moist habitats 

growing on woodland floor, shaded stream banks, 
waterfalls or rocks; often epiphytic and often occur 
with mosses IBryophytal. Widespread in cold 
temperate regions, present in Antarctica, but species 
diversity highest in tropics Often among first plants 
to colonize burned or newly exposed substrates. 



Lycophyta ♦ # 

Club mosses * 

No. species described ca 1 000 

Proportion of group known Moderate 

Size Mainly low-growing herbaceous plants 

Nutrition Photosynthetic 

Mode of life Terrestrial, in moist and dry habitats 



Small seedless evergreen vascular plants found in 
temperate and tropical habitats, typically on forest 
floor in temperate regions although most tropical 
species are epiphytic. A few occur in and areas. 
Lycophytes were prominent in Paleozoic plant 
communities before evolution of flowering plants; 
although all living species are small, trees up to AO 
m in height were dominant in Carboniferous coal 
forests. Some similarity to mosses and conifers but 
unrelated to either 



238 WORLD ATLAS OF BIODIVERSITY 



Ar 



EUKARYA: PLANTAE 



EUKARYA: 
PROTOCTISTA 



Psilophyta 9 

Whisk fern 

No. species described 10 

Proportion of group l(nown Moderate/fiigfi 

Size Small herbaceous plants 

Nutrition Photosynthetic and symbiotic with fungi 

Mode of life Terrestrial 



A very snnall group of vascular plants, the only 
ones lacking both roots and leaves. Similar to 
earliest simple, leafless land plants of late Silurian 
and Devonian times, 400 million years ago. and 
conceivably direct descendants of them. Present as 
epiphytes or ground-living species with a restricted 
range in subtropics and temperate areas. These 
plants have a mycorrhizal association lalso seen 
in earliest fossil forms! with fungal hyphae that 
increase the flow of soil nutrients to the non- 
photosynthetic plant cells. 



Splienophyta 

Horsetails 

No. species described 15 

Proportion of group knovKn High 
Size Herbaceous plants 

Nutrition Photosynthetic 
Mode of life Terrestrial 



A small phylum of seedless vascular plants, 
with jointed ridged stems and tiny scale-like 
leaves. Found in moist or disturbed areas, including 
urban areas and roadsides; more typically in moist 
woods and wetland margins, and also in salt flats. 
Historically consumed as food in Europe and North 
America, poisonous to livestock. As with lycophytes, 
sphenophytes were diverse and abundant in 
Devonian and Carboniferous forests, with tree-like 
forms up to 15 m high. 



Actinopoda T ♦ • 

Radiolarians 

No. species described ca 4 000 

Proportion of group known Low 

Size Ivlicroscopic 

Nutrition Heterotrophic but most hold symbiotic 

photosynthetic haptomonads 
Mode of life Mostly marine, although the Heliozoa is 

mainly freshwater 



Relatively large, generally unicellular protoctists with 
radial symmetry. Some form large colonies in which 
many individuals are embedded in a jelly-like matrix. 
Some occur in open ocean waters, some are benthic. 
Many have silicaceous skeletons with spines or oars 
used for swimming. Most acantharians include 
photosynthetic grass-green haptomonad or yellow or 
green algae symbionts. 



Apicomplexa ? ? • 

Sporozoa 

No. species described ca 5 000 

Proportion of group known Low 

Size Single-celled 

Nutrition Heterotrophic 

Mode of life Symbiotic with or parasitic on animals 



Many of these spore-forming protoctists are 
bloodstream parasites with complex life cycles. 
Coccidians are the best-kno«in group because 
infection often causes serious or fatal intestinal 
tract infection. Many e.g. Eimena. infect livestock; 
kospora hominis is the only direct coccidian parasite 
of humans. Plasmodium causes malaria, probably at 
present the single most important infectious disease 
affecting humans. 



Archaeoprotista T ^ # 

Amitochondriates 
No. species described ' 

Proportion of group known Low'' 
Size Single-celled 

Nutrition Mostly heterotrophic 
Mode of life Free-living in aquatic habitats, and 
symbionts, often parasitic, in animals 



Anaerobic and lacking mitochondria. Many forms are 
parasitic or symbiotic in the intestines of animals, 
eg. wood-eating termites and cockroaches. Giardia 
causes giardiasis in humans. 



APPENDIX 1 239 



Chlorophyta T ♦ 

Green algae 

No. species described ca 16 000 

Proportion of group known Low 
Size Range from single-ceiled to 

macroscopic green seaweeds 
Nutrition Photosynthetic 
Mode of life Diverse, mostly marine and freshwater 

algae; a few symbiotic with other 

organisms 



Chlorophyles include unicellular and complex 
multicellular species as well as forms with many 
nuclei sharing the same cytoplasm, |v|a|or primary 
producers, they are estimated to fix over 1 billion 
tons of atmospheric carbon annually Symbiotic 
forms include Ptatymonas in the flatworm Convolula 
roscoffensis. Some forms are resistant to al least 
periodic desiccation. Some early form of chlorophyta 
almost certainly gave rise to plants. 



▼ ♦ 



Chrysomonada 

Chrysophyta 

No. species described i 

Proportion of group known Low' 

Size fvlost single-celled; some form large 

branching colonies 
Nutrition Photosynthetic 
Mode of life Free-living, mainly in freshwaters 



A large and diverse group of algae with golden- 
yellow pigments. The silicotlagellates are a 
component of marine plankton and extract 
silica from sea water to form shells. 



Chytridiomycota 

Chytridiomycota 

No. species described ca 1 000 

Proportion of group known Low 
Size Ivlicroscopic 

Nutrition Heterotrophic 
Mode of life Decomposers or parasites in 
freshwater or moist soils 



♦ • Feed by extending threadlike hyphae into living 

hosts or dead material. Simplest forms grow entirely 
within the cells of their hosts. Some are associated 
with plant diseases, e.g. Physoderma zea-maydis 
causes brown-spot in maize. Cell walls of chitin; 
" some with cellulose also. Chytrids may be ancestral 

to fungi. 



EUKARYA: 
PROTOCTISTA 



Ciljopfiora T ♦ 

Ciliates 

No. species described ca 10 000 

Proportion of group known Low 
Size Ivlostly microscopic and single-celled 

Nutrition Mostly heterotrophic 
Mode of life Mostly free-living; some symbionts or 
parasites 



Although most are unicells, a few multicellular 
forms resembling slime molds exist. Ciliates feed 
on bacteria or absorb nutrients from the surrounding 
medium, Entodmiomorphs live as symbionts in the 
stomachs of ruminants; the parasite Balantidium 
sometimes causes disease in humans. The free- 
living Paramecium and Slentor are well studied 
and much used in research and education. 



Cryptomonada 

Cryptophyta 

No. species described ' 

Proportion of group known Low'' 

Size Single-celled 

Nutrition Some heterotrophic; others 

photosynthetic 
Mode of life Mostly free-living 



T ♦ Cosmopolitan in moist areas. Most cryptomonads are 

flattened, ellipticaL free-swimming cells in freshwater. 
Marine species may form blooms on beaches; others 
are intestinal parasites. Heterotrophs ingest bacteria 
and protoctisls. Some of the photosynthetic forms 
possess yellow and red pigments in addition to 
chlorophyll, and some also contain blue-red 
phycocyanin pigments. Some form colonies of 
non-mobile cells embedded in a gel-like matrix. 



Diatoms 


T*« 


Widely distributed in the photic zone of marine and 
inland waters worldwide. Some occur in moist soils. 


No. species described ca 1 000 




Diatoms have distinctive paired tests or shells of 


Proportion of group known Low 




organic material impregnated with silica extracted 


Size Single-celled; some colonial 




from surrounding water Important basal 


Nutrition Mostly photosynthetic; 




components or marine and freshwater food webs. 


some saprophytes 






Mode of life Mostly free-livmg 







2*0 WORLD ATLAS OF BIODIVERSITY 



^r 



EUKARYA: 
PROTOCTISTA 



Dinomastigota ▼ ♦ 

Dinoflagellafes 

No. species described ca 4 000 

Proportion of group known Low 
Size Single-celled, up to 2 mm. 

occasionally colonial 
Nutrition Some are heterotrophic; 

others photosynthetic 
Mode of life Mostly free-living marine plankton 



Typically planktonic; some symbiotic with or live 
on marine animals or seaweed, some occur in 
freshwalers. Many adopt very different forms at 
different life stages. Gymnodintum microadnaticum 
IS the most common intracellular photosynthesizing 
symbiont in corals. Some produce powerful toxins 
and are an important cause of fish mortality le.g. 
Pleistena piscicida] and may form toxic 'red tides' 
le.g. Gonyaulax tamarensis]. Ciguatera poisoning in 
humans is caused by accumulations of dinoftagellate 
toxins in fishes and marine invertebrates. Many 
species le.g. Noctiluca] are bioluminescent. 



Oiscomitochondria ▼ ♦ • 

Flagellates, zoomastigotes 
No. species described ? 

Proportion of group known Low? 
Size Mostly unicellular 

Nutrition Generally heterotrophic; most 

euglenids are photosynthetic 
Mode of life Mainly free-living in a wide range of 

aquatic and terrestrial habitats 



All formerly regarded as protozoan animals, and in 
medical literature are commonly termed flagellates. 
Most feed on bacteria or absorb nutrients directly 
from surroundings; some, i.e. euglenids. are usually 
photosynthetic. Some are symbiotic or parasitic, the 
latter including organisms I rfypanosomal responsible 
for sleeping sickness and Chagas disease. 



T* 



Eustigmatophyta 

Green eyespot algae 

No. species described ' 

Proportion of group known Low? 

Size Single-celled 

Nutrition Photosynthetic 

Mode of life Free-living algae, mostly freshwater 



Planktonic algae with yellowish-green pigments, 
typically at the base of freshwater food webs. A few 
multicellular forms are known. Nine genera 
described. 



Gamophyta i 

Conjugating green algae 

No. species described Several thousand 

Proportion of group known Low 

Size Multicellular forms are macroscopic 

Nutrition Photosynthetic 

Mode of life Freshwater algae 



Multicellular filament-forming or unicellular green 
algae found in freshwaters. Many contribute to algal 
blooms and pond scum. Filamentous forms include 
Spirogyra. Desmids consist of paired cells joined 
at a narrow bridge through which their cytoplasm 
IS continuous. 



Granuloreticulosa ▼ ♦ 

Foraminifera and reticulomyxids 

No. species described ca 4 000 

Proportion of group known Low 

Size Mostly microscopic, but some several 

centimeters in diameter 
Nutrition Heterotrophic, some with 

photosynthetic symbionts 
Mode of life Mostly benthic, but some are free- 

swimmmg planktonic organisms; nearly 

all marine 



Foraminifera have multipored shells ItestsI 
composed of organic matter reinforced with 
minerals Isand or calcium carbonatel. Important 
in marine food webs. Many marine sediments are 
composed largely of foraminifera, and fossil species, 
about 40 000 of which are known, are important in 
stratigraphy Some of latter, e.g. Nummulites. can 
be up to 10 cm in diameter Reticutomyxids lack 
shells and form soft reticulate masses. 



APPENDIX 1 241 



Haplospora 

No. species described 33 

Proportion of group known Low 
Size Single-celled 

Nutrition Heterotrophic 
Mode of life Unicellular symbionts living in tiie 
tissues of marine animals 



Life history incompletely known, but characterized by 
production o( spores into water or host tissue. Many 
are benign symbionts, and exist in multinucleate 
"Plasmodium form, but several are parasitic and 
damage host tissues. Host animals include 
mollusks, nematodes, trematodes and polychaetes. 
Often found as parasites of parasites, e.g. within 
Irematode parasites of oysters. Formerly regarded 
as sporozoans. 



Haptomonada ▼ ♦ 

Prymnesiophytes 
No. species described 
Proportion of group known Low 
Size Mostly single-celled 

Nutrition Photosynthetic 
Mode of life Aquatic, with free-living and resting 
stages 



Most are marine; some occur in freshwaters. Two 
distinct life stages: a motile, golden-colored alga 
and a resting, coccolithophorid stage, covered 
in distinctive calcareous plates Icoccolithsl. 
Coccolithophorids are important in calcium 
carbonate sediments and in stratigraphic studies. 
Some are endosymbionfs of radiolaria lActinopodal. 



Hyphochytriomycota ♦ # 

Hyphochytrids 

No. species described 23 

Proportion of group known Low 
Size Microscopic 

Nutrition Heterotrophic 
Mode of life Present in freshwaters and soil 
moisture, saprophytic or parasitic 



Feed by extending threadlike hyphae into host 
tissue, typically algae or fungi, or into organic 
remains, e.g. insect or plant debris, where digestive 
enzymes are released and nutrients absorbed. 
Formerly regarded as fungi. 



Labyrinthulata ▼ 

Slime nets and thraustochytrids 

No. species described ' 

Proportion of group known Low? 

Size Colonies up to a few centimeters long 

Nutrition Heterotrophic 

Mode of life Colonial marine protoctists 



Slime nets consist of a complex colonial network of 
cells that move and grow within an extracellular 
slime matrix of their own making. Labynnthula 
grows on eel grass \Zostera\ where possibly 
pathogenic. Eight genera described. 



Microspora ? ? • 

Microsporans 

No. species described ca 800 

Proportion of group known Low 

Size Single-celled 

Nutrition Heterotrophic 

Mode of life Intracellular parasites of animals 



Anaerobic and lacking mitochondria Frequently 
orm large single-cell tumors in host animals, some 
highly pathogenic, some harmless. Nosema causes 
pebrine, a disease of silkworm larvae. 



Myxomycota ? ? • 

Plasmodial slime molds 

No. species described ca 500 

Proportion of group known Low 

Size Microscopic cells but macroscopic, up 

to several centimeters, in plasmodial 
form 

Nutrition Heterotrophic 

Mode of life Free-living organisms in damp 
terrestrial habitats 



Similar in some respects to cellular slime molds 
IRhizopodal. Myxomycotes have a sexual stage, 
and the Plasmodium that develops from the zygote 
is multinucleate. Fruiting stage develops in drier 
conditions. Feed by enveloping bacteria and 
protoctists growing on decaying vegetation. 
Key organisms in studies of cell motility 



EUKARYA: 
PROTOCTISTA 



242 WORLD ATLAS OF BIODIVERSITY 



A'- 



EUKARYA: 
PROTOCTISTA 



Myxospora T ♦ 

Myxospondians 

No. species described ca 1 100 

Proportion of group known Low 

Size Infected tissue may tiave growths of 

several centimeters in diameter 

Nutrition Heterotrophic 

Mode of life t»lulticellular symbionts, mostly 

parasites of fishes but also of marine 
and freshwater invertebrates 



Myxospondians penetrate the host integument 
and travel to the intestine where amoeboid forms 
carried to target organs are released. Many form 
large plasmodial masses attached to internal 
organs. Hosts include sipunculans and freshwater 
oligochaete worms. Most appear benign, including 
the fish symbionts, but some are important 
pathogens, e.g. Myxostoma cerebralis causes 
twist disease of salmon. Formerly regarded 
as sporozoans. 



Oomycota ♦ • 

Oomycetes 

No. species described Several hundred 

Proportion of group l<nown Low 
Size Microscopic 

Nutrition Heterotrophic symbionts 
Mode of life Mostly in freshwaters or soil; some 
parasitic on land plants 



Feed by extending threadlike hyphae into host tissue 
where release digestive enzymes and absorb 
nutrients. Familiarly known as water molds, white 
rusts and downy mildews. Many oomyctes are very 
important crop pests, e.g. Phytophthora inlestans 
causes potato blight; Saprolegnia parasitica attacks 
freshwater and aquarium fishes. Formerly regarded 
as fungi. 



Paramyxa 



6 

Low 



No. species described 
Proportion of group known 
Size Microscopic 

Nutrition Heterotrophic 
Mode of life Obligate symbionts living within the 
cells of marine invertebrates 



Characterized by production of multicelled spores 
within host tissue Live within annelids, crustaceans, 
mollusks and probably other groups of marine 
invertebrates. Formerly regarded as sporozoans. 



Phaeophyta T -^ 

Brown algae 

No. species described ca 900 

Proportion of group known Moderate/high 
Size Macroscopic plant-like organisms, 

mostly a few centimeters; sometimes 

much larger 
Nutrition Photosynthetic 
Mode of life Most live anchored to the substrate on 

rocky coasts 



Most widespread in temperate regions, where they 
usually dominate the mtertidal zone. Generally fixed 
but some, e.g. Sargassum, form large floating mats 
far out to sea. The largest proloctists: Pacific giant 
kelp iMacrocysds pynfera] sometimes reach 65 m in 
length. Brown algae are major primary producers in 
inshore environments and also provide habitat for a 
large number of macroscopic marine organisms. 



Plasmodiomorpfia 


? ? 


• 


Zoospores occur in soil and infect the host; a 
Plasmodium with many cell nuclei but no dividing 


No. species described 29 






walls develops within the host cell. Most species do 


Proportion of group known Low 






not appear to harm their hosts but Plasmodiophora 


Size Microscopic 






brassicae causes clubroot disease of brassicas and 


Nutrition Heterotrophic 






Spongospora sublerranea powdery scab of potatoes. 


Mode of life Obligate intracellular syml 


lonts, mainly 




of terrestrial plants; some 


parasitic 







APPENDIX 1 243 



■#> 



Rhizopoda ▼ ♦ • 

Amastigote amoebas and cellular slime molds 

No. species described ca 200 

Proportion of group known Low 

Size Single-celled or multicellular 

Nutrition Heterotrophic 

Mode of life Mainly benthic in aquatic habitats, or in 

water film on land; some amoebas are 

parasitic 



Often abundant in soil, where cyst-forming types 
highly resistant to desiccation. Entamoeba histolytica 
IS responsible for some forms of amoebic dysentery 
in humans. Some amoebas construct a coating ItestI 
from detritus and these have a fossil record from 
Paleozoic times, some fossil acntarchs Isee Chapter 
31 may represent testate amoebas. Cellular slime 
molds typically exist amid decaying vegetation, on 
logs or bark, and feed by enveloping bacteria and 
protoctists. The reproductive form of slime molds is 
an aggregation of cells each formerly having 
independent existence. Key experimental organisms 
in studies of cell communication and differentiation. 



Rtiodophyta T ♦ • 

Red algae 

No. species described ca i 000 

Proportion of group known Moderate/high 
Size tvlacroscopic plant-like organisms, up 

to 1 meter in size 
Nutrition Nearly all photosynthetic; a few are 

symbionts on other red algae 
Mode of life Virtually all are marine, a few species 

are freshwater or terrestrial 



Red algae occur attached to substrate on beaches 
and rocky shores worldwide. Most abundant in 
tropics. Many forms become encrusted with calcium 
carbonate; calcified red algae have a fossil record 
from the early Paleozoic. Agar jelly is extracted from 
red algae, and other extracts are used in food 
manufacture. Along with the Phaeophyta Ibrown 
algael the largest and most complex protoctists. 



Xanthophyta T ♦ • 

Yellow-green algae 

No. species described ca 600 

Proportion of group known Low 

Size Single-celled or colonial 

Nutrition Photosynthetic 

Mode of life Free-living, mostly freshwater algae 



Free-swimming unicells, or highly structured 
multicellular or multinucleated organisms, with 
gold-yellow xanthin pigments. Often form scum in 
pond water and margins. Typically form pectin-rich 
cellulosic cell walls; cysts often rich in iron or silica. 



Xenophyophora 

Xenophyophores 

No. species described i2 

Proportion of group known Low 

Size Sometimes several centimeters in 

diameter 
Nutrition Heterotrophic 
Mode of life Benthic marine forms 



Little-known bottom-living marine protoctists from 
deep sea and abyssal regions. Make shells itestsl 
from detritus leg. forammiferan shells, sponge 
spicules). Xenophyophores are the most abundant 
macroscopic organisms in some deep-sea 
communities, with several individuals per square 
meter Some acritarch fossils Isee Chapter 3| may 
have been xenophyophores. 



Zoomastigota ▼ ♦ • 

Zoomastigotes 

No. species described ^ 

Proportion of group known Low'' 

Size Single-celled, some colonial 

Nutrition Heterotrophic 

Mode of life Some are free-living in marine and 
freshwater environments, others are 
symbionts m the intestines of 
vertebrates 



Many feed by ingesting bacteria. Parasitic forms 
occur in the intestine of aquatic vertebrates, e.g. the 
opalinids. found in frogs and toads. One group of 
colonial forms, the choanomastigotes, may be 
ancestral to the sponges. 



EUKARYA: 
PROTOCTISTA 



244 WORLD ATLAS OF BIODIVERSITY 



CEREALS 



APPENDIX 2: 
IMPORTANT FOOD CROPS 

FOOD CROPS OF MAJOR SIGNIFICANCE 

These species and groups of species are those 
that, according to national food supply data 
maintained by the Food and Agriculture 
Organization of the United Nations IFAOI', provide 



5 percent or more of the per capita supply of 
calories, protein or fat in at least ten countries. 

Source- List ol crops from Prescott-AUen and Prescott-AUen ; other 

J i 5 

information from f^abberley , Smartt and Simmonds .Smith et at- , 
Vaughan and Geissler ; conservation status from Walter and Gillett . 



Millets 

Echinochloa frumentacea, Eieusine coracana, Panicum miliaceum, Pennisetum glaucum, 
Setaria italica (Gramineael 



Main uses: Ec frumentacea: Quicl<est growing of all 
millets, available in six weeks. Used for human 
consumption in India and East Asia and for animal 
fodder in the United States. f( coracana: Important 
staple in East and Central Afnca and India. Wild cereal is 
harvested dunng times of famine. In Africa it is the 
preferred cereal for brewing beer Seed heads may be 
stored for ten years. Pa. miliaceum: Cereal cultivated for 
human consumption, mainly in northern China, Russia, 
Mongolia and Korea, and as animal feed elsewhere. In 
Europe it is grown mainly as bird seed. Millets are 
generally tolerant of poor soils, low rainfall and high 
temperatures, and are quick matunng. Pe glaucum: 
Most widely grown of the millets. The mam cereal of the 
Sahel and northwest India. Heat and drought resistant. 
May contribute to incidence of goitre. 5. italica: A once 
important cereal that has declined in popularity, but still 
grown on a relatively large scale in India and China, 
mainly for home consu.mption Used also for animal 
fodder and bird seed. Early maturing and stores well. 

Origins: Echinochloa: Different strains are thought to 
have partially different origins. Approximately 35 spp. 
exist in the genus, distributed in warm areas. Eieusine: 
Eastern and southern Afnca highlands. Nine spp. in the 
genus in Africa and South America. Panicum: Unknown 
in wild state. The closest relative P. miliaceum var. 
ruderale is native to central China. At least 500 spp. in 
the genus in tropical to warm temperate areas. 
Pennisetum: Cultigen originated in West Africa from P. 
violaceum. Total of 130 spp, in the genus, found in 
tropical and warm areas. Setaria: Native to temperate 
Eurasia. Approximately 1 50 spp. in the genus in tropical 
and warm areas. 

Related species: Echinochloa: E. pyramidalis (tropical 
and southern Africa and Madagascar], used as fodder 
and locally as flour; £ turnerana channel millet 
lAustralial is a promising forage and gram crop. 
Several other spp. are weeds, Panicum: P. hemiotum 
[pifine grass. North America] and P. texanum (Colorado 



grass. North Amencal used as fodder. P. maximum 
(Guinea grass. Afnca, naturalized Amencal is cultivated 
as a forage crop, P. sonorum (Mexico! a minor grain; 
P. sumatrense (little millet, Malaysia! a minor gram. 
Pennisetum: Used as fodder lawn grasses, some 
grains. P. hohenacken (moya grass. East Africa to 
Indial IS suggested lor papermaking; P. clandestinum 
(Kikuyu grass, tropicat Afnca! pasture grass, erosion 
control, lawns; P. purpureum (elephant or Napier 
grass. Africa! fodder and paper P. violaceum (Africa! 
harvested during famines. Setaria: S. glauca (yellow 
foxtail! cattle fodder; S. paWdilusca is a cereal in 
Burkina Faso; S patmifolia (India! shoots are eaten in 
Java; S. pumita is cultivated as cereal; S. sphacelata 
(South Atncal is an important silage crop. 

Genetic base: Echinochloa: One sp. listed as 
threatened in 1797. Eieusine: Five races of cultivated 
finger millet recognized from Africa and India. 
Excellent prospects for improvement. Significant 
annual yield increases in India, mainly due to the 
incorporation of Afncan germplasm. Panicum: 16 spp. 
listed as threatened in 1997. Pennisetum: P. violaceum, 
the wild progenitor, is an aggressive colonizer and may 
be found in large populations around villages in West 
Afnca. The cultivated crop is relatively undeveloped. 
Open-poUinated cultivars are popular in Afnca and 
India. Five spp. listed as threatened in 1997. Setaria: 
Largely a crop of traditional agnculture systems. Two 
spp. listed as threatened in 1997. 

Breeding: El. coracana: Wild spp. in Africa cross with 
domesticated finger millet to produce fertile hybnds 
which can be obnoxious weeds. Pe. glaucum: Genetic 
exchange with related wild forms in same geographical 
area is possible, S, i'(a(/ca; Hybridizes easily with wild 
relative S. italica var viridis to produce fertile offspring. 

Germplasm collections: 90 500 general millet 
accessions. 45-60 percent of landraces and 2-10 
percent of wild spp. represented. 



APPENDIX 2 245 




Main uses; Early maturing gram with high yield 
potential; can be grown where other crops fail, e.g. 
above Arctic circle, at high altitude and in desert and 
saline areas. Most important as animal feed, also for 
brewing beer and human food. Mam producers in 
Europe. North Africa. Near East. Russia, China, India, 
Canada, United Slates. 

Origins: Southwestern Asia. Approximately 20 spp. in 
the genus, distributed m the north temperate region. 

Related species: H. distichon |2-rowed barley] is 
possibly H. vulgare x H. spontaneum 



Genetic base: Landraces have been almost completely 
replaced by pure line cultivars and the change in 
genetic structure of barley populations has been 
profound. Important contribution of Ethiopian barleys 
highlights need to broaden genetic base. Two spp were 
listed as threatened in 1997. 

Breeding: Fertile hybrids between wild and cultivated 
forms occur naturally where ranges overlap. Crosses 
possible with other spp. in the genus but not utilized in 
barley cultivars. Ethiopian barleys have been important 
as genetic source of disease resistance and improved 
nutritional value. 




Main uses: Highest world production of all grains. 
Primary source of calories and protein in humid and 
subhumid tropics. The gram is relatively low in protein; 
brown rice a source of some B vitamins Rice bran is 
used m animal feeds and industrial processes. Can 
grow in flood-prone areas Mam producers are China, 
India, Indonesia, Bangladesh, Viet Nam. Only 4 percent 
of world production is exported. Mam exporters 
Thailand, Viet Nam, Pakistan, United States. 

Origins: Two cultigens appear to have been 
domesticated independently. The origin of 0. sativa 
is uncertain, possibly derived in several centers from 
0. rufipogon (selected weed in Colocasia fields). 
Archeological evidence suggests origin m China or 
Southeast Asia. Center of diversity of 0. glabernma is 
the swampy area of the Upper Niger Approximately 
18 spp. in the genus, distributed in the tropics. 



Genetic base: Following agricultural intensification 
many populations of wild relatives have disappeared or 
intergraded with domesticated rice. Reduced genetic 
base has also led to repeated pest epidemics. Great 
genetic diversity exists in 0. sativa cultivars, much less 
in 0. glabernma. Rapid spread of improved rice 
varieties has displaced tens of thousands of landraces. 
many now extinct. 0. glabernma is rapidly being 
replaced by 0. sativa. Genetic erosion is reported in 
China, Philippines. Malaysia, Thailand and Kenya. 
Three spp. were listed as threatened in 1997. 

Breeding: sativa has formed numerous hybrids with 
wild spp. 0. nivara and 0. rufipogon. Genes improving 
tolerance to diseases or adverse conditions have been 
derived from African rices and wild relatives. 



CEREALS 



246 WORLD ATLAS OF BIODIVERSITY 



CEREALS 




Main uses: Cereal crop used as animal feed and for 
human consumption. Eaten mainly as rye bread and 
cnspbread Higher in minerals and fiber than wheat 
bread- Previously more popular as a bread flour; now 
largely replaced by wheat. Still important in cooler 
parts of northern and central Europe and Russia, 
cultivated up to the Arctic circle and up to k 000 m 
altitude. Tolerates poor soils. Also used in brewing 
industry and young plants produce animal fodder Ivlain 
producers are Russia and Europe, 



Origins; Probably originated from weedy Secale types 
in eastern Turkey and Armenia. S, montanum is 
possible ancestor Total of three spp in the genus in 
Eurasia, 

Genetic base; A number of weed ryes, found 
associated with agriculture throughout the Near East. 
are now considered to be subspecies of S, cereale. A 
complex of subspecies of 5. montanum extends from 
Ivlorocco east to Irag. Five spp, were listed as 
threatened in 1997, 



Sorghum 

Sorghum b/co/or (Gramineae 



Main uses: Staple cereal in semi-arid tropics. Mostly 
grown in developing countries, especially for domestic 
consumption by small farmers in Africa and India. 
Used in brewing beer and as animal fodder Gram 
stores well. Main producers are United States, India, 
China, Nigeria, Sudan, 

Origins; Developed primarily from the wild 

S arundinaceum in Africa, Total of 24 spp, in the 

genus in warm areas of the Old World and Mexico. 

Related species; Backcrosses with 5. arundinaceum 
gave S drummondii cultivated for forage; S, halepense 
IMediterraneanI is a widely naturalized fodder plant, 
often weedy. 




Genetic base: Wide variation in landraces. Genetic 
erosion reported in Sudan. Modern varieties have not 
been widely popular for use as human food. Two spp. 
listed as threatened in 1997. 

Breeding: Wild relatives may be an important source 
for disease resistance. 

Germplasm collections: 168 500 accessions. 21 
percent in the International Crops Research Institute 
for the Semi-Arid Tropics. 80 percent of landraces, 1 
percent wild spp. represented. 



APPENDIX 2 247 



Wheat 

Triticum aestivum, T. turgidum iGramineael 



Main uses: Most widely cultivated crop. Gram 15 gluten 
ricti and tiigtily valued for bread making. 90 percent of 
wheat grown 15 T. aestivum. Wheatgerm oil is highly 
unsaturated and high in vitamin E. Durum wheat, 
T. turgidum, has higher protein content. Used for 
making pasta. High nutritive value; easy processing, 
transport and storage. Ivlain producers are China, India, 
United States, France and Russia. 

Origins: tvlediterranean and Near East. Origin is 
complex and not fully understood, probably involving 
Aegitops spp. Total of four spp. in the genus, 
distributed from the Ivlediterranean to Iran. 



Genetic base: Large variation in the crop, around 
25 000 different cultivars. However, large areas are 
planted with genetically uniform crops and the inflow of 
landrace material into breeding programs is low 
18 percentl. Genetic erosion is reported in China, 
Uruguay, Chile and Turkey 

Germplasm collections: Approximately 850 000 
accessions. Largest collection (13 percentl in Centre 
for Maize and Wheat Improvement. 95 percent of 
landraces and 60 percent of wild spp. collected. 




Main uses: Mostly grown for human consumption in 
parts of Africa and Latin America; elsewhere mainly 
for animal fodder Starch may be extracted and used in 
food processing. The germ oil is important. Also used 
in the brewing industry. Mam producers are the United 
States, China, Brazil, Mexico, France. 

Origins: Probably derived from teosmte, Zea mays ssp. 
mexicana. Total of four spp. in the genus, confined to 
Central America, 

Genetic base: Most of the world's maize crop is 
derived from a few inbred lines. Landraces represent 
40 percent of the crop grown in developing countries. 
Genetic erosion is reported m Mexico, Costa Rica, 



Chile, Malaysia, Philippines, Thailand. Z. perenn/s was 
presumed extinct in the wild until its rediscovery m 
1977. Z- diploperennis was recently discovered and is 
now protected m the Sierra de Manantlan Biosphere 
Reserve, Mexico. Three spp. were listed as threatened 
in 1997. 

Breeding; Teosinle crosses readily with maize to produce 
fertile offspring, Tnpsacum crosses with less success. 
Neither has been widely used in breeding programs. 

Germplasm collections: 277 000 accessions, the 
largest existing at Indian Agricultural Research 
Institute. 95 percent of landraces and 1 5 percent 
wild spp. are represented. 



CEREALS 



2A8 WORLD ATLAS OF BIODIVERSITY 



TUBERS 




Yams 

Dioscorea spp. D. alata, D. bulbifera, D. cayenensis, D. dumetorum, D. esculenta, 
D. rotundata, D. trifida (Dioscoreaceael 



Main uses: Edible stem tuber, 28 percent starch and 
limited vitamin C. Important staple in the humid and 
subhumid tropics. Also major ingredient in oral 
contraceptives. Religious and cultural role. Good 
storage properties. West Africa produces 90 percent of 
world production; Nigeria atone produces 70 percent. 

Origins: Three mam independent centers of diversity or 
domestication in Asia, Africa and America. 
Approximately 850 spp. exist in the genus, distributed 
in tropical and warm regions. 



Genetic base: Predominantly a subsistence crop. 
Apparently little genetic erosion. Large genetic 
variability in wild edible forest yams. 68 spp. were listed 
as threatened in 1997. 

Breeding: New World and Old World spp. show strong 
genetic isolating barriers and crosses between them 
are not successful. 



Cassava 



Manihot esculenta (Dioscoreaceae! 



Main uses: Edible tuber containing 35 percent starch 
and vitamin C. Cultivated in almost all tropical and 
subtropical countries, mainly by smallholders. One of 
the most efficient crops for biomass production. A good 
famine reserve, able to withstand harsh conditions. 
Also animal feed. Mam producers are Brazil, Nigeria, 
Dem. Rep. of Congo, Thailand, Indonesia. 

Origins: Unknown in the wild state. Total of 98 spp. exist 
in the genus, occurring between southwest United 
States and Argentina. Most diversity occurs in northeast 
Brazil and Paraguay and in west and south Mexico. 

Related species: M. glaziovii is the source of Ceara or 
Manicoba rubber and oilseeds. 

Genetic base: Estimated 7 000 landraces. Local 
preferences in flavor, root texture and growth habit vary 




greatly, many farmers retain traditional cultivars 
despite improvements in new cultivars. Genetic erosion 
reportedly a risk in South and Central America, 
Thailand and China. 65 spp. were listed as threatened 
in 1997. 

Breeding: Variability of cultivated forms has probably 
increased through crosses with wild forms. M. glaziovii 
and M. melanobasis have contributed to improvement 
of cultivated form. High diversity in germplasm 
provides good improvement potential. Interspecific 
crossing with wild relatives may be employed further to 
broaden tolerance of different conditions. 

Gernnplasnn collections: 28 000 accessions, mostly in 
international centers of research. 35 percent of 
landraces and 5 percent wild spp. collected were listed 
as threatened in 1997. 



APPENDIX 2 249 




Potato 

Solanum tuberosum (Dioscoreaceael 



Main uses: One of the most important world crops. 
Cultivated in 150 countries, mainly for local 
consumption. Little international trade. Tubers are 
cooked or processed into a range of products. Starcfi, 
alcohol, glucose and dextrin are also ma|or products. 
Tubers also make animal feed. Potatoes are 80 percent 
water, 18 percent carbohydrates, with range of 
minerals, and a good source of vitamin C. I^ain 
producers are Russia, China, Poland, Germany, India. 

Origins: Maximum diversity in cultivated and wild spp. 
on the high plateau of Bolivia and Peru A number of 
ancestral spp involved. The gene pool consists of 
S. tuberosum ssp. andigenum and tuberosum, S. 
stenotomum, S. aj3nt)wn. S. goniocalyx, S. x cbauca, S. 
X juzepczukii, 5. x curtilobum, 5 phure/a. Total of 1 700 
spp in the genus, distributed worldwide. 

Related species: S. mebngena llndial (eggplant). S. 
centrale land Australia! and 5. muncatum Ipepinol 
lAndesI have edible fruit; 5. quitoense Inaranjidol (AndesI 



IS used for fruit juice; S. melanocerasum ('cultigenl 
Icultivated in tropical West Africa! fruit; S hyporhodium 
lupper Amazon!; 5. amencanum lyerba mora!. 

Genetic base: Between 3 000 and 5 000 varieties of 
potato are recognized by farmers in the Andes. Genetic 
erosion is reported in centers of origin, including Chile 
and Bolivia. In Peru, of the 90 wild potato spp. 35 are 
now extinct in the wild. Wild spp. and ancient cultivars 
largely replaced by modern varieties. Attempts to 
broaden the narrow genetic base have been slow. 
1 25 spp. were listed as threatened in 1 997. 

Breeding: Much introgression from wild relatives has 
been attempted, improving disease resistance and 
other traits. 

Germplasm collections: 31 000 accessions worldwide. 

20 percent are held by the Centre Internacional de la 
Papa, Lima. Peru. 95 percent of landraces and 40 
percent of wild spp. are collected. 




Main uses: Mainly temperate vegetable crop, but grown 
worldwide. Large number of edible and ornamental 
varieties, including cauliflower, calabrese and kohlrabi. 
Important component of human nutrition throughout 
the world: a good source of fiber, vitamins E, B and C, 
and also Vitamin A in the greener parts. Main 
production in Europe including Russia. 

Origins: The wild cabbage is native to Europe; 
development of cultivars took place in the 
Mediterranean region. Total of 35 spp. exist in the 
genus, distributed in Eurasia. 

Related species: Wide range of crops [variously leaves, 
buds, florets, stems and roots! eaten; also used for oil 
production. S. campestns and 6. napus Irapeseedl. 
B.cannata [Texsel greens! (northeast Africa!; 6. hirta 



(white and yedow mustard! (Mediterranean!; 6. juncea 
(Indian mustard! (Eurasia(; S.yuncea var cnspifolia 
(Chinese mustard!. 

Genetic base: Outbreeding nature. Large amounts of 
genetic variation in most crops, where not highly 
selected. Continuing emphasis on uniformity in recent 
decade and controls on release of new cultivars have 
led to significant reduction in genetic variation in 
commercial cultivars Wild relatives in Mediterranean 
are threatened; U spp. listed as threatened in 1997. 

Germplasm collections: Efforts made to ensure 
different crops are represented, including obsolete and 
locally popular varieties. Cultivars from southern 
Europe are less well collected. 



TUBERS 



LEAF VEGETABLES 



250 WORLD ATLAS OF BIODIVERSITY 



BEANS 




Main uses: L purpureus. Young pods and young and 
mature seeds of lablab are eaten; pulse contains 25 
percent protein, little fat and 60 percent carbotiydrate. 
Mam producers: India, Souttieast Asia, Egypt, Sudan. 
P lunatus: Dried or immature seeds of lima bean 
are used as pulses; seeds contain 20 percent protein, 
1 .3 percent fat, 60 percent carbohydrate; flour also 
obtained from seed. Mam producer is the United 
States. P. vulgaris: Most widely cultivated of all beans; 
in temperate areas grown mainly for the pod, which 
contains 2 percent protem and 3 percent carbohydrate 
with vitamins A, B, C and E; seeds have 22 percent 
protein, 50 percent carbohydrate, 1.6 percent fat and 
vitamins B and E. Mam producer: Brazil. 
V. unguiculata: Cov^pea is a nutritionally important 
minor crop in subsistence agriculture in Africa; dry and 
green seeds, green pods and leaves are eaten; highly 
resistant to drought. 

Origins: Lablab: African or Asian origin; only one 
species m the genus (previously Dolichos lablabl. 
Phaseolus: It is thought that separate domestications 
occurred in Central and South America from 
conspecific races; total of 36 spp. in the genus, found 
in tropical and warm America, Vigna: Center of 
diversity of wild relatives in southern Africa; greatest 
diversity of cultivated form exists m West Africa; 
subspecies dekindtiana is probable progenitor; total of 
1 50 spp in the genus, mainly in the Old World tropics. 

Related species: Phaseolus: Five cultigens exist in the 
genus: apart from P. lunatus and P. vulgaris, there are 
P. aculitolius Itepary bean, America); P. coccineus 
Iscarlet runner. Central America] and P. polyanthus 
lyear bean. Central America!. Various other spp. are 
important pulse crops, previously listed as Vigna spp. 
Vigna: Other spp. are used for forage and green 



manure, etc. Other pulses include: V. aconitilotia Imoth 
bean. South Asia; V. angulans lAduki bean. Asia]; 
V. mungo lurad, tropical Asia); I', radiata (mung bean, 
'Indonesia!; V. subterranea IBambara groundnut. West 
Africa). V. umbellata Ince bean, southern Asia); 
V. unguiculata )cowpea. Old World); V. vexillata (tropical 
Old World) which has edible roots. 

Genetic base: Lablab: Mainly grown in small plots and 
home gardens; larger areas under cultivation in 
Austraha. No threat of genetic erosion. Phaseolus: 
Much dry bean cultivation in the United States depends 
on very small germplasm base; improved varieties also 
widely adopted by smallholder farmers. Relatively wide 
genetic base provided by landrace groups, if conserved; 
most wild relatives widespread but populations of 
several taxa being lost to overgrazing in southwest 
United Slates and northern Mexico. Two spp. listed as 
threatened in 1997. Vigna: Breeding relies on narrow 
genetic base and hybridization with other Vigna spp. is 
important; more variability in wild relatives m the 
primary gene pool than in cultivated cowpea. Four spp. 
were listed as threatened in 1997. 

Breeding: Phaseolus: Several wild relatives are fully or 
partly compatible; populations of wild hma bean with 
larger seeds recently discovered in northwest Peru and 
Ecuador Vigna: cowpea crosses successfully with wild 

subspecies of V. unguiculata. 

Germplasm collections: Lablab: 1 1 500 accessions in 
Africa and Caribbean. Phaseolus: 268 500 accessions 
of Phaseolus spp. in total. 15 percent are held by 
Centro Inlernacional de Agricultura Tropical. Call, 
Colombia. On average 50 percent diversity m the genus 
IS represented. 



APPENDIX 2 251 



Groundnut 

Arachis hypogaea ILeguminosael 



Main uses: The edible nut contains 50-55 percent oil, 30 
percent protein, and is good source of essential minerals 
and E and B vitamins. Cultivated for ttie nut or for oil in 
many tropical and subtropical countries. Seed residue 
useful as animal feed. Nutshells are used as fuel and in 
industry. Stems and leaves used as forage. Mam 
producers are India. China. United States, Argentina, 
Brazil. Nigeria. Indonesia. Myanmar. Mexico. Australia. 

Origins: Mato Grosso in Brazil is the primary center 
of origin and diversity for the genus. The cultivated 
groundnut is thought to have originated in southern 
Bolivia and northv^est Argentina. Total of 22 spp. in the 
genus, all from South America. 



Coconut 

Cocos nucifera IPalmae) 



Main uses: The endosperm of the nut contains 65 
percent saturated oil, used in manufacture of 
margarine, soap, cosmetics and confectionery. Also 
eaten fresh, desiccated or as a coconut milk. Residue 
IS a high-protein animal feed. There are many more 
uses: source of naturally sterile water, fiber, wood, 
thatch. Mainly a smallholders' crop. Mam producers 
are the Philippines, Indonesia, India. Sri Lanka, 
Malaysia, Mexico. Pacific Islands. 

Origins: Possibly originated in Melanesian area of 
Pacific. Wild types predominate on the African and 




Genetic base: Cultivated as a marginal crop with 
relatively little selection pressure. Many varieties exist 
worldwide with broad adaptability 

Breeding: A. monlicola freely crosses with 

A. hypogaea- Wild Arachis material confers resistance 

on domestic form. 

Germplasm collections: 13 000 accessions at the 
International Crop Research Institute for the Semi- 
Arid Tropics. 




Indian coasts of the Indian Ocean, and scattered in 
Southeast Asia and the Pacific. Single species in 
the genus. 

Genetic base: Tendency to plant uniform, improved 
hybrids is reducing genetic variation particularly in 
domesticated types. 

Breeding: Wild and domestic coconuts are fully 
compatible. Hybridization with wild types has increased 
genetic diversity of cultivated crups. 




Main uses: The mesocarp on the fruit yields oil for 
human consumption. Unrefined oil is high in vitamin A. 
Oil may also be extracted from the kernel. An export 
crop and important for local consumption. Very high 
yielding. Malaysia supplies 70 percent of world exports. 

Origins: West Africa, originally a species of the 
transition zone between savannah and rainforest. 
Only 2 spp. exist in the genus. 



Related species: £ oleifera (tropical Americal is less 
important as an oil crop than £ guineensis. 

Genetic base: Populations in Africa are semi-wild. They 
are being thinned to make way for other crops. 
Plantations in Malaysia were based on material from 
only four specimens. New material is being introduced 
to broaden the genetic base. 

Breeding: Fertile offspring produced with £ oleilera. 



OIL CROPS 



252 WORLD ATLAS OF BIODIVERSITY 



OIL CROPS 



Soybean 

Glycine max iLeguminosael 




Main uses: The most important oil crop and gram 
legume in terms of production and international trade. 
An important basis of Asian cuisine, developed into 
various forms of food from soy sauce to tofu. Immature 
green beans and sprouts also eaten. Seeds contain 18- 
23 percent oil and 39-45 percent protein. Oil is used in 
various forms. Most of the meal is used as a high- 
protein animal feed fvlain producers are the United 
States, Brazil, China, Argentina, India. 

Origins: A cultigen, not known m the wild. Soybean is 
thought to have arisen as a domesticate in the eastern 
half of northern China about 3 000 years ago probably 
from S. soja. Total of 18 spp. exist in the genus, 
distributed from Asia to Australia. 



Genetic base: The genetic base of varieties is narrow 
worldwide Conservation of traditional landraces is 
urgently needed. Two spp. listed as threatened in 1997. 

Breeding; Wild spp. are increasingly used for 
improvement and there is good potential for further 
valuable characteristics to be found in wild Glycine spp, 
6. soj3 easily crosses with soybean. 

Germplasm collections: 174 500 accessions, 9 percent 
in Institute of Crop Germplasm Resources. Chinese 
Academy of Agricultural Sciences, Beying, China. 
60 percent of landraces and 30 percent wild spp. 
are represented. 



Cotton seed 

Cossypium barbadense, G. hirsutum (Malvaceael 



Main uses: Cotton is the second most valuable oil crop, 
as well as being the most important textile fiber Crop 
development is concentrated on fiber production 
because value is three or four times greater New 
World cottons took over from Old World forms after the 
European exploration of the Americas. Ivlain producers 
of G. barbadense are Russia, Egypt, Sudan, India, 
United States, China. 

Origins: Unique in that four spp. were domesticated 
independently for the same use as a fiber and oil crop, 
in Africa and India: G. arboreum and G. herbaceum; in 
Central and South America: G. hirsutum and 6, 
barbadense. Total of 39 spp, in the genus, found in 
warm temperate to tropical zones. 

Related species: 6. arboreum is still important in India 
and Pakistan. G, herbaceum is grown only on a small 
scale in Africa and Asia. 



Genetic base: Modern cultivars of G. hirsutum are 
responsible for over 90 percent of world production. 
New Gossypium spp, possibly occur in Arabia and 
Africa, Wild forms of G, herbaceum, G. hirsutum and 6. 
barbadense are known. Past breeding involved much 
introduction of genetic material from different 
geographic regions, but a severe narrowing of the 
genetic base has occurred in the production of modern 
G, hirsutum varieties Large amounts of fertilizers and 
pesticides required in modern cotton production. Eight 
spp. listed as threatened in 1997. 

Breeding: At least six related spp, have contributed 
genes of importance to the cultivated crop Ivlaterial 
from wild gene pool used in genetic engineering, G, 
herbaceum and G, arboreum are able to interbreed, 
although later generations have a high probability of 
failing reproductively. 



APPENDIX 2 253 



Sunflower seed 

Helianthus annuus (Compositae 



Main uses; Seeds contain 27-iO percent 
potyunsaturated oil and 13-20 percent protein. Oils and 
margarines used for human consumption, and for 
industrial uses, and waste products useful in animal 
feed. Pollinating bees frequently used for fioney 
production, (ulam producers are Russia, United States. 
Argentina. 

Origins: Probably originated in southwest North 
America, Total of 50 spp. exist in the genus, distributed 
in North America. 

Related species: Also ornamental; H. tuberosus 




{Jerusalem artichoke! is also eaten. H. petiolaris used 
for hybridization. 

Genetic base: Increased yields in hybrids led to 
increased interest and production in 1960s. Large gene 
pool exists in wild and weed sunflowers in North 
America, although habitat loss is resulting in 
population declines. 16 spp. listed as threatened in 
1997. 

Breeding: Resistance to several diseases was secured 
through hybridization with H. tuberosus. 




Main uses; Fruit with iO percent oil content. Highly 
superior oil for cooking, margarines, dressing; also 
used in cosmetics and pharmaceutical industry. Fruit 
eaten pickled. Despite competition with more modern 
oil-producing crops, olive oil still commands premium 
price. Recent rise in popularity and recognition of 
nutritional value. Mam producers are Spam, Italy. 
Greece. Turkey, Tunisia. 

Origins: Olive is a cultigen. evolved in eastern 
Mediterranean, 0, europaea ssp. oleasfer recognized 
as progenitor Total of 30 spp. in the genus, in tropical 
and warm temperate parts of Old World. 

Related species: Related species provide good timber 



Genetic base: A long-lived tree. The turnover of clones 
should be slow. Hundreds of distinct cultivars, found in 
different geographic groups. Olive production still relies 
on traditional cultivars. Few new varieties have been 
released- Decline in area under cultivation. Marginal 
groves have been abandoned with serious 
consequences for Mediterranean wildlife. Wild 
populations outside area of cultivation under pressure 
from cutting and land clearance; two spp. were listed 
as threatened in 1998. 

Breeding; Closely related to wild subspecies in the 
Mediterranean, Africa. Arabia, Iran and Afghanistan. 



OIL CROPS 



254 WORLD ATLAS OF BIODIVERSITY 



SUGAR CROPS 



FRUITS 




Sugarcane 

Saccarhum officinarum (Gramineae 



Main uses: Ma|or source of calories worldwide. 
Cultivated in about 70 countries, mainly in tropics. 
Requires good rainfall and rich soil for successful 
growth. Stems are easily transported. Mam producers 
are Brazil, India, China, Thailand, Pakistan. 

Origins; A cultigen with origin and center of diversity in 
New Guinea. Between 35 and iO spp. in the genus, 
distributed in tropical and warm zones. 

Related species: Other cultivated sugar canes include 
S barben, 5 edule and 5 sinense. S.robustum and S. 
spontaneum are wild sugar canes. 

Genetic base: Risk of genetic erosion reported in 
Assam and suspected in Indonesia, Papua New Guinea 
and Thailand, where monocrop plantations have taken 



Banana, plantain 

Musa spp. IMusaceael 



Main uses: One of the most popular dessert fruits in 
industrial nations, a major source of calories and export 
earnings in developing countries. Bananas and plantains 
are high in carbohydrates and potassium; bananas are a 
good source of vitamins C and 06, and plantains contain 
high levels of vitamin A. Numerous other uses. Mam 
producers are India, Brazil, Ecuador. Philippines and 
China for the banana; Uganda, Colombia. Rwanda, 
Dem. Rep. of Congo and Nigeria for the plantain. 

Origins: Bananas evolved in Southeast Asia from M. 
acuminata or combinations of M. acuminata and M. 
balbisiana. Plantains probably originated in southern 
India. Primary areas of diversity exist in Southeast Asia. 
Secondary areas also occur in tropical Africa, Indian 
Ocean islands and the Pacific. Fe'i bananas I2nl, 
thought to be derived from M. mactayi and possibly 
other related spp. Greatest diversity of fe'i bananas is 
on Tahiti. Total of 35 spp. in the genus, distributed 
throughout the tropics. 

Related species: Fe'i bananas are a significant source 
of food in New Guinea and the Pacific. M. textilis recent 
domesticate in Philippines used for Manila hemp. 
Related Ensete ventncosum cultivated in Ethiopia for 
starchy pseudostem. M. balbisiana produces edible 
fruit and contributed to present-day cultivars. 

Genetic base: About 500 genetically distinct cultivars. 



over from indigenous spp. Modern hybrids have a 
narrow genetic base Plantations are prone to severe 
pest and disease epidemics. Attempts to incorporate 
more genetic diversity is slowly having effect. Only 10 
percent wild germplasm used in breeding. S. robustum 
exhibits the most genetic diversity, but has had little 
application in breeding 

Breeding: Commercial varieties are derived from 
interspecific crosses with other wild and cultivated 
sugarcane spp. 

Germplasm collections: 19 000 accessions in total, 
nearly a quarter of them in Centra Nacional de 
Pesquisa de Recursos Geneticos e Biofecnologia, 
Brasilia, Brazil. 70 percent of landraces, 5 percent of 
wild spp. represented. 




90 percent of global banana production is from 
smallholdings International trade in bananas relies on 
very few cultivars, based on the Cavendish type. 
Dangerously narrow genetic base and very susceptible 
to diseases. Increased disease resistance is extremely 
important given the economic importance Of the export 
crop. The number of Fe'i banana cultivars has declined 
severely as a result of human demographic changes in 
the Pacific and the spread of pests. Banana is an 
aggressive weed. Wild populations of Musa benefit 
from forest clearance if succession is allowed to take 
place. Three spp. listed as threatened in 1997. 

Breeding: An extensive contact zone between cultivated 
and weedy types exists in several areas, e.g. Sri Lanka. 
Much mtrogression is believed to have enriched the 
gene pool of cultivated types. M. balbisiana has 
valuable traits. Several other wild relatives have useful 
characteristics. Germplasm collections have been 
poorly used; better selections could be made to suit 
subsistence farmers. 

Germplasm collections: Edible bananas, being 
seedless, are not storable. Seeds from wild spp. may 
be stored. Field gene banks hold collections. 10 500 
accessions in lotal. The International Network for the 
Improvement of Bananas and Plantains holds 10 
percent. Most of the diversity of wild and cultivated 
bananas is thought to be covered. 



APPENDIX 2 255 




Main uses: Seeds are fermented and roasted to 
produce cocoa powder and chocolate Waste goes lo 
produce animal feed, mutcfi or fertilizer Cocoa is a 
nutritional beverage: tfie powder is 25 percent 
saturated fat, 16 percent protein and 12 percent 
carbohydrates, fvlain producers are West African 
countries, Brazil, fvlalaysia. 

Origins; Upper Amazon basin. Center of cultivation in 
Central America. Total of 20 spp. in the genus, confined 
to tropical America. 

Related species: All the following are cultivated: 
T. grandiflorum Icupuacu, Amazonia): 
r. speciosum Icacaui, Central and South America!: 
T. subincanum (South Americal: T. obovatum (AmazonI: 
r. angus(/fo//um (Central Americal: T. b/co/or (Central 
and South Americal: T. glaucum (Amazonial, 

Genetic base: Undoubted genetic erosion has occurred 
In recent years. Currently cacao plantations are 



established by seed with varying degrees of genetic 
heterogeneity Production in West Africa is based on a 
particularly narrow gene pool. Originally three mam 
cultivated types. Criollo yields the most superior 
chocolate but has been largely replaced because of 
low yields. Forastero dominates world production. 
Wild cacao is highly variable, especially in its core 
area. Dramatic increase in plantations of coca and 
pulp-producing spp in various parts of the Amazon, 
agricultural expansion and movement of human 
populations have caused severe losses to the wild 
gene pool. 

Breeding: Little use of or research into wild genetic 
reserves because they are relatively hard to cross. 

Germplasm collections: Seeds do not remain viable for 
long, i 000 to 5 000 accessions kept in field gene 
banks. International Cocoa Genebank in Trinidad has 
the most comprehensive collection. Close relatives are 
poorly represented. Vegetative germplasm is collected. 



BEVERAGE CROPS 



256 WORLD ATLAS OF BIODIVERSITY 



CEREALS AND 
PSEUDO-CEREALS 



FOOD CROPS OF SECONDARY OR LOCAL IMPORTANCE 

These species and groups of species are those that, 
according to national food supply data maintained 
by the FAO' ■', provide a significant amount of the 
per capita supply of calories, protein or fat, but on 
the criteria followed here are not of equal 
importance to the crops in the previous table 



(i.e. they provide below/ 5 percent of the total 
per capita supply and/or do so in fewer than 
ten countries]. 

Source: List of crops from Prescotl-AUen and Prescott-Allen ; 
ottier information from Mabberley . Smartt and Simmonds . 
Smitti et 3l , Vauqtian and Geissler ; conservation status from 
Vifalterand Gillett , 



Oats 


■ 


Avena sativa IGraminael 


n 


Origin; West and north Europe from weed oat 


A. byzantina also cultivated. Genetic erosion from 


components of wheat and barley crops 


intensive breeding has resulted in efforts to 




conserve landraces and early varieties. Crosses 


One of the fnajor temperate cereals, althoiigh currently 


between A. sativa and A. byzantina have led to 


declining in production and generally regarded as a 


numerous cultivars. Fertile hybrids obtained from 


secondary crop, f^ostly used for animal feed. Oat 


crosses between cultivated oats and weed species. 


kernel is higher in high-quality protein and fat than 


Some success in incorporating desirable genes 


any other cereal. Oat bran is a good dietary fiber 


from more distant relatives. 



Quinoa 

Chenopodium quinoa IChenopodiaceae) 

Origin: High Andes 

An important and sacred pseudo-cereal in Inca times. 
Remains a staple in large parts of South America. 
Nutrient composition is superior to other cereals, being 
high in lysine and other essential amino acids, calcium, 
phosphorus, iron and vitamin E. Can grow in marginal 
conditions. Greatest diversity of genotypes in the 




highlands of southern Peru and Bolivia. Cultivation 
declined with Spanish conquest until 1970s when 
grown as a sole crop only in parts of Peru and Bolivian- 
Peruvian Altiplano. Agricultural and nutritional benefits 
have now been recognized and acreage has increased 
significantly Improvement so far has been based on 
inbred populations and pure lines. Considerable 
potential for improvement, both in the crop and its use. 



Fonio 

Digitaria exitis (Gramineae 



Origin: West Africa; thought to be a cultigen 




Popular cereal in parts of West Africa. Adapted to 
marginal agricultural land. Several species are 
harvested as cereals during times of famine. 



APPENDIX 2 257 



' Taro ^^^^^^ 


^^■^^^^■^H| 


Colocasia escutenta lAraceae) "^ 


^m^^^^i 


Origin: India 


products. Young leaves eaten as spinach. Tolerates high 




temperatures and poor soils. More than 1 000 cultivars 


Edible tuber; 25 percent starch, low protein, good 


have arisen through subsistence farming. Lack of 


vitamin C source. Probably cultivated before rice. Widely 


interest and germplasm exchange at a more 


cultivated in China and staple in many Pacific islands. 


commercial level. Serious danger of genetic erosion. 


Also used by food and beverage industries, and in pasta 





Carrot M 


^^^^^^^^^^^H 


Caucus carota (Umbelliferael '% 


^^^mmiiii 


Origin: Afghanistan 


replaced by hybrids in United States, Japan and 




Europe. Environmental health concerns over level of 


Root crop, grown worldwide, and eaten raw, cooked or 


pesticide has led to interest in genetic source of pest 


processed. The best plant source of provitamin A; low 


resistance. D. capiUifolius has passed some pest 


in other nutrients. Numerous wild and cultivated 


resistance to cultivated crop. 


subspecies. Open pollinated crops almost entirely 





Sweet potato 

Ipomoea batatas (Convolvulaceae 



Origin: Not known in the wild. Greatest species 
diversity occurs between Yucatan and the mouth of the 
Orinoco. Major variation is found in Guatemala. 
Colombia, Ecuador and Peru. 

The tuberous root is an important staple in the tropics. 
Able to grow in high temperatures with low water and 
fertilizer input. Good source of fiber, energy and 
vitamins A and C. Also industrial source of starch and 
ethanol. Although acreage has declined, increases are 
likely as a crop able to respond to population growth in 



Tannia 

Xanthosoma sagittifoUum (Araceae 

Origin: New World 




marginal areas. China accounts for 80 percent of 
production. Until recently, material used in breeding 
programs represented a fraction of existing diversity. 
Genetic base has now broadened but requires further 
increase. Little work has been done on cultivar 
improvement in areas of highest production li.e. where 
sweet potato is a staple). Countries where breeding 
programs exist have replaced native cultivars with 
improved varieties. Sweet potato is thought to have 
more potential for yield improvement than any other 
major crop in Asia. 




Similar use and nutritional composition as taro. but 
starch is more difficult to digest. Used in preparation of 
fufu in West Africa. 



ROOTS AND 
TUBERS 



258 WORLD ATLAS OF BIODIVERSITY 



BEANS AND OTHER 
LEGUMES 



Pigeonpea "^^^H 


IH^HI^^Hl 


Cajanus cajan (Leguminosae) ^^^| 


^mufii^mii 


Origin: Cultigen, India 


species wiffi good potential in agroforestry systems and 




on marginal lands, India contributes more than 90 


One of the major pulse crops of tlie tropics, feature 


percent of world production. Domestication has not 


seeds contain 20 percent protein, 60 percent 


altered the species as much as other crops. 


carbofiydrate and little fat. Important in small-scale 


C cajanifolius is closest relative;! 2 spp may be 


farming in mainly semi-arid regions, A multipurpose 


crossed with pigeon pea. 



Chickpea 

Cicer arietinum ILeguminosae 



Origin: Southeast Turkey; C. reticulatum is probably 
the progenitor 

One of the most important pulse crops. The seeds 
contain less protein 1 1 7 percent or morel but more fat 
15 percent! than other pulses. Grown over large area 
from Southeast Asia to Ivlediterranean, Only 2-i 




percent of world production is exported. Recently 
discovered C, reticulatum is confined to ten 
populations in Turkey Two main cuUivars have 
emerged. Traditional landraces have been selected to 
suit local ecological conditions. Commercial breeding 
is a recent phenomenon, C, reticulatum and C. 
echinospermum are compatible with the chickpea. 




Origin: Near East; wild progenitor L orientalis 

Seeds contain 25 percent protein. 56 percent 
carbohydrate and 1 percent fat. Young pods also eaten. 
Seeds are commercial source of starch for textiles and 



printing. Residues used as animal feed. Unique 
assemblages of landraces in different geographic 
regions. The crop has been altered little by modern 
breeding, ti4uch variation in the crop unexploited. 




Origin: Wild progenitor is unknown. Possible centers of 
origin are Ethiopia, the Ivlediterranean and Central Asia. 

The second most important pulse, 90 percent 
production as dried peas. Seed coats are source of 
protein, used in bread or health foods. Russia and 



China produce 80 percent of the world production of 
dried peas. United States and United Kingdom are 
largest producers of green peas. Breeding relies on a 
fairly narrow genetic resource base and efforts to 
conserve genetic variability of the cultivated crop have 
been fairly limited. 



APPENDIX 2 259 



Lupin 

Lupinus mutabilis ILeguminosael 



Origin: Andes; other lupins originated in two mam 
centers of genetic diversity in Mediterranean and in 
Americas 

A relatively minor pulse crop, obtained from several 
Lupinus spp. Seed contains U percent protein, 
17 percent oil Seed is human food in subsistence 
agriculture. Also used as coffee substitute and 




high-protein animal feed. Seed flour used as soya. 
Species also important in soil improvement Related 
spp. have ornamental value, used as fodder, coffee 
substitute, green manure or to stabilize sand dunes. 
May act as substitute for soybean, where climate is 
unsuitable for soybean growth. Other Lupinus spp. 
potentially suitable for cultivation. 




Broad bean 

Vicia faba iLeguminosae' 



Origin: Cultigen; wild ancestor unl^nown, possibly fronn 
central Asia 

A temperate pulse crop. Both immature seeds and dry 
mature seeds are eaten. The latter are also used as 



Mustard seed 

Brassicajuncea (Cruciferae) 



Origin: Central Asia: Himalaya; probably 6. n/gra x 
6. campestns and other Brsssica spp. 

The most important spice in the world in terms of 
quantity. Four species contributing to mustard exist. 



animal feed. Dried seed contains 25 percent protein, 
1.5 percent fat, 47 percent carbohydrate; the immature 
bean has much less of these nutrients but more 
vitamin A and vitamin C Also used as green manure. 




6. juncea took over from 6. nigra in 19505 as it allowed 
completes mechanization of harvesting. Also valuable 
as oil crop and salad crop, vegetable and fodder Long- 
lived seed allows easy maintenance of large 
collections. Wild material is widely distributed. 




Origin: Probably a hybrid of B. oteracea x 6. 
campestris 

The seed is an important, relatively recent source of oil, 
containing 40 percent unsaturated fat. with industrial 
and culinary applications. The root crop provides animal 
and human food Iswedel. Uncertain whether 6. napus 



exists in wild form. Domestication was a relatively 
recent event. The crop is tolerant of inbreeding, and 
landraces have been replaced by improved cultivars 
since the 19th century. Swedes, of which there are 
only a few varieties, are the result of hybridization with 
6- campestns- Various valuable contributions to oilseed 
rape also from 6. campestns and B. oteracea. 



BEANS AND OTHER 
LEGUMES 



OIL CROPS 



260 WORLD ATLAS OF BIODIVERSITY 



OIL CROPS 



LEAF 
AND FLOWER 
VEGETABLES 




Safflower seei 

Carthamus tinctorius ICompositae 



Origin: Turkestan, Turkey, Iran, Iraq, to Israel and 
Jordan 

Oilseed crop, produces two types of oil for margarine 
and also cooking oils. Ingredient of animal feeds. Dried 
flowers serve as substitute for saffron. Applications in 



cosmetic industry and as medicine. Originally 
domesticated for use as dye plant. Mucfi diversity 
developed as the species was cultivated over a wide 
area. Large-scale cultivation in few countries. No 
reported genetic erosion. Related species cross easily 
witli cultivated crop and form natural hybrids. 



Sesame ^^^^^^^^^^^^^^^^^^^^^^^^^^^H 


Origin: Origin and ancestors unknown, possibly 
Ethiopia or peninsular India 

An ancient oilseed crop. Seeds contain 50 percent 
unsaturated oil and 20-25 percent protein, and are 
used widely in bread and confectionery. Oil used in 


cooking, margarine, soaps and other industries. 
Residues are valuable animal feed. Interest in the crop 
is in decline as difficulty mechanizing harvesting and 
low seed yields compared with other oil crops. Good 
genetic diversity in related species. 5. malabaricum 
produces fertile offspring with S. indicum. 



Shea nut 

Viteilaria paradoxa ISapotaceael 

Origin: Dem. Rep. of Congo, Sudan, Uganda 

The roasted kernels are used to make purified shea 
butter, rich in vitamin E, used in cooking and as an 
alternative to cocoa butter for chocolate manufacture. 
Also has commercial use in manufacture of soap, 
cosmetics, candles. Various local uses. Fruit is eaten 



Artichoke 

Cynara scolymus ICompositae] 

Origin: Mediterranean. Canary Islands 



and is a good source of carbohydrates, iron and 
B vitamins. A monospecific genus. Threatened by 
overexploitation as a timber and source of fuel, and 
also by land clearance. Stands may be conserved for 
their valuable seed, but no official protection exists. 
Mostly grown for local consumption. 




Flowerheads and the receptacle are eaten. Small 
amount of vitamin C. 



APPENDIX 2 261 




Origin: Probably evolved in Asia Ivlinor or Middle East 
from L. sernota 

Lettuce leaves are a useful source of fiber, minerals 
lespecially potassium), vitamins A, E and C, Ivlay be 



grown year round. Stem is boiled as a vegetable in China. 
A flighty variable crop, resulting probably from long 
history of selection. Increasing diversity of lettuce types 
consumed. Wild species, including L sernola, L saligna 
and L. wrosa have been used in breeding programs. 




Origin: Southv^est Asia 



Edible leaves contain range of minerals, vitamins A, E 
and C and the B vitamin range. 



Pineapple 

Ananas comosus (Bromeliaceae 




Origin: Obscure origin in South America, probably on 
fringes of Amazon 

Seedless fiber-rich fruit, source of vitamins C. A and B. 
Highly suited for canning and as a juice. Unigue in that 
timing of harvest can be controlled by externally applied 
groviith hormone. Over 65 countries grow pineapple for 
domestic consumption and export. No wild populations. 



Genetically variable species, but genetic base of 
commercial plantations very narrow. 70 percent of world 
production and 96 percent of cannery industry comes 
from one variety. Highest diversity of near relatives in 
Paraguay and Brazil Poorly known, but A. ananassoides 
has contributed several characteristics to cultivated 
crop. A. erectilolius also considered for improvement 
programs. 




Origin; Obscure, probably hybrid of several Carica spp., 
arising in lowland tropical forest in eastern Andes or 
Central America 

Easily digested fruits produced all year round. Good 
source of vitamin C; red-fleshed fruits also rich in 
vitamin A. Papain extract is exported as a meal 
tenderizer; also used medicinally, to tan leather and in 
brewing beer May be produced by biotechnology in 
future Commercially produced in over 30 countries. 



mostly for domestic consumption. High diversity in 
eastern Andes. At least six other spp. domesticated and 
1 2 spp. are harvested for their fruit. Several commercial 
cultivars come from highly inbred hermaphrodite lines. 
Most production from backyard papaya trees, where 
local variation is high. Many wild species have desirable 
characteristics, useful in breeding. Hybridization already 
carried out with five wild Canca spp. Highly susceptible 
to viral and fungal diseases; some resistance detected in 
wild relatives but conventional crossing impossible. 



LEAF 

AND FLOWER 

VEGETABLES 



FRUITS 



262 WORLD ATLAS OF BIODIVERSITY 



FRUITS 



Lime, lemon, pomelo, tangerine, sweet orange 

Citrus aurantiifolia, C. limon, C. maxima, C. x paradisi, C. reticulata, C. sinensis (Rutaceae) 



Origin: Lime: cultivated tiybnd with obscure origins, 
possibly a hybrid of C. medica with another sp- Lemon: 
probably a hybrid of lime with C. medica. Pomelo: 
probably a native of the tvlalay peninsula. Grapefruit: 
probably hybrid between orange and pomelo, arising in 
the Caribbean, Tangerine: possibly Indo-Chma. Orange: 
probably introgressed hybrids of C. maxima and C. 
reticulata, perhaps originating in China. 

Fruits contain nearly 90 percent water, potassium, 
vitamins A, B, E and high vitamin C. They are eaten 



fresh, used as a flavoring and in marmalade. Orange 
accounts for 70 percent of C/(rus production. Various 
other spp. are cultivated. Wild populations located 
in northern India. Wild species threatened by forest 
clearance. In Southeast Asia wild groves are 
being replaced by oil palm and cacao plantations. 
C. taiwanica is critically endangered in Taiwan, mainly 
because of extensive habitat loss but also because of 
use as a rootstock for citrus plantations. Wide variation 
within the genus. Can be crossed with several genera. 
Economic Citrus spp. are highly interferlile. 



Pumpkin, squash, gourd 

Cucurbita moschata, C. maxima, C. argyrosperma, C. pepo, C. ficifolia ICucurbitae) 




Origin: C. moschata is most like the wild species and 
was domesticated independently in Central and South 
America. 

Fruits, containing 90 percent water, small amounts of 
starch, sugars, protein, fat and vitamins A, B and C, are 
used as vegetables and as animal feed. Leaves and 
flowers may be cooked. Seeds eaten and sometimes 
processed for oil. Grown worldwide in temperate and 
tropical zones, commonly in home gardens and as 



subsistence crops as well as commercially Long 
shelf life. Broad gene pool because of wide use of 
traditional or unimproved varieties in subsistence 
farming and home gardens. Many Cucurbita spp. have 
restricted geographical ranges. Disease resistance is 
found in wild relatives, with some transfer to cultivated 
species through interspecific crosses. Crosses between 
crop species and wild or feral relatives have occurred 
and genetic exchange takes place where their ranges 
overlap. 



Strawberry ^^^^^^^^^^^^^^^^^H 


Origin: A hybrid between two American species, F. 
chiloensis and F. virginiana 

Soft fruit, 90 percent water, high vitamin C, eaten 
fresh or in jams and confectionery. Grown in most 
temperate and subtropical countries. All Fragaria spp. 
produce palatable fruit. Hundreds of cultivars with 


wide ecological adaptability. Considerable genetic 
diversity lost in cultivated strawberries in the last 
1 00 years. Attempts are being made to extend the 
genetic base of the crop. Much unused genetic 
variation in wild species. 



APPENDIX 2 263 



M 




Origin: Eastern Mediterranean 

Fruits contain 1 percent sugar when fresh and 50 
percent when dried. Also substantial amounts of 
potassium, especially in the dried fruit. Ivlost world 
trade as dried figs. Figs are widely distributed in 
tropical, subtropical and warm temperate areas 
throughout the world. Ficus spp. are also source of 
rubber, fibers, paper, medicines and ornamental plants. 
Fig is largely grown for domestic consumption using 



traditional cultivars, hundreds of which exist, with local 
clones occurring in distinct geographical groups in the 
Mediterranean basin. Closely related wild forms are 
distributed throughout the Mediterranean basin. Fig 
culture is in decline. Many old groves have been 
abandoned or cleared. A number of wild relatives are 
considered threatened. 27 Ficus spp. were listed as 
threatened in 1998. Reproductive isolation is dependent 
solely on the specificity of the wasp pollinator Artificial 
crosses can be made between species. 




Origin: An aggregate of over 1 I 
and complex hybrid origin 



) cultivars. of ancient 



Apples, with pears, are the most important fruit crops of 
cooler temperate regions. Fruit is eaten fresh or cool<ed, 
as a |uice or brewed as cider Potassium is the mam 
mineral with small amounts of vitamin C. Breeders in 
the 19th century used wild species in breeding. Genetic 



diversity accumulated in North America was greater 

than in Europe because propagation was by seed 
rather than by grafting. Current trend depending on 
few varieties has caused rapid loss in genetic diversity 
and potential breeding material. Widespread elimination 
of wild stands is also talking place. Three spp. were 
listed as threatened in 1998, Hybridization with many 
wild species within the genus occurs readily. 




Origin: Northeast India 

Fleshy edible fruit; a good source of vitamins A and C. 
Thrives on infertile marginal soils. Important tree in 
Hindu mythology and religion. Kernel oil may be used 
in chocolate manufacture. Demand for the fruit and its 
juice IS increasing worldwide. India accounts for two 
thirds of production. Fruits of more than 12 wild spp. 
collected. Several are cultivated. The majority of fruit- 
bearing trees are more or less wild. Genetically highly 
heterogeneous. Over 1 000 cultivars exist, many in 



Borneo and the Malay peninsula. Feral populations 
are distributed throughout the tropics. Of the 40 to 
60 spp. in the genus, many are poorly l<nown, severely 
threatened or possibly extinct. 35 spp. were listed 
as threatened in 1998. Logging, forest clearance and 
replacement with commercial species in Southeast 
Asia are largely responsible for population extinctions. 
Various species are suitable for cultivation given 
further selection. Many display valuable traits, 
such as tolerance of waterlogged soils and more 
regular fruiting. 



FRUITS 



264 WORLD ATLAS OF BIODIVERSITY 



FRUITS 




Origin: Mexico to northwest Colombia 

A highty nutritious fruit, containing 15-25 percent 
monounsaturated fat and vitamins C. B and E- Ttie 
oil IS used in cosmetics. Trees fruit year round. 
Importance tias increased over recent decades and tfie 
crop IS now grown in most tropical and subtropical 
countries. Ivlost production is for domestic 
consumption. Other Persea spp. are used for timber 
and fruit. There are three geographically distinct 
varieties, which are able to interbreed. Commercially 
important cultivars have arisen mostly in private 



orchards by chance rather than as a result of 
germptasm manipulation. Increasing use of grafting 
and uniform varieties. Serious genetic erosion in 
traditional varieties. Diversity appears greater in 
traditional growing areas, where farmers still 
propagate by seed. Genetic exchange occurs between 
cultivated forms and wild populations. Wild populations 
of the avocado and its close relatives are small and 
becoming increasingly isolated. Deforestation poses a 
severe threat to their survival. 15 spp. were listed as 
threatened in 1 998. A number of wild relatives show 
resistance or tolerance to disease, drought and frost. 




Origin: Western India or Arabian Gulf 

Edible fruit with sugar content of 30 to 80 percent, 
corresponding to soft and dry dates. Vitamins are 
relatively low in quantity. Eaten as an ingredient in a 
variety of foods or as a juice. A staple where produced. 



Good storage. One of the oldest cultivated tree crops. 
Current cultivars resulted from thousands of years of 
selection. Perhaps over 3 000 cultivars exist; only 60 
grown widely All commercial cultivars are female. Wild 
populations of some related spp. are highly restricted in 
geographical range. All Phoenix spp. intercross freely. 



Apricot, cherry, plum, almond _^^^^^^^ 

Prunus armenica, P. avium, P. domestica, P. du/c/slRosaceae 




Origin: Apricot: west China. Cherry: west Asia. Plum: 
an ancient 6n cultigen with complex origin, possibly in 
southwest Asia; North American plums may be native 
American spp. or hybrids with P. saticina. Almond: 
central to west Asia. Peach: west China, possibly a 
cultigen derived from P. davidiana. 

Apricot, cherry, plum and peach are soft fruit with up to 
10 percent sugar, good potassium and Vitamin A in the 
case of apricots, but low vitamin C. Consumed fresh, 
dried or as an ingredient in jams and confectionery. 
Almond is the most important tree nut crop. The kernel 
contains '10-60 percent unsaturated oil and 20 percent 



protein. Eaten as a dessert nut and in confectionery and 
marzipan. A major trading commodity. Many other 
Prunus spp. have edible fruit. Many cultivars and much 
genetic diversity. Plums are genelically central to the 
genus and harbor the most useful genetic material 
Narrow breeding has led cherries to be more isolated 
from the rest of the genus. Increasing loss of genetic 
diversity Developing countries are tending to replace 
indigenous types and wild stands with western varieties, 
e.g. the switch from seed to vegetatively propagated 
almonds in Turkey A number of wild relatives are 
confined to narrow ranges. 23 spp. were listed as 
threatened in 1998. 



APPENDIX 2 265 




Origin: Asia minor, the Caucasus, central Asia and 
China. Cultivars have come from P bretschneideni, P. 
pynlolia, P. sinkiangensis and P. ussunensis- P. nivalis 
for perry production. 

With apples, pears are the most important fruit crops 
of cooler temperate regions. The fruit is eaten fresh or 
cooked, as a juice or brewed as perry. The fruit are a 
good source of dietary fiber, potassium and reasonable 
amounts of vitamin C. Currently about 20 spp. and 



5 000 recorded cultivars. li/lajor loss of genetic diversity 
through concentration on few varieties. Several wild 
species in Turkey are under threat. Five spp. were 
listed as threatened in 1998. Hybridization with high 
proportion of wild species within the genus is possible, 
providing useful rootstocks and possibly disease 
resistance. Much use of wild species in breeding in the 
past. Evolution of new varieties will be seriously limited 
unless stands of wild species conserved. 



Blackcurrants, redcurrants 

Ribes nigrum, R. rubrum (Grossularlaceae 



Origin: Europe and northern Asia, viiith the 
blackcurrant extending to the Himalayas 




Fruits with high vitamin C content. A luxury crop, 
largely produced for processing into juice. Many spp. 
with edible fruits, cultivated and wild. Wide use has 
been made of wild or near-wild relatives. 



Grape 


^^■■j^^^^^H 


Vitis vinifera (Vitaceael 


^BHill 


Origin: Eurasia 


commercial products include grapeseed oil and vine 




leaves. Various other spp. produce edible grape. One 


The fruit has high sugar content. 68 percent of grape 


estimate suggests there are 10 000 cultivars of grape. 


production is for the manufacture of wine. 20 percent 


Wild species still occurs in Middle Asia. Wild relatives 


for dessert grapes, 1 1 percent for dried fruit - raisins. 


are suffering genetic erosion m the United States. All 


sultanas, currants - 1 percent for juice. Other 


known Wdsspp. produce fertile offspring. 




Melon seed/watermelon 

CitruUus lanatus ICucurbitae) 



Origin: Tropical and sub-tropical Africa, domestication 
took place in Mediterranean 

The flesh of the fruit is 90 percent water; also contains 
vitamin C and A. The seeds contain 40 percent 



unsaturated oil and 40 percent protein. Wild plants still 
harvested in Kalahari. C. cotocynthis is fertile with the 
watermelon. An African watermelon with extraordinarily 
long storage life has been identified. 



FRUITS 



FRUIT 
VEGETABLES 



266 WORLD ATLAS OF BIODIVERSITY 



FRUIT 
VEGETABLES 



BEVERAGE CROPS 




Melon, cucumber 

Cucumis melo, C. sativus (Cucurbitael 



Origin: Wild melon populations appear to be distributed 
south of the Sahara to Transvaal in South Africa. 
Cucumber's wild or feral relative and possible 
progenitor, var hardwickii, is native to the southern 
Himalayan foothills. 

Melon IS grown worldwide in temperate and tropical 
countries, 90 percent water some sugar and vitamin C, 
Pink or orange-colored fruit have a high percentage of 
vitamin A, Also grown for their fragrance or ornamental 
value. Cucumber produces edible fruits, containing 96 



Tomato 

Lycopersicon escuientum ISolanaceael 

Origin: Cultigen. from Mexico 

There are few growing areas, from the tropics to the 
Arctic circle, where the tomato is absent. Fruit, 
containing potassium, vitamins A, B, C and E, is eaten 
fresh, dried or cooked as a vegetable, or processed in a 



percent water, some vitamin C and reasonable amounts 
of Vitamin A. Also used in production of fragrances, 
cosmetics and medicines. Young leaves and shoots may 
be cooked. Also cultivated C. angaria West Indian 
gherkinl and C. metuliferus (African horned cucumber 
or jelly melon). Wild and feral populations of melon 
occur throughout Africa and southern Asia. 
Cucumber produces fertile hybrids with its wild 
counterpart C. saf/vusvar hardwickii. No interspecific 
hybridizations have been used to improve crops. 




wide range of food products. Disease is a common 
threat. The wild relatives of the tomato have limited 
ranges. Wild gene pools are prone to erosion by habitat 
destruction. Tomato can be hybridized with all spp. in 
the genus and wild relatives have been used as source 
of numerous useful traits, including disease resistance. 



Eggplant 

Solarium melongena 




Origin: India; wild progenitor, 

S, incanum, occurs throughout Africa 

and Asia. 



Fruit IS eaten as a vegetable, contains over 90 percent 
water, large amount of potassium, some vitamins A, E. 
B. C. Highly productive and useful smallholders crop. 
Various spp. cultivated and used as grafting stock. 




Origin: Probably lower Tibetan mountains or central 
Asia 

Tender shoots are used to make tea. Important 
plantation and smallholder crop throughout the tropics 
and subtropics Planted commercially in at least 30 
countries. Increasing consumption in developing 



countries. High diversity of forms or species in East and 
Southeast Asia. Ivlany distinct forms, hybrids and species 
continue to be discovered. Recent trend to propagate 
the plant vegetatively, which has led to large areas being 
planted with one or few clones. No threat yet of genetic 
erosion in the crop. 1 1 Camellia spp. in China and Viet 
Nam were recorded as threatened in 1998. 



APPENDIX 2 267 




Origin: Arabica coffee originated in montane forest in 
southwest Etfiiopia and the neighboring Boma Plateau 
in Sudan; possibly also in Marsabit forest in Kenya, 
Robusta grows wild in West and Central Africa. 

Important sources of foreign currency in many 
developing countries. Roasted seeds used for beverage 
containing 1-2.5 percent caffeine, and niacin and 
potassium. A number of other Collea spp. also 
cultivated as coffee or as a source of edible berries. 
Commercial arabica cultivars have very narrow genetic 
base, especially in Latin America and the Caribbean. 



Mate 

//ex paraguariensls 

Origin; South America , 

Tea IS made from leaves, contains 2 percent caffeine. 
Little use in export market. 



and are highly susceptible to disease. Robusta 
outcrosses and has wider variation, t^luch production 
by smallholders, but iO percent of coffee from 
Americas and Caribbean from intensive monocrop 
plantations. Recent recognition of importance of 
conserving species-rich shade coffee systems. 
Signihcant percentage of Ethiopian coffee from uniform 
commercial cultivars; 400 000 ha remain of wild coffee, 
accounting for half of Ethiopia's coffee production. 
Several populations of wild species increasingly 
restricted in distribution and fragmented. Nine spp. 
from mainland Africa were listed as threatened in 1998. 




^^■^—■■—■ffi II in H 

Onion, garlic 

Allium cepa, Allium sativum (Alliaceaei 



Origin: Onion exists only in cultivation; may have come 
from Afghanistan, Iran, and former USSR area. 
Possible progenitors of garlic are A lonqicuspis or wild 
A. ampeioprasum. The greatest number of AUium spp. 
are in North Africa and Eurasia. 

Underground bulbs are more important for their flavor 
and antimicrobial properties than nutritional value. 
Garlic contains large amounts of potassium and 
significant vitamin C. Important component in the diet 
of a wide range of cultures. Numerous medicinal 




functions. The allins contained in Allium spp., 
especially garlic, may protect against cancer and 
cardiovascular disease. Seven economically important 
cultivated Ailium spp.; many other species consumed 
on a lesser scale. Open pollinated populations still 
represent most of the production in tropical and 
subtropical countries. The habitat of some wild Album 
spp. IS severely threatened. Poor results from 
interspecific hybridizations The genetic variability 
available in wild and cultivated relatives has not been 
extensively used in crop improvement. 



BEVERAGE CROPS 



SPICES AND 
FLAVORS 



268 WORLD ATLAS OF BIODIVERSITY 



SPICES AND 
FLAVORS 



Chili pepper, sweet pepper 

Capsicum annuum (Solanaceael 



Origin: Tropical America 

Fruits of varying pungency are used either fresh as a 
vegetable or dried or powdered as a spice. Fresh fruits 
contain large quantities of vitamin A, plus vitamin C. 



Cardamom 

Elettaria cardamomum IZingiberaceael 

Origin: India 



Seeds used as a spice. Essential oil used in perfume 
and as flavor for liqueurs. Wild populations exist in 




Breeding has generally depended on pure lines. Wild 
peppers are still collected and sold locally. Some 
interfertility with other Capsicum spp. Wild spp. offer 
valuable new traits. 




monsoon forests in south India and Sri Lanka. There 
are no essential differences between wild and cultivated 
forms. Collection from the wild contributes to the 
commercial trade. Wild populations are disappearing. 




Pimento 

Pimenta dioica IMyrtaceae 



Origin: West Indies and Central America 

Plants grow in semi-wild state. Populations of wild 
relatives are confined to small areas of remaining dry 



forest and coastal habitat in the Caribbean, especially 
Cuba. They are poorly studied and are severely 
threatened by habitat loss. 




Origin: Western Ghats, India 

The dried fruits, high in alkaloid content, represent one 
of the oldest spice crops. Several other Piper spp. 
important for local pepper production. Grown as a 



smallholder crop in tropical countries. Large-scale 
planting is based on one clone and is dangerously 
vulnerable to disease. Wild pepper still grows in the 
Western Ghats. 



APPENDIX 2 269 




Origin: Evolved from sea-beet (fi. vulgaris V3r. 
maritima] in Europe and west Asia- 
Swollen taproot provides nearly half tfie world 
production of sucrose. Forms of ttie same species 
include leaf beets and ctiards used as garden 
vegetables, and other beets with swollen taproots, eg. 



beetroot and mangold, for human consumption and 
animal feed. All forms within the species may be 
crossed. Wild relatives have already provided some 
disease resistance. The only source of resistance 
against the beet cyst nematode is detected in relatives 
from a different section in the genus. 




Origin: Europe and western Asia 

Edible tree nut and ornamental. Kernels are used in 
confectionery; 18 percent protein and 68 percent oil. All 



Corytus spp. have edible nuts. C. colurna is also 
cultivated for nuts. Populations of C. chinensis in China 
have declined, largely because of overexploitation. 



Walnut 


m 


Juglans regia Uuglandaceae) 


1 


Origin: Balkans to China 


timber, ornamentals. No apparent threat of genetic 




erosion in the crop. Major cultivation in the United 


Edible nut. containing vitamins E. C and B. Use as 


Stales but enormous unexplored potential elsewhere. 


dessert and in confectionery; oil also extracted. The 


Wild walnut forest has declined and become 


l<ernel contains 15 percent protein and 70 percent 


fragmented throughout its native range. Of the 21 


unsaturated oil. Leaves matte good fodder The timber 


species in the genus, seven were listed as threatened 


IS highly valued. All Juglans spp. produce edible seeds. 


in 1997. 



Pistachio 

Pistacia vera (Anacari 




Origin: Near East and western Asia 

Tree nut: low in sugar, more than 20 percent protein. 
50 percent oil. Important food for nomads during 
migration in Iran and Afghanistan. Highly drought 
resislant. Trees used for ornamental and shade 
purposes, also as a source of resin, dye. turpentine, 
mastic and medicine. Cultivated in the Mediterranean 
and western Asia for 3 000-4 000 years. None of the 



related species has value as a nut crop, although seven 
spp. are used as rootstocks and also for pollination. 
Largely harvested from wild in Afghanistan and parts 
of Pakistan. Iran has had commercial plantations for 
hundreds of years. Wild species may have a role in 
future improvements. Many wild populations have 
been destroyed by forest clearance, over-cutting 
for charcoal and grazing. Three spp. were listed as 
threatened in 1998. 



SUGAR CROPS 



TREE NUTS 



270 WORLD ATLAS OF BIODIVERSITY 



TREE NUTS i 



Brazil nut 

Berthotletia excelsa ILecythidaceael 

Origin: Tropical South America 



Kernel contains 1 7 percent protein, 65-70 percent 
monounsaturated oil. Largely an export crop. Also a 
staple for indigenous people and important ecological 
component of rainforest in the Amazon basin. Oil used 
for cooking or as fuel or animal feed. Valuable timber 
Attempts to establish plantations have generally failed 
Well-managed plantations have the potential of 
producing yields far exceeding natural groves. Almost 
all nut production is from wild trees. Distribution and 
density of groves may have been largely influenced by 




indigenous groups in the past. Little information exists 
on genetic variation. Populations appear to tolerate 
different soil types. Sustainable system of harvesting in 
extractive reserves, but considerable habitat loss and 
illegal tree felling continues elsewhere. Development in 
the Tocantins valley where there is high concentration 
of brazil nut trees continues to cause population 
decline. Developments elsewhere are also resulting in 
serious genetic losses. The species was listed as 
vulnerable in 1998. Protected populations are found in 
biological reserves, Indian and extractive reserves and 
corporate property 



REFERENCES 

1 Prescott-AUen, R. and Prescott-Allen, C. 1990. How many plants feed the world? 
Conservation Biology A{A]: 365-37ii. 

2 FAQ. FAOSTAT database. Available at Food and Agriculture Organization of ttie United 
Nations website, http://apps.tao.org/ (accessed February 20021. 

3 Mabberley, D.J. 1997. The plant-book. A portable dictionary of ttie vascular plants. 2nd 
edition. Cambridge University Press, Cambridge. 

4 Smartt, J. and Simmonds, N.W. 1995. Evolution of crop plants. 2nd edition. Longman 
Scientific and Technical, Harlow. 

5 Smith, N.J.H. et al. 1992. Tropical forests and their crops. Cornell University Press, Ithaca 
and London. 

6 Vaughan, J.G and Geissler, C.A. 1997. The new Oxford bool< of food plants. Oxford 
University Press, Oxford. 

7 Walter, K.S. and Gillett, H.J. (eds) 1998. 1997 lUCN Red List of threatened plants. 
Compiled by the World Conservation Ivjonitoring Centre. lUCN-the World Conservation 
Union, Gland and Cambridge. 



APPENDIX 3 271 



APPENDIX 3: 
DOMESTIC LIVESTOCK 

This table presents information on the major 
domestic mammals used in agriculture and 
related activities, such as hunting or transport, 
and on the number and status of closely related 
wild species. 

At the local level, a great many wild animal 
species are used primarily to meet subsistence 
needs, the kind depending largely on availability, 
convenience and tradition. Far fewer are used in 
livestock production: breeds of goat, sheep, cattle 
and pigs are cosmopolitan in distribution and, 
along with domestic fowl, form the basis for most 
of the world's agricultural animal food production 
on land. These four principal mammalian 
livestock species have diversified into more than 
4 000 recognized breeds. While intensification of 
production has typically gone hand in hand with 
narrowing of the genetic base, such that semen 
from individually documented and tested lines 
commands a premium, there is increasing 
recognition of the genetic potential resident in 
less commercially developed breeds and blood 
lines, and of the often neglected value of locally 
adapted stock in comparison with commercial 
stock from advanced industrial countries. The 
pool of genetic resources represented by 
domestic animal diversity is an essential basis for 
efficient and sustainable food production, and is 
likely to be of increasing importance in the more 
demanding production environments. 



There is no universally accepted system for 
naming domestic stock. Some authorities 
apply the earliest valid name to both the 
wild species and to domestic stock derived from 
it; others prefer to retain separate names for 
domestic stock where such a name has been in 
common use, and apply the next available valid 
name to the wild species. In the first case, for 
example, Capra hircus Linn. 1758 would be 
applied to the wild goat and all domestic 
derivatives; in the latter case, that name would be 
restricted to domestic stock and Capra aegagrus 
Erxleben 1 777 applied to the wild goat of Eurasia. 
The second approach is adopted below. 

In 'Number and status of breeds' the 
first figure is the number of breeds given in the 
Food and Agriculture Organization of the United 
Nations database. Subsequent figures are, in 
order: the number of threatened breeds in the 
critical' category, the number endangered' and 
the number 'extinct'. See Scherf for definitions. 
For both threatened categories, the counts 
include breeds maintained by active conservation 
programs and by institutions. 



Source: For general intormation. see Clutton-Brock' and Mason^; 
for number and status of breeds see Scherf^; for number of 
congeneric wild species see Wilson and Reeder*; and for status of 
wild species see HiUon-Taylor^. 



272 WORLD ATLAS OF BIODIVERSITY 



Bos taurus Ihumpless, mainly European cattle, 

taurine type] 

Bos indicus Ihunnped, mainly Asian cattle, zebu type! 

IBovidael 

Meat, milk, transport, draught, dung. etc. 

Domestic longhorn cattle from around 8 000 years 
ago at several Middle East sites, later in Nile region, 
and circum-Mediterranean by 3 000 years before the 
present, European cattle probably of Middle East 
origin. Humps, assumed result of artificial selection, 
at base of neck or over shoulder in zebu type. Zebu 
generally heat and parasite resistant, dominant in 
Asia and Africa Isome longhorns persist, e.g. 
trypanosome-resistant N'Dama in West Africal. 
Cattle were first draught farm animals; in Europe 



only specialized for meat or nnilk when replaced as 
power source by horse. Very high breed diversity, 
many now rare. British breeds to North America and 
Australia in the IVth century; Iberian breeds earlier 
to South America, Cattle certain to continue as major 
farm animals for meat and milk. Much potential In 
tropics for development of local stock, e.g. zebu dairy 
breeds. Several feral herds. 

Number and status of breeds: U79; C 106, E 193, 
Ex 255 

Derived from: 60s pnmigenius wild ox or aurochs 
lextinctl 

Congeneric wild species: i; all 4 listed as 
threatened in 2000. 





Bos frontalis iBovldae] 


freely in forest during day or for months, restrained 




at intervals, lack human control over breeding. May J 


Ceremonial sacrifice, barter 


breed with cattle and gaur j] 


No firm evidence but probably of early origin. 


Number and status of breeds: None formally \ 


Restricted to Bhutan, hills in northeast India 


recognized '.t 


bordering China and Myanmar, and Chittagong hills 




of Bangladesh. Typically higher elevation than cattle 


Derived from: Bos gaurus gaur. South and ' 


and lower than yak. Kept mainly by hill tribes. 


Southeast Asia 


usually by men of high status, for use in ceremonial 




sacrifice, exchange and trophy display. Not much 


Congeneric wild species: 4; all 4 listed as 


used for draught or milk. Mithan generally forage 


threatened in 2000 las for domestic cattle].' 



Bos grunniens IBovidael 

Milk, transport, meat, dam of 'dzo' Icattle x yak 
hybrid draught animal] 

Possibly domesticated at same time as cattle, 
probably on Tibetan plateau or the Himalayas. Most 
yak in west China, many in Mongolia, fewer in 
Tajikistan. Kyrgyzstan, Nepal, Bhutan. Afghanistan, 
India. Usually at 3 000-5 000 meters altitude- 
Variable size and pelage, usually smaller than wild 
yak. Yak tail in trade for centuries; white tips favored 
for ease of dyeing. Yak can graze where other 



livestock cannot. Much medical or religious use in 
Tibet, where milk and butter most important; used 
as meat source in Mongolia. Hair used for rope, felt; 
skin for leather; dung for fuel; important pack 
animal. 

Number and status of breeds: 13; C 0, E 1. ExO 

Derived from: Bos mulus yak, China: north of Tibet 
plateau lAltun Shan. Qilian ShanI 

Congeneric wild species: 4; all 4 listed as 
threatened in 2000 las for domestic cattlel. 



APPENDIX 3 273 





Bos javanicus [Bowdae] 


fertile, little fat, uses poor pasture in hot humid 




conditions. Good draught animal for small fields and 


Draught, meat 


terraced slopes; much potential as meat or crossing 




stock. Feral herd in Cobourg Peninsula lAustralial. 


Domestic cattle present in Souttieast Asia ca 5 500 




years before present, Banteng possibly domesticated 


Number and status of breeds: No data 


in prehistory in Southeast Asia or Java. Now in many 




parts of Indonesia; small herds in Malaysia, 


Derived from: Bosjavanicus banteng, Southeast Asia 


Philippines, Australia. Uniform in type. Organized 




selection in 20th century: no entire males exported. 


Congeneric wild species: 4; all i listed as threatened 


no crossing with other cattle. Small size, highly 


in 2000 las for domestic cattlel. 





Bubalus bubalis IBovidael 


for draught, and river buffalo in South Asia, 




mainly for milk. Do better than cattle on swamp and 


Draught, milk 


lloodplam grazing. Much potential for development 




as meat producer. Milk rich in fat. Large feral herds 


Probably domesticated earlier than 4 500 years ago 


in Australia. 


in Middle East. Wild ancestor occurred from 




Mesopotamia east to Southeast Asia; by the 19th 


Number and status ot breeds: 86; C 3, E 8, Ex 


century restricted to India and adjacent areas, where 




local. Domestic buffalo reached southeast Europe by 


Derived from: Bubalus arnee wild water buffalo. 


12th century where from Uth century much used in 


Bhutan, India, Nepal. Thailand 


Muslim communities; later taken to the Americas 




and Australia, and Africa in 20th century. Breed 


Congeneric wild species: 4; alU listed as 


development centered in India and Pakistan. Broadly 


threatened in 2000. 


divided into swamp buffalo in Southeast Asia, mainly 







Capra hircus IBovidael 


where often adverse impact on native biota. Much 




potential for further breed development, e.g. for 


Meat, milk, hair 


specialized tropical dairy animals. Ruminant 




physiology allows efficient use of coarse vegetation 


Goats and sheep next to be domesticated after dog. 


in semi-arid and and regions. 


Domesticated around 10 000 ago in Zagros 




Mountains of western Iran; to Europe by mid- 


Number and status of breeds: 587; C 31 , E 70. Ex 17 


Neolithic. Worldwide distribution. Great variety in 




form of horns and ears, hair color, etc. Highest 


Derived from: Capra aegagruswild goat, southwest 


numbers in South Asia. Milk breeds developed in 


Asia: Turkey east to Pakistan 


Switzerland have influenced many milk breeds 




worldwide. The Boer ISouth Africal is major meat 


Congeneric wild species: 9; 7 listed as threatened in 


breed. Two fleece breeds: angora ITurkeyl and 


2000. 


cashmere Icentral Asial. Many feral populations. 





274 WORLD ATLAS OF BIODIVERSITY 



Ovis anes IBovidael 

Meat, milk, wool 

Sheep and goats next to be domesticated after dog. 
Sheep in use in Mesolithic; evidence for 
domestication around 1 1 000 years ago in Middle 
East; to North Africa (where no wild sheepi by 6 000 
years ago; to Annericas in 16th century. Worldwide 
distribution; important in Europe, Middle East. 
Central Asia. Coat of wild sheep has outer hairs over 
woolly inner coat; hairs lost during domestication to 
produce fine fleece breeds. Wool and mill< often 
more important than meat. Wool trade basis of great 
wealth in medieval and early modern Europe. Many 
breeds: some multipurpose, others specialized for 
milk, fleece or meat. Sheep numbers in decline in 



some developed countries e.g. United States and 
Australia, but elsewhere provide vital support to 
human life in marginal and rangeland environments. 
Ruminant physiology allows efficient use of coarse 
vegetation in semi-arid and arid regions. 



Number and status of breeds; 1 '195; C t 
181 



3, E 1V9, Ex 



Derived from: Ovis onentslis mouflon, southwest 
Asia: Turkey east to Iran; Mediterranean populations 
(Corsica, Sardinia, Cyprus! possibly feral primitive 
domestic stock 

Congeneric wild species; 6; 4 listed as threatened in 
2000. 





Bus domesticus (Suidael 


Production increasingly specialized, but still an 




important role for local varieties in utilizing liouseliold 


fvleat 


waste and wild foods. Pigs have a major cultural 




significance in parts of Southeast Asia and Melanesia. 


First evidence of domestic pigs by 9 000 years ago in 




Anatolia; widespread in Eurasia, incl. Egypt, by 5 000 


Number and status of breeds: 6't9; C 58, E 1 06. Ex 


years ago. Worldwide; nearly half the world's pigs 


151 


occur in non-Muslim Asia, mostly in China. 




Management varied: may free-range in woodland or 


Derived from; Sus scrofa, Eurasian wild pig, North 


be sty-fed. Pigs introduced to the Americas from 


Africa. Europe, Asia 


Europe; few in Africa or Australia, New Zealand. 




Several feral herds. Large number of breeds. 


Congeneric v»ild species: 10; 6 taxa listed threatened 


Commercial production now dominated by few lines. 


in 2000 



Came/us bactnanus (Camelidael 

Draught, transport, meat, milk, wool, dung 

Fossil camels known from North America (where no 
extant camels! and Eurasia west to North Africa. 
Rock drawing in Mongolia of two-humped camel may 
be 10 000 years old. First evidence of domestication 
in Iran and Turkmenistan about 5 000 years before 
the present. Widespread in central Asia by ca 3 000 
years ago. Mam transport on Silk Route between 
Mesopotamia and China but replaced by dromedary 
in west and south from ca 2 000 years ago. 



Restricted to Central Asia, incl. Mongolia and China. 

Numbers probably in decline. 

Number and status of breeds: 1 1 ; C 1 , E 1 , Ex 

Derived from: A domesticated form of wild Bactrian 
camel, southwest Mongolia, northwest China 

Congeneric virild species: Non-domesticated 
populations in central Asia, listed threatened 
(endangered) in 2000. 



APPENDIX 3 275 



Camelus dromedanus ICamelidael 

Transport Idraught, meat, milk, wool, dung) 

Remains of dromedary or similar species at 
Paleolithic sites in North Africa about 80 000 years of 
age. Wild camels apparently extinct in Africa by 5 000 
years ago but persisted in Saudi Arabia Iwhere 
perhaps first domesticated! until ca 2 000 years ago. 
Domestic camel to Horn of Africa around 4 000 years 
ago. Reached present importance with rise and 
spread of Arab power from 7th century onwards. 
Most camels in northeast Africa and Afghanistan/ 
Pakistan/India, where numbers rising; fewer and 



Lama glama ICamelidael 

Transport, wool Icoarsei. meat, dung 

Domesticated by b 000 years before the present In 
high-altitude Andean pastures, possibly centered 
around Lake Titicaca basin of south Peru and west 
Bolivia, Alpaca textiles known from 2 500 years ago. 
Domestic camelids spread to lower altitudes and 
along Andean chain by i 000 years ago and reached 
greatest extent during Inca period; in decline since 
Spanish conquest in early 16th century and 
introduction of European stock. Remain important to 
Andean culture and for superior adaptation to poor 



Lama pacos ICamelidael 

Wool Ifinel 

See llama for background. 



decreasing in Middle East, Primarily for transport; 
specialized pack and riding breeds exist. Introduced 
to Canaries and Australia (where feral herds]. Ability 
to withstand long periods without drinking and use 
thorny browse key to human use of hot deserts. 

Number and status of breeds: 52; CI . E 2. Ex 

Derived from; Ancestral form unknown, presumed 
extinct Camelus species; Bactrian camel closest 
living relative 

Congeneric VKild species: See Bactrian camel. 



high-altitude grazing. Pad feet may cause less 
pasture damage than hoofs of sheep. Llamas and 
most alpacas held by small-scale pastoralists on 
communal grazing; some alpaca kept in large herds 
by cooperatives In Peru, Not milked. Alpaca wool has 
high commercial value. Llama flocks in the United 
States and Europe, 

Number and status of breeds: 8; C 0. E 0. Ex 

Derived from: Probably Lama guanicoe guanaco, 
south Peru, west Bolivia, northwest Argentina 

Congeneric wild species: 1; not listed threatened. 



Number and status of breeds: 6; C 0, E 1 . Ex 

Derived from: Unknown in wild, presumed Lama sp. 
or Lama x Vicugna hybrid 

Congeneric wild species: See llama. 



276 WORLD ATLAS OF BIODIVERSITY 





Rangifer tarandus ICervidaej 


potential tor better use; numbers have been 




increasing but with local indications of overgrazing. 


Meat, milk, transport 


Lichens, the mam winter feed, are vulnerable to 




atmospheric pollution. 


Fossil evidence for use of reindeer from 80 000 years 




ago, domesticated by 2 500 years before ttie present- 


Number and status of breeds: No data 


Management varies: riding or miilk animals may be 




separated from herd and fed, or herds may roam 


Derived from: Semi-domesticated forms of Eurasian 


widely and be gathered annually for marking or 


Rangifer tarandus. reindeer lEurasial, caribou INorlh 


slaughter Reindeer industry important in north 


America! 


Scandinavia, northwest Russia and Siberian Russia, 




less so in North America, Reindeer exploitation key 


Congeneric wild species: See above; \ North 


to settling the far north. Wild reindeer include four 


American subspecies listed threatened in 2000. 


major types, all used in husbandry systems. Some 





.JVggg r donkey 

Equusas/nuslEquidae) 

Transport, draught, sire of mule lass x horse hybrid! 

Probably domesticated in northeast Africa; records 
from about 6 000 years ago in Egypt. The only 
domestic animal certainly of African origin. 
Widespread in Middle East by ca 2 000 years ago. To 
Americas in 16th century. Much more important than 
horse in Africa where present in north and west. 
Common in south and central Asia; also present in 
south Europe. Mostly lor transport; specialized riding 
and pack breeds exist. Formerly milked, meat 



sometimes used. Feral asses widespread incl. 
Socotra, Galapagos, United States. Australia, Sahara, 
etc. Numbers worldwide likely to decline, but 
because of hardiness and low cost will retain 
importance in less developed areas. 

Number and status of breeds: 1 03; C 5, E 1 6, Ex 6 

Derived from: Equus afncanus, African wild ass. 
North Africa to Somalia 

Congeneric wild species: 9; 8 taxa listed threatened 
in 2000. 



Equus caballus lEquidae! 

Transport, draught, sport, dam of mule lass x horse 
hybrid! 

Some evidence of domestic horses 6 000 years ago in 
central Eurasia lUkraine!. Spread through Eurasia 
during Bronze and Iron Ages. Important early military 
use. to draw chariots and lor riding, especially after 
invention of stirrups before the start of the 6th 
century. Wild horses present with Amerindians in 
North America but extinct there by 12 000 years ago; 
domestic horses introduced by European colonists. 



Most horses occur in South America where numbers 
also highest in relation to humans; numbers high in 
North America and Asia. Specialized for draught or 
riding, but both uses in decline. Feral horses on all 
continents lexcept Antarctica!. 

Number and status of breeds: 820; C 1 27. E 1 78. 
Ex 94 



Derived from: Equus ferus If. przewalskiil wild 
horse, formerly Americas, Europe. Asia 

Congeneric wild species: See ass. 



I 

i 



APPENDIX 3 277 





Cams lamiliaris ICanidael 


Number and status of breeds: A few hundred, no 




data on status 


Companion, hunting, security, food 


Derived from: Cams lupus wolf. North America, 


Domestication may have begun long before that of 
agricultural stock, possibly 100 000 years ago; first 
direct evidence U 000 years ago in Middle East; 
distinct kinds of dog evident by 7 000 years ago. The 
dingo is a feral domestic dog taken to Australia 


Europe. Asia 

Congeneric wild species: 7; 3 taxa listed threatened 
in 2000. 


around 12 000 years ago. 







Cavia porceltus ICaviidael 


Number and status of breeds: No data 


Meat, laboratory, companion 


Derived from: Cavia aperea. widespread in 




South America, or C. tschudil Peru, south 


One of the few domestic animals of South American 


Bolivia, northwest Argentina, north Chile 


origin. Probably domesticated 3 000-6 000 years ago. 




but in use long before. Taken to Caribbean and Europe 


Congeneric wild species: i; 1 listed threatened 


by mid-16th century. Some planned selective breeding 


in 2000 


during past 30 years. Potential for more development 




as meat source, especially in original Andean range. 




but broiler fowl increasingly used instead. 


■BK 



Number and status of breeds: fO; C 0, E 4, Ex 

Derived from: Oryctoiagus cumculus European 
rabbit, west and south Europe to northwest Africa 

Congeneric wild species: None; a monospecific 
genus (several related genera listed threatenedl. 



Oryctoiagus cumculus iLeporidael 

Meat, fur, laboratory, companion 

Kept enclosed jin leporaria by Romans! for more 
than 2 000 years. Kept by medieval monks; newborn 
or unborn young were permissible food during Lent. 
Distributed worldwide by manners; many feral 
populations. Some development of meat breeds 
since the second world war; much potential as low- 
cost converter of surplus vegetation into meat. 



REFERENCES 

1 Clutton-Brock, J. 1999.4 natural history of domesticated mammals. 2nd edition. 
Cambridge University Press, Cambridge, and the Natural History Ivluseum, London. 

2 Mason, LL. led.) 1984. Evolution of domesticated animals. Longman, London and New York. 

3 Scherf, B.D. led.) 2000. World Watch List for domestic animal diversity. 3rd edition. 
Food and Agriculture Organization of tfie United Nations. Rome. 

4 Wilson, D.E. and Reeder, D.M. (eds) 1993. Mammal species of the world: A taxonomic and 
geographic reference. 2nd edition. Smithsonian Institution Press, Washington DC and London. 

5 Hilton-Taylor, C. (compiler) 2000. 2000 lUCN Red List of threatened species. lUCN-the 
World Conservation Union, Gland and Cambridge. Available online at 
http://www.redlist.org/ (accessed April 20021. 



278 WORLD ATLAS OF BIODIVERSITY 



APPENDIX 4: 

RECENT VERTEBRATE EXTINCTIONS 

This list provides information on vertebrate 
species thai have been listed as globally extinct. It 
is based in part on lists produced using the lUCN 
threat category system, but mammals, birds and 
fishes have been revised following more rigorous 
criteria developed by the Committee on Recently 
Extinct Organisms |CREO|. The OREO system 
relates to extinctions since AD1500. Where 
information meets all OREO criteria, the 
extinction event is considered fully resolved; these 
species appear on a darker background. To date, 
extinct and possibly extinct mammals and fishes 
have been evaluated in this way, and data are 
available at the CREO website. The bird list is 
based on Threatened Birds of the World which 
itself informally follows CREO guidelines. The 
reptiles and amphibians listed have been regarded 
as extinct, but in several cases data needed to 
meet CREO criteria le.g. on taxonomy or survey 
effort! appear to be unavailable. A number of 



species have been removed, as new evidence 
suggests that extant populations of that taxon 
persist or because extinction took place before 
AD1500. The Lake Victoria cichlid fishes in the 
genus Haplochromis [sensu lata] at the end have 
been widely regarded as extinct, but data available 
leave some uncertainty whether these species are 
extinct in the wild, or persist in small numbers 
outside sampled areas. These fishes are listed as 
questionably extinct and are distinguished by a 
paler background. The Period column indicates, 
very approximately, the probable period during 
which extinction occurred. 'Early' refers to the 
first four decades of a century ICl, 'mid' the next 
three decades, 'late' the final three decades. 
Where a more precise date is available, this is 
given in parentheses. 



Source: Data on mammals and fishes. CREO ; on birds, BirdLife 
International ; on reptiles and amphibians, Hilton-Taylor . 



Class MAMMALIA 








Order DASYUROMORPHIA 








Family Thylacinidae 








Thylacinus cynocephalus 


Thylacine 


Tasmania 


Early 2l)thC 119361 


Order PERAMELEMORPHIA 








Family Peramelidae 








Chaeropus ecaudatus 


Pig-footed bandicoot 


Australia 


Mid 20th C 


Macrotis leucura 


Lesser bilby 


Australia 


Mid20thCI1960sl 


Perameles eremiana 


Desert bandicoot 


Australia 


Mid20thCI19(i0sl 


Order DIPROTODONTIA 








Family Macropodidae 








Cabprymnus campestris 


Desert rat-kangaroo 


Australia 


Early 20th C 11932) 


Lagorchestes asomatus 


Central hare-wallaby 


Australia 


Mid 20th C1 19605) 


Lagorchestes leporides 


Eastern hare-wallaby 


Australia 


Late 19th C 118901 


Macropus greyi 


Toolache wallaby 


Australia 


Late 20th C 11972) _ 


Onychogalea tunala 


Crescent nailtail wallaby 


Australia 


Mid 20th C 11956) ■ 


Polorous ptatyops 


Broad-faced potoroo 


Australia 


Late 19th C 11875)^ 



Order INSECTIVORA 
Family Nesophontidae 
Nesophontes hypomicrus 

Nesophontes longirostns 



Atalaye island-shrew Hispaniola Early 16th C? "^ 

Long-nosed island-shrew Cuba Post-1500 



APPENDIX U 279 



I^^^H 




^^B 


■M^g 


Nesophontes mapr 




Cuba 


Post- 1500 


Nesophontes micrus 


Western Cuban island-shrew 


Cuba 


Post- 1500 


||[ Nesophontes paramicrus 


St Michel island-shrew 


Hispaniola 


Early 16th C ' 


Nesophontes submicrus 




Cuba 


Post-1500 


W Nesophontes superstes "^H 




Cuba 


Early 16th C? ; 


t Nesophontes zamicrus 


Haitian island-shrew 


Hispaniola 


Early 16th C? I 


Nesophontes sp. 1 


Grand Cayman island-shrew 


Grand Cayman 


Post- 1500 


Nesophontes sp. 2 


Cayman Brae island-shrew 


Cayman Brae 


Post- 1500 


Family Solenodontidae 








1 Solenodon marcanoi 


Marcano's solenodon 


Hispaniola 


Early 16th C | 


Order CHIROPTERA 








Family Molossidae 








E Mystacina robusta 


New Zealand lesser 
short-tailed bat 


New Zealand 


Mid 20th C (1965! ] 

i 


Family Pferopodidae 


* Dobsonia chapmani 


Dobson's fruit bat 


Negros 1. 


Late 20th C 






(Philippines) 


(1970s! 


Nyctimene sanclacrucis 


Nendo lube-nosed fruit bat 


Santa Cruz 
ISolomonsI 


Late 19th C (1892! , 


Pteropus brunneus 


Dusky flying fox 


Percy 1. 


Late 19lhC (1874! 


i Pteropus pilosus 




(Queensland. Australia! 


Large Palau flying fox 


Palau 


Lalel9lhC(1874! , 


Pteropus subniger 


Reunion flying fox 


Mauritius, 


Mid 19th C (1840s! ' 


I Pteropus tokudae 




Reunion 




Guam flying lox 


Guam 


Mid 20th C (1968! 


Family Vespertilionidae 








Kemoula slncana 


Dobson's painted bat 


Tanzania 


Pre-1878 


1 Pharolis imogene 


Large-eared nyctophilus 


Papua 
New Guinea 


Late 19th C (1890! ^ 

J 


Order PRIMATES 








Family Megaladapidae 








f Megaladapis cf. M, edwardsi 


Tretretretre 


Madagascar 


Early 16lhC? 


Family Paleopropithectdae 


|| Palaeopropithecus ingens 


Large sloth lemur 


Madagascar 


Early 16th C? | 


Family Xenotrichidae 


J Xenothrix mcgregori 


Jamaican monkey 


Jamaica 


Early 16th C | 


Order CARNIVORA 








Family Canidae 








1 Dusicyon australis 


Falkland island dog 


Falkland Is 


Late 19th C 11876| ^ 


'Sl 



280 WORLD ATLAS OF BIODIVERSITY 



^Kecres^^^^^^H 


' UvmnW^^^^^^^^^^^^^^I 




Perio^^^^B 


Family Mustelidae 


- 






Mustela macrodon 


Sea mink 


USA 


Late 19th C 118601 


Family Phocidae 


MonachuB Iropicalis 


Caribbean monk seal 


Caribbean Sea 


Mid 20th C 119505] 


Order SIRENIA 
Family Dugongidae 






1 


Hydrodamalis gigas 


Steller's sea cow 


Russia. USA 


Mid 18th C 117681 "■ 


Order ARTIODACTYLA 

Family Bovidae 


Gazella rufina 
Hippotragus leucophaeus 


Red gazelle 
Bluebuck 


Algeria 
South Africa 


Late 19th C 11894] 
Early 19th C 11800] 


Family Hippopotamidae 


Hippopotamus lemedei 


Malagasy dwarf hippo 


Madagascar 


Early 16th C 


Hippopotamus 
madagascanensis 

Order RODENTIA 
Family Capromyidae 
Capromys sp. 1 


Common Malagasy hippo 
Cayman hutia 


Madagascar 
Cayman Is 


Post- 1500 :m 
Post- 1500 ^ 


Geocapromys colombianus 
Geocapromys thoractus 


Cuban coney 
Swan Island hutia 


Cuba Little 
Swan 1. 
IHondurasI 


Early 16th C? W 
Mid20thCll950sia 


Geocapromys sp. 1 
Geocapromys sp, 2 


Great Cayman coney 
Cayman Brae coney 


Grand Cayman 
Cayman Brae 


Post- 1500 
Post-1500 


Hexolobodon phenax 
Isolobodo!) montanus 
Isolobodon portoricensis 
Plagiodontia ipnaeum 
Rtwzopiagiodontia lemkei 


Imposter hutia 
Montane hutia 
Allen's hutia 
Johnson's hutia 
Lemke's hutia 


Hispaniola 
Hispaniola 
Hispaniola 
Hispaniola 
Hispaniola 


Early 16th C? 
Early 16th C? _ 
Early 16th C? fl 
Early 16th C ■ 
Early 16th C? J 


Family Echiimyidae 
Boromys offetla 
Boromys torrei 


Cuban esculent spiny rat 

De la Torre's esculent spiny rat 


Cuba 

Cuba 


Post- 1500 1 

Post-1500 ' 


Brotomys voratus 


Muhoy 


Hispaniola 


Early 16th C? J 


Family Heptaxodontidae 
Quemisia gravis 

Family Mundae 


Quemi 


Hispaniola 


Post- 1500 


Coniiurus albipes 


White-footed rabbit rat 


Australia 


Late 19th C 11875] 


Craleromys paulus 
Leimacomys buetiner, 


llin bushy-tailed cloud-ral 
Groove-toothed forest mouse 


Philippines 
Ghana 


1953 

1890 


Leporilius apicalis 


Lesser stick-nest rat 


Australia 


Late 20lh 0(1970] ^ 





APPENDIX U 281 



|Bli^"™ 






W^^ 


Malpaisomys insulans 


Volcano mouse 


Canary Is 


Post- 1500 


Megalomys audreyae 


Barbuda giant rice rat 


Barbuda 
and Antigua 


Pre- 1890 


P Megalomys desmarestii 


Martinique rice rat 


Martinique 


Early 20th C (1902) 


Megalomys tuciae 


St Lucia giant rice rat 


Sainl Lucia 


Late 19th C (18811 


Megaoryzomys cunoi 
L 


Curio's giant rice rat 


Santa Cruz. 
Galapagos 


Mid 18th C (1740) ' 


Megaoryzomys sp 1 


Isabela giant rice rat 


Isabela. Galapagos 


Post- 1500 


1 Nesoryzomys darwini 


Darwin's rice rat 


Santa Cruz, 


Mid 20th C 11940s) 


1 




__J3alapagos 




Nesoryzomys sp 2 


Isabela Island nee rat B' 


Isabela. 
Galapagos 


Post- 1500 


Noronhomys vespuccii 


Vespucci's rice rat 


Fernando da 
Noronha 1. (Brazil) 


Early 16th C : 


Notomys amplus 


Short-tailed hopping-mouse 


Australia 


Late 19th C (1896) 


^ Notomys bngicaudatus 


Long-tailed hopping-mouse 


Australia 


Early20thC(19Gl) 


Notomys macrotis 


Big-eared hopping-mouse 


Australia 


1843 


Notomys mordax 


Darling Downs hopping-mouse 


Australia 


1846 


Notomys sp 1 


Great hopping-mouse 


Australia 


Pre- 1900 


1 Oligoryzomys victus 


St Vincent pygtITy rice rat 


Saint Vincent 


Late 19th C (1892) 


1 Oryzomys antillarum 


Jamaican rice rat 


Jamaica 


Late 19th 0(1877) 


Oryzomys liypenemus 


Barbuda rice rat 


Barbuda 
and Antigua 


Post- 1500 


f Oryzomys nelsoni 


Nelson's rice rat 


Maria Warire 1. 
(Mexicol 


Late l9thC (1897) ; 


Oryzomys sp. 1 


Barbados nee rat 


Barbados 


Pre- 1390 


W Peromyscus pembertoni 


Pemberton's deer mouse 


San Pedro 
Nolasco 1. [Mexicol 


Early 20th C (1931) 


Pseudomys gouldii 


Gould's mouse 


Australia 


Early 20th C (1930) 


Rattus madeari 


Maclear's rat 


Christmas 1. 


Early 20th 01903) 


■ 




lAustralial 




' Rattus nativitatis 


Bulldog rat 


Christmas 1. 
[Australial 


Early20thC (19031 


Uromys imperator 


Giant naked-tailed rat 


Guadalcanal 


Mid 20th C (1960s) 






(Solomons! 




Uromys porculus 


Little pig rat 


Guadalcanal 


Late 19th C (1887) 






(Solomons) 




Order LAGOMORPHA 








Family Leporidae 








Sylvilagus insonus 


Omilteme cottontail 


Mexico 


Late 20th C (1991) 


Family Ochotonidae 


f Prolagus sardus 


Sardinian pika 


Corsica, 
Sardinia 


Late 18lhC (1777) 





282 WORLD ATLAS OF BIODIVERSITY 



||M|^|pHHHi^^pHI^^^^B 


Class AVES 




1 


■■^H 


Order CASUARIIFORMES 






^^^^^^^H 


Family Dromaiidae 






-^^^^^^^H 


Dromaius ater 


King Island emu 


King 1. 
(Australia! 


Early 19th C fl 


Dromaius baudmianus 


Kangaroo Island emu 


Kangaroo 1. 
(Australia] 


Early 19th C 


Order PODICIPEOIFORMES 








Family Podicipedidae 








Podiceps andinus 


Colombian grebe 


Colombia 


Late 20th C (1977! 


Podifymbus gigas 


Atitan grebe 


Guatemala 


Late 20th CI1986! 


Order PROCELLARIIFORMES 






M 


Family Procellariidae 






9 


Bulweria bifax 


St Helena bulwer's petrel 


St Helena 


16th C M 


Oceanites maorianus 


New Zealand storm petrel 


South 1. 
(New Zealand! 


^■fl 


Pterodroma rupinarum 


St Helena gadfly petrel 


St Helena 


16th c ^^m 


Order PELECANIFORMES 






^H 


Family Ardeidae 






^^B 


Ixobrychus novaezelandia 


New Zealand little bittern 


New Zealand 


Late 19th C? ■ 


Nycticorax duboisi 


Reunion night-heron 


Reunion 


Late 17th C? 


Nycticorax mauritianus 


Mauritius night-heron 


Mauritius 


Early 18th C? 


Nycticorax megacephalus 


Rodrigues night-heron 


Rodrigues 


Mid 18th C 


Family Phalacrocoracidae 








Phalacrocorax 


Pallas's cormorant 


Bering Straits 


Mid 19th C(1850s| 


perspiciUatus 




(Russia! 


m 


Family Threskiornithidae 






1 


Threskiornis solitahus 


Reunion fhghtless ibis 


Reunion 


Early 18th C ■ 


Order ANSERIFORMES 






m 


Family Anatidae 








Alopocben mauribania 


Mauritian shelduck 


Mauritius 


Late 17th C (16981 _ 
Early 19th C' 1 
(1793! 


Anas marecuta 


Amsterdam Island duck 


Amsterdam 1. 






(France! 


Anas Iheodon 


Mauritian duck 


Mauritius, 


Early 18th C? 






Reunion? 


(I710I 


Camptorhyncbus 


Labrador duck 


Canada, USA 


Late 19th 0(18781 


labradohus 








Mergus australis 


Auckland island 
merganser 


New Zealand 


Early 20th C11902! 



APPENDIX 4 283 




Order FALCONIFORMES 
Family Falconidae 
Cracara lutosus 

Order GALUFORMES 
Family Phasianidae 
Argusianus bipunctatus 
Coturnix novaezelandiae 

Order GRUIFORMES 

Family Rallidae 
Aphanapteryx bonasia 
Aphanapteryx leguati 
Aramides gutturalis 
Atiantisia elpenor 
Atlantisia podarces 
Cabalus modestus 
Fulica newloni 

GaWnula nesiotis 

GaWrallus dielfenbachii 

GaWrallus pacificus 

GaWrallus sharpei 

Gallirallus wakensis 
Nesodopeus poecilopterus 
Porphyrio a/bus 

Porphyria coerutescens 
Porphyria hochsletten 

Porphyria kukwiedei 
Porzarta astnctocarpus 
Parzana monasa 

Porzana nigra 

Parzana palmen 
Porzana sandwichensis 

Order CHARADRIIFORMES 
Family Charadriidae 
Haematopus meadewaldoi 



Common name 



Guadalupe caracara 



Double-banded argus 
New Zealand quail 



Red rail 
Rodrigues rail 
Red-throated wood rail 
Ascension flightless crake 
St Helena crake 
Chatham rail 
Mascarene coot 

Tristan moorhen 

Oieffenbach's rail 

Tahiti rail 

Sharpe's rail 

Wake Island rail 
Bar-winged rail 
Lord Howe swamphen 

Reunion gallinule 
North Island takahe 

New Caledonia gallinule 
St Helena rail 
Kosrae crake 

Miller's rail 

Laysan crake 
Hawaiian crake 



Canary Islands oyslercatcher 




Place 



Guadalupe 
IMexicol 



Southeast Asia' 
New Zealand 



Early 20th C 
11900) 



Late 19th C? 
Late 19th C 118751 



Mauritius 


Early 18th C 117001 


Rodrigues 


Mid 18th C 11761) 


Peru 


19th C 


Ascension 1. 


Early 19th C? 


St Helena 


Early 16th C 


New Zealand 


Late 19th C 11900') 


Mauritius. 


Early 18th C 


Reunion 


116931 


Tristan da 


Late! 9th C 


Cunha 




Chatham Is 


Late 19th C 


iNewZealandl 


118721 


Society Is 


Early 20th C 


IFrench Polynesial 


11930sl 


Indonesia? 


Late 19th 




or early 20th C? 


Wake 1. lUSAI 


Mid 20th C 119451 


Fill 


Late 20th C 119731 


Lord Howe 1. 


Early 19th C? 


lAustralial 




Reunion 


Early 18th C 117301 


North 1., 


Late 19th C 


New Zealand 




New Caledonia 


17th C 


St Helena 


Early 16th C 


Fed, States 


Mid 19th C 


Micronesia 




Society Is 


Late 18th C 


IFrench Polynesial 




Hawaii 


Mid 20th C|)9W1 


Hawaii 


Late 19th C 118841 



Canary Is 



Mid or late 20th C 



284 WORLD ATLAS OF BIODIVERSITY 



Species I^^^^^H 


{^Common name ^^^H 




^^1 




^^H 


■^-TTr5'rs?57S»?r5«S5sa3Si 


Family Laridae 








Pinguinus impennis 


Great auk 


North Atlantic 
coasts 


Mid 19th C (18521 


Family Scolopacidae 






■ 


Prosobonia ellisl 


White-winged sandpiper 


Society Is 
(French Polynesia] 


Late 18th C 


Prosobonia leucoptera 


Tahitian sandpiper 


Society Is 
(French Polynesia! 


Late 18th C? 

,1 


Order COLUMBIFORMES 






' 


Family Columbidae 






■It 


Alectroenas nitidissima 


Mauritius blue pigeon 


Mauritius 


Early 19th C (1830s! 


Alectroenas rodericana 


Rodrigues pigeon 


Rodrigues 
(Mauritius! 


Early 18th C? 


Columba duboisi 


Reunion pigeon 


Reunion 


Early 18th C' 


Columba jouyi 


Ryukyu pigeon 


Nansei-shoto 
(Japan! 


Early 20th C (19361 

J 


Columba versicolor 


Bonin wood pigeon 


Ogasawara-shoto 
(Japan! 


Late 19th C (1889! J 


Dysmoropelia dekarchiskos 


St Helena dove 


St Helena 


Early 1 6th C? 


Ectopistes migratorius 


Passenger pigeon 


USA 


Early 20th C (19001 


Gallicolumba ferruginea 


Tanna ground dove 


Vanuatu 


Late 18th C ■S 


Gallicolumba norfolaensis 


Norfolk Island ground dove 


Norfolk 1. 
(Australia! 


Late 18th C? 1 


Microgoura meeki 


Choiseul pigeon 


Choiseul 
(Solomon Isl 


Early 20th C (190^1! 


Ptilinopus mercieni 


Red-moustached fruit-dove 


Marquesas Is 


Early 20th C J 






(French Polynesia! 


119221 ^jl 


Family Raphidae 






^ 


Pezophaps solitana 


Rodrigues solitaire 


Rodrigues 


Late 18lh C ^ 
(1760s! 


Raphus cucutlatus 


Dodo 


Mauritius 


Late 17th C (1665! 


Order PSITTACIFORMES 








Family Psittacidae 








Amazona martinicana 


Martinique parrot 


Martinique 


Late 18th C 


Amazona violacea 


Guadeloupe parrot 


Guadeloupe 


Late 18th C 
(1779! 


Ara atwoodi 


Dominican green-and-yellow 
macaw 


Dominica 


Early 19th C? 


Ara erythrocephala 


Jamaican green-and-yellow 
macaw 


Jamaica 


Early 19th C 


Ara gossei 


Jamaican red macaw 


Jamaica 


Late 18th C? 


Ara guadeloupensis 


Lesser Antillean macaw 


Guadeloupe, 
Martinique 


Late 18th C? 



APPENDIX 4 






B!? ^^^Hl 




4ra Incotor 


Cuban red macaw 


Cuba, 
Hispaniola 


Late 19th C (1885! 


Aratinga tabati 


Guadeloupe parrot 


Guadeloupe 


Late 18th C 


Conuropsis carolmensis 


Carolina parakeet 


USA 


Early 20th C (1904! 


Cyanoramphus ulietanus 


Raiatea paral^eet 


Society Is 


Late 18th C? 






(French Polynesia) 


(1773! 


Cyanoramphus zealandicus 


Black-fronted parakeet 


Society Is 

(French Polynesial 


Mid 19th C (1844! 


Lophopsittacus bensont 


Mauritius gray parrot 


Mauritius 


Late 18th C (1764! 


Lophopsittacus mauritianus 


Broad-billed parrot 


Mauritius 


Late 17th C (1675! 


Mascahnus mascannus 


Mascarene parrot 


Reunion 


Late 18th C (1770s! 


Necropsitlacus rodencanus 


Rodrigues parrot 


Rodrigues 
(Mauritius! 


Late 18th C (17611 


Neslor productus 


Norfolk Island kaka 


Phillip 1, 


Early 19th C , 






(Australia! 


Psephotus pulcherrimus 


Paradise parrot 


Australia 


Mid 20th C (19271 ' 


Psitlacula exsul 


Newton's parakeet 


Rodrigues 
(Mauritius! 


Late 19lhC (1875! 


Psittacula wardi 


Seychelles parrot 


Seychelles 


Late 19th CdeSOs! 


Order CUCUUFORMES 








Family Cuculidae 








Coua delalandei 


Snail-eating coua 


Madagascar 


Mid 19th 0(18341 


Nannococcyx psix 


St Helena cuckoo 


St Helena 


18th C? 


Order STRIGIFORMES 








Family Strigidae 








Mascarenotus gwcheti 


Reunion owl 


Reunion 


Early 17th C 


Mascarenotus munvows 


Rodrigues owl 


Rodrigues 


Early 18th C (1726! 


Mascarenotus sauzieri 


Mauritius owl 


Mauritius 


Mid 19th C (18371 


Sceloglaux albifacies 


Laughing owl 


New Zealand 


Early 20thC (1914! 


Order AP0DIF0RME5 








Family Trochilidae 








Chbrostilbon bracei 


Brace's emerald 


Bahamas 


Late 19th C (18771 


Chlorostilbon elegans 


Gould's emerald 


Bahamas? 
Jamaica? 


Late 19th C? 


Order UPUPIFORMES 








Family Upupidae 








Upupa antaois 


St Helena hoopoe 


St Helena 


Early 16th C? 


Order PASSERIFORMES 








Family Acanthisittidae 








Traversia lyalli 


Stephens Island wren 


Stephens 1 
(New Zealand! 


Late 19th 0(18941 


Xenicus bngipes 


Bush wren 


New Zealand Late 20th C (1972! 



286 WORLD ATLAS OF BIODIVERSITY 







^^^Place 


Periot^^^^^l 


'^^SiSS 




F 


. .-i-ijU 


Family Cailaeidae 




E 




Heleralocha acutirostris 


^^^^^^^M 


1 New Zealand 


Early 20th C 11907] 

i 


Family Drepanididae 




^ 


1 


Akialoa ellisiana 


Oahu 'akialoa 


Oahu. Hawaii 


Early 20th C 7 118371 


Akialoa lanaiensis 


Maui Nui 'akialoa 


Lanai. Hawaii 


Early 20th C 7118921 


Akialoa obscura 


'Akialoa 


Hawaii 


Mid 20th C 7119401 


Akiaba stejnegeri 


Kaua'i akialoa 


Kaua'i, Hawaii 


Late 20th C 7 119691 ; 


Ciridops anna 


Ula-ai-hawane 


Hawaii 


Late 19th C (18921 ; 


Drepanis funerea 


Black mamo 


Hawaii 


Early 20th C (19071 .' 


Drepanis pacifica 


Hawaii mamo 


Hawaii 


Late 19lhC 118991 ] 


Dysmorodrepanis munroi 


Lana'i hookbill 


Hawaii 


Early 20th 0(19201 ' 


Paroreomyza flammea 


Kakawihie 


Hawaii 


Mid 20th 0(19631 


PsiWrostra kona 


Kona grosbeak 


Hawaii 


Late 19th 0(18941 i 


Rhodacanlhis flaviceps 


Lesser koa-finch 


Hawaii 


Late 19th 0(18911 | 


Rhodacanthis palmen 


Greater koa-finch 


Hawaii 


Late 19th 0(18961 


Viridonla sagittirostns 


Greater 'amakihi 


Hawaii 


Early 20th 0(19011 j 


Family Fringillidae 






1 


Chaunoprodus ferreorostns 


Bonin grosbeak 


Ogasawara-shoto 
(Japan) 


Late 19th 0(189071 


Family Icteridae 








Quiscalus palustris 


Slender-billed grackle 


Mexico 


Early 20th 0(19101 


Family Meliphagidae 








Chaetoptila angustipluma 


Kioea 


Hawaii 


Late 19th 0(18591 


Moho apicalis 


Oahu 00 


Hawaii 


Mid 19th G (18371 


Moho bishopi 


Bishop's 00 


Hawaii 


Early 20th 0(19041 


Moho braccatus 


Kaua'i 00 


Kaua'i, Hawaii 


Late 20th C (19871 


Moho nobilis 


Hawaii 00 


Hawaii 


Early 20th 07(19341 


Family Muscicapidae 








Anlhomis melanocephala 


Chatham Island bellbird 


New Zealand 


Early 20th 0(19061 


Bowdlena rufescens 


Chatham Island fernbird 


Chatham Is 


Early 20th ■> 






(New Zealand! 


(19001 


Gerygone insulans 


Lord Howe gerygone 


Lord Howe 1.. 
Australia 


Early 20th 


Myadestes oabensis 


Amaui 


Hawaii 


Mid 19th C7 118251 


Myiagra freycineti 


Guam flycatcher 


Guam 


Late 20th C 119831 


Nesillas aladabrana 


Aldabra bush-warbler 


Aldabra 
ISeychellesI 


Late 20th 0(19831 


Pomarea pomarea 


Maupiti monarch 


Maupiti, 
French Polynesia 


Early 19th 07 (18231 


Turdus ravidus 


Grand Cayman thrush 


Cayman Is 


Early 20th 0(19381 


Turnagra capensis 


South Island piopio 


New Zealand 


Mid 20th 0(19631 


Turnagra lurnagra 


North Island piopio 


New Zealand 


Mid 20th 0(19551 



APPENDIX 4 287 



^^^^ffife^H^^^^H^^jyCAm m on^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^B 


Zoothera terrestns Bonin thrush 


Ogasawara-shoto 


Mid 19th C 118281 






IJapanj 




Family Sturnidae 








Apbnis corvina 


Kosrae mountain starling 


Kosrae 


Mid)VlhCll828| 






(Fed. States Micronesial 


Aplonis fusca 


Norfolk Island starling 


Norfolk 1. 
(Australia] 


Early20thC 119231 


Apbnis mavornata 


Mysterious starling 


Cook Is 


Mid 19th C 11825] 


Fregilupus vanus 


Reunion starling 


Reunion 


Mid 19th C 118505] 


Necrospar rodencanus 


Rodrigues starling 


Rodrigues 


Early 18th C 117261 


Family Zosteropidae 








Zosterops strenuus 


Robust white-eye 


Lord Howe 1. 
lAustralial 


Early 20th C (1928) 


Class REPTILIA 








Order SAURIA 






il 


Family Anguidae 








Cetestus occiduus 


Jamaican giant galliwasp 


Jamaica 


Mid 19th 0(1840] 


Family Gekkonidae 






. 


Hoplodadylus dekourti 




New Zealand I'l 


Mid 19th C 


Phelsuma gigas 


Giant day gecko 


Rodrigues 


Late 19th 


Family Iguanidae 








Leiocephalus eremitus 




Navassa 1. 
lUSAl 


Early 20th C (19001 


Leiocephalus herminieri 




Martinique 


Early 19th C (1830s] 

i 


Family Scmcidae 






] 


Leiolopisma mauntlana 




Mauritius 


Early 17th 0(1600] 


Macwscincus coctei 


Cape Verde giant skink 


Cape Verde 


Early 20th C 


Tachygia microlepis 


Tongan ground skink 


Tonga 


17th 0? 


Tetradactylus eastwoodae 


Eastwood's longtailed seps 


South Africa 


Early 20th C? ; 


Family Teiidae 






^V 


Ameiva cineracea 




Guadeloupe 


Early 20th J 


Ameiva major 


Martinique giant ameiva 


Martinique 


17th C? ^^ 


Order SERPENTES 






^H 


Family Boidae 






4H 


Bolyeria multocannata 




Round 1. 
iMauriliusI 


Late 20th 0(1975] 

1 


Family Colubridae 








Alsophis sanclicruas 


St Croix racer 


Virgin Is lUSAl 


Mid 20th J 



288 WORLD ATLAS OF BIODIVERSITY 



OT^^H 


m 


^^^ 


mmmsm 


m^^ 


Family Typhlopidae 




1 


^p 




TypMops canei 






' Mauritius 


17th C . 


Order TESTUDINES 








m 


Family Testudinidae 








^n 


Cylindraspis borbonica 






Reunion 


Early 19th C (1800! 


Cylindraspis indica 






Reunion 


Early 19th C (1800) 


Cylindraspis inepta 






Mauritius 


Early 18th C 


Cylindraspis peltasles 






Rodrigues 


Early 19th C (18001 


Cylindraspis tnserrata 






Mauritius 


Early 18th C 


Cylindraspis vosmaen 






Rodrigues 


Early 19th C (1800! 


Class AMPHIBIA 








i 


Order ANURA 








Family Discoglossidae 








< 


Discoglossus nigmenter 


Israel 


jainted frog 


Israel 


Mid 20th C (1940! 


Family Myobatrachidae 








, 


Uperoleia marmorata 


Marbled toadlet 


Australia 


Late 19th C 


Family Ranidae 








1 


Arthroleptides dutoiti 






Kenya 


Early 20th C 


Rana fisheri 


Relict 


leopard frog 


USA 


Mid 20th C (1960! 


Rana tlaloci 






Mexico 


Late 20th 0(1 990sl 


Class ACTINOPTERYGII 










Order CYPRINIFORMES 










Family Catostomidae 










' Xflas'mstes~n^u^iei 


Snake River sucker 


USA 


Early 20th Cl1927r^ 


^ Moxostoma lacerum 


Harelip sucker 


USA 


Late 19th C (1893! ] 


Family Cyprinidae 










Acanthobrama hulensis 






Lake Huleh 
(Israel) 


Late 20th C ? 


Anabanlius alburnops 






Lake Dianchi 
IChinal 


20th C 


Anabarilius polylepis 






Lake Dianchi 
(China! 


20th C 


Barbus microbarbis 






Lake Luhondo 
(Rwanda! 


Mid 20th C 


' Cephalakompsus pachychsilus 






Lake Lanao 
(Philippines! 


Mid 20th C' (post 19211' 


Chondrostoma scodrensis 






Lake Skadar 
(Albania. Yugoslavia 


20th C (post 18811 

j 


Cyprinus yilongensis 






Lake Yilong 
(China! 


Late 20th C (19771 j 





APPENDIX i 289 





P^gtnfilftBff'rianie 




Pefiofd ^ 




Evarra buslamantei 


Mexican dace 


Mexico 


Mid 20th C 11957! 


Evarra eigenmanni 


Plateau dace 


Mexico 


Mid 20th C11V54! 


Evarra tlahuacensis 


Endorheic dace 


Mexico 


Mid 20th C (19571 , 


Gita crassicauda 

1 


Thicktail chub 


USA 


Mid 20th C (19571 


Hybopsis amecse 


Ameca shiner 


Mexico 


Late 20th C 11969! 


Hybopsis aubdion 


Durango shiner 


Mexico 


Late 20th C 


Lepidomeda altivelis 


Pahranagat spinedace 


USA 


Mid 20th C 11933! 


Mandibularca resinus 


Bagangan 


Lake Lanao 
IPhilippinesI 


Early 20th C 11922! 


Notropis amecae 


Anneca shiner 


Mexico 


Late 20th C (1970s?! 


Notropis aubdion 


Durango shiner 


Mexico 


Mid 20th C 11961! : 


Notropis orca 


Phantom shiner 


Mexico. USA 


Late 20th C '> 


j Ospatutus pataemorphagus 




Lake Lanao 


Early 20th C 119241 ', 


i 




(Philippines) 


Early 20th C 11921! 


Ospatulus twnculatus 


Bitungu 


Lake Lanao 


I 




(Philippines! 


J 


Phoxinellus egndin 


Yag baligi 


Lake Egridir 
ITurkeyl 


Mid 20th C 119551 


Phoxinellus handbrschi 


Cicek 


Lake Egridir 
ITurkeyl 


Mid 20th C (1955! 


Pogonichthys ciscoides 


Clear Lake splittail 


USA 


Late 20th C 11970! | 


Puntius amarus 


Pait 


Lake Lanao 
IPhilippinesI 


Early20thC 11910! 


Punbus baoulan 


Baolan 


Lake Lanao 
(Philippines! 


Early 20th C 119261 


Punbus demensi 


Bagangan 


Lake Lanao 
(Philippines! 


Early 20th C 119211 


Punbus disa 


Disa 


Lake Lanao 
(Philippines! 


Early 20th C 119321 


Puntius flavifuscus 


Katapa-tapa 


Lake Lanao 
(Philippines! 


Early 20th C 119211 


Puntius tierrei 




Lake Lanao 
(Philippines! 


Early 20th C 119081 


Punbus tanaoensis 


Kandar 


Lake Lanao 
(Philippines! 


Early 20th C 119221 


Puntius man3lal( 


Manalak 


Lake Lanao 
IPhilippinesI 


Early 20th C 119241 


Puntius Strang 


Sirang 


Lake Lanao 
IPhilippinesI 


Early 20th C 119321 


Puntius tras 


Tras 


Lake Lanao 

IPhilippinesI 


Early 20th C 119251 


Rhinichthys deaconi 


Las Vegas dace 


USA 


Mid 20th C (19401 


Sprattlicypris palata 


Palata 


Lake Lanao 
(Philippines! 


Early 20th C (19221 


StvDodon sianifer 


Slumptooth minnow 


Mexico 


Early 20th C (19031 


^ 



290 WORLD ATLAS OF BIODIVERSITY 





■fJ'^iJyl^i.-X'V.f-, /■ ,- ^,-.:^.^■..■..:»l^^., 


^^Hj^H 


^rl^^^^H 


Order CHARACIFORMES 








Family Characidae 








Brycyon acuminatus 




Brazil 


20th C 


Order OSMERIFORMES 






^^B 


Family Retropmnidae 






^ 


Prototrocles oxyrhyncbus 


New Zealand grayling 


New Zealand 


Early 20th 0119231^ 


Order SILURIFORMES 






\ 


Family Scliilbeidae 








Plalytropius siamensis 




Thailand 


Late 20th C 119661 


Family Tnchomycteridae 






1 


Rhizosomichthys totae 




Lake Tola 
IColombial 


Mid 20th C 119571 

j 


Order SALMONIFORMES 






d 


Family Salmonidae 






^1 


Coregonus alpenae 


Longiaw cisco 


Great Lakes 
(Canada. USA] 


Late 20th C 119751 


Coregonus confusus 


Parrig 


Lake Moral 
ISwitzerlandl 


Mid 20th C ■: 
'i 


Coregonus fera 


Fera 


Lake Geneva 
ISwitzerlandl 


Mid 20th C 


Coregonus gutturosus 


Kilch 


Lake Constance 
ISwitzerlandl 


Mid 20th C 

1 


Coregonus hiemalis 


Gravenche 


Lake Geneva 
ISwitzerlandl 


Mid 20th C i 


Coregonus johannae 


Deepwater cisco 


Great Lakes 


Mid 20th C 






ICanada, USAI 




Coregonus restnctus 


Fent 


Lake Moral 
[Switzerland! 


Mid 20th C 


Salvelinus agassizi 


Silver trout 


Dublin Pond 
lUSAl 


Early 20th C 1193011 


Salvetinus mlramundis 


Orkney char 


Hoyl.lUKI 


Early 20th C 119091 


Salvelinus scharlfi 


Scharffs char 


Lough Owel 

llrelandl 


Early 20th C 119081 


Order ATHERINIFORMES 






1 


FanfiilyAtherinidae 








Chirostoma compressum 


Cuitzeo silverside 


Lake Cuitzeo 
IMexicol 


Mid 20th C 7119401 • 
! 


Rheocles sikorae 


Zona 


Madagascar 


Mid 20th C 119661 


Order CYPRIN0D0NTIFORME5 








Family Cyprinodontidae 










"T3SR5FFitb de la Presa 


Mexico 


Late 20lh CI19881 ■ 
Late,25tJ)C119£l4)J 


, Cyprinodon mmemomm 


Cachorrito de la Trinidad 


. Mexico 



APPENDIX k 291 



^^^^^^^^^^^Hr Common name ^^^^^^^^^^^^^^^^^^^^^^| 


wr_ 


'''^^prinodon iatifasciatus 


Pernio de Parras 


Mexico 


Early 20th C 119031 


Cypnnodon sp. ' 


Monkey Spring puplish 


USA 


Late 20th C 


Leptolebias marmoralus 
Orestias cuvien 


Ginger pearllish 
Lake Tilicaca oreslias 


Brazil 

Lake Tilicaca 
IBolivia. Peru] 


Mid 20th C [mil] 
Mid 20th C 119371 


Family Fundulidae 


Fundulus albolineatus 


Whileline lopminnow 


USA 


Early 20th C ■> 1)899] 


Family Goodeidae 


Characodon garmani 
Empetrichthys merriami 


Parras characodon 
Ash Meadows kiUilish 


Mexico 
USA 


Late 19th C (18891 1 
Mid 20th CI19A8I j 


Family Poeciliidae 


Gambusia amistadensis 


Amislad gambusia 


USA 


Mid 20th C 119631 


Order BELONIFORMES 
Family Adrianichthyidae 


Adriankhthys kruyti 


Duck-billed buntingi 


Lake Poso 
llndonesial 


Late 20lh C 119831 i 


Order GASTEROSTEIFORMES 
Family Gasterosteidae 


' Pungitius kaibarae 


Kyoto ninespine stickleback 


Japan 


Mid 20th C 119591 j 



Order 5C0RPAENIF0RMES 








Family Cottidae 








Cottus echinatus 


Utah Lakesculpin 


Utah Lake 
lUSAI 


Early 20lh 0(19281 


Order PERCIFORMES 






m 


Family Cichlidae 






m 


Tristramella magdalenae 




Israel 


Late 20th 0(19971 


Family Gobiidae 












■ Lake Poso. 
1^ Sulawesi (Indonesia 


Late 20th 0(19851 


Hapbchromis attigenis 




Lake Victoria 


Mid/late 20th C ? 


Hapbchromis apogonoides 




Lake Victoria 


Mid/late 20th C ? 


Hapbchromis arcanus 




Lake Victoria 


Mid/late 20th C ? 


Hapbchromis argenteus 




Lake Victoria 


Mid/late 20th C ? 


Hapbchromis artaxerxes 




Lake Victoria 


Mid/late 20th C ' 


Hapbchromis barbarae 




Lake Victoria 


Mid/late 20th C ^ 


Hapbchromis bareli 




Lake Victoria 


Mid/late 20th ■> 


Hapbchromis barbni 




Lake Victoria 


Mid/late 20th ? 


Hapbchromis bayoni 




Lake Victoria 


Mid/late 20th ? 


Hapbchromis boops 




Lake Victoria 


Mid/late 20th ? 



J 



292 WORLD ATLAS OF BIODIVERSITY 



^^^^m^^^ 


^^^^^^ Place 


^^^H 


Haplochromis cassius 


Lake Victoria 


Mid/late 20th C '' 


Haptochromis cinctus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis cnester 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis decticostoma 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis dentex group 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis diplotaenia 


Lake Victoria 


Mid/late 20th C "> 


Haplochromis estor 


Lake Victoria 


Mid/late 20th C 


Haplochromis llavipinnis 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis gilberti 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis gowersi 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis guiarti 


Lake Victoria 


Mid/late 20th C ■? 


Haplochromis heusinkveldi 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis hiatus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis ins 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis longirostris 


Lake Victoria 


Mid/late 20th C '' 


Haplochromis macrognathus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis maculipinna 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis mandibularis 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis martini 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis megalops 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis michaeli 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis microdon 


Lake Victoria 


Mid/late 20th C '' 


Haplochromis mylergates 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis nanoserranus 


Lake Victoria 


Mid/tale 20th C ? 


Haplochromis nigrescens 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis nyanzae 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis obtusidens 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis pachycephalus 


Lake Victoria 


Mid/late 20th C '' 


Haplochromis paraguiarti 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis paraplagiostoma 


Lake Victoria 


Mid/late 20th C? 


Haplochromis parorthostoma 


Lake Victoria 


Ivlid/late 20th C '' 


Haplochromis percoides 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis pharyngomylus 


Lake Victoria 


Mid/late 20th C ? 


Haptochromis prognathus 


Lake Victoria 


Mid/late 20th C •> 


Haplochromis pseudopellegrini 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis pyrrhopteryx 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis spekii 


Lake Victoria 


Mid/late 20th C '' 


Haplochromis teegelaari 


Lake Victoria 


Mid/late 20th C '^ 


Haplochromis thuragnathus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis tridens 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis victorianus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis xenostoma 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'bartoni-like' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'bicolor' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis big teeth' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'black cryptodon' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'back pectoral' 


Lake Victoria 


Mid/late 20th C ? 



APPENDIX 4 293 



HHHI^^H^Hi^l^HH 


Haplochromis 'chlorocephalus 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'citrus' 


Lake Victoria 


Mid/late 20th C ■> 


Haplochromis 'coop' 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'elongate rockpicker' 


Lake Victoria 


Mid/late 20th C '> 


Haplochromis 'fiiamentus 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'fleshy lips' 


Lake Victoria 


Mid/lale 20th C ' 


Haplochromis 'gray pseudo-nigricans' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'large eye guiarti' 


Lake Victoria 


Mid/late 20th C '> 


Haplochromis 'lividus-frels' 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'longurius' 


Lake Victoria 


Mid/late 20th C '' 


Haplochromis 'macrops like' 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'micro-obesus' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'morsei' 


Lake Victoria 


Mid/late 20th C -> 


Haplochromis 'orange cinereus' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'orange macula' 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'orange yellow big teeth' 


Lake Victoria 


Mid/late 20th C ■> 


Haplochromis 'orange yellow small teeth' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'paropius-like' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'pink paedophage' 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'pseudo-morsei' - 


Lake Victoria 


Mid/late 20th C ' 


Haplochromis 'purple head' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'purple miller' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 11] 'purple rocker' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'red empodisma' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'red eye scraper' 


Lake Victoria 


Mid/late 20th C ? 


Haptochromis 'reginus' 


Lake Victoria 


Mid/late 20th C ^ 


Haplochromis 'regius 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'short supramacrops' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'small blue zebra' 


Lake Victoria 


Mid/lale 20th C ? 


Haplochromis 'small empodisma' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'smoke' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis "soft gray' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'stripmac' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'supramacrops' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'theliodon-like' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'tigrus' 


Lake Victoria 


Mid/late 20th C ■? 


Haplochromis 'too small' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'twenty' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'two stripe white lip' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'wyber' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'xenognathus-like' 


Lake Victoria 


Mid/late 20th C ? 


Haplochromis 'yellow' 


Lake Victoria 


Mid/late 20th C i 


Haplochromis 'yellow-blue' 


Lake Victoria 


Mid/late 20th C ? 


Hoplotilapia retrodens 


Lake Victoria 


Mid/late 20th C ? 


Psammochromis cryplogramma group 


Lake Victoria 


Mid/late 20th C ? 



294 WORLD ATLAS OF BIODIVERSITY 



REFERENCES 

1 BirdLlfe International 2000. Threatened birds of the world. Lynx Edicions and BirdLife 
International, Barcelona and Cambridge. 

2 CREO. Connmittee on Recently Extinct Organisms ICREO] website: http://creo.amnh.org/ 
(accessed January 20021. 

3 Hilton-Taylor. C. (compiler! 2000. 2000 lUCN Red List of threatened species. lUCN, Gland and 
Cambridge. Online at http://www.redlist.org/ (accessed April 2002). 



APPENDIX 5 295 



APPENDIX 5: 

BIODIVERSITY AT COUNTRY LEVEL 

This table includes estimates of the number of 
mammals, breeding birds and vascular plants in 
each country of the world, together with estimates 
of the number of these endemic to each country. 
The threat status of mammals and birds has been 
comprehensively assessed, and for these classes 
the number and percentage of globally threatened 
species present in each country are given. In the 
mammals, most estimates of the total number 
present relate to native non-marine species only, 
but the threatened species counts for many 
countries include marine species (a few island 
countries have a small number of non-marine 
mammals but a higher number of threatened 
species - the percentage figure is omitted in such 
casesl. The richness and endemism data should 



Methodology notes 

Numerical indices representing national biodiversity 
have been derived from available estimates of species 
total and species endemism. Taxonomic groups covered 
are: mammals, birds, reptiles, amphibians and vascular 
plants Iferns to flowering plants!. For birds alone, the 
database includes estimates of botfi the total number 
recorded li.e. including non-breeding migrants and 
accidental visitors! and the number of breeding bird 
species; the latter has been used in the analysis. All 
countries belovi/ 5 000 km^ in area have been excluded 
from the analysis, leaving 169 countries. 

Four arbitrary assumptions are made: the four 
vertebrate classes included are of equal importance; 
plants are of equal importance to the vertebrates 
combined; richness and endemism are reasonably 
vtfell correlated at country level across the taxonomic 
groups covered; vertebrates plus plants provide 
a valid surrogate for biodiversity in general. 

Because interest lies in relative biodiversity rather 
than absolute values, data in each column are first 
normalized. Each estimate N/ for each country /is 
divided by Nmax where Nmax is the highest value for 



be taken as provisional; they are subject to 
change with new taxonomic treatments and 
new surveys. The columns headed Dl and Al, 
respectively, contain the unweighted national 
diversity indices, and the diversity indices adjusted 
for area [see methodology notes below for 
details!. The index is based on data for the groups 
shown here, plus reptiles and amphibians, not 
included here. The Al column is the basis for 
Map 5.4. 



Source: WCMC database; data derived from a large number of 
published and unpublished sources, including country reports and 
regional checklists Numbers of threatened species retrieved 
from the online version of the 2000 tUCN Red List of threatened 
species, at http;//www.redlist.org/ [accessed f^larch 20021. 



that parameter This transforms the data to within the 
range 0-1, with the most important country having the 
value Nmax/Nmax- 1, and the least important having 
the value closest to zero. Estimates for mean vertebrate 
richness fV/?i and mean vertebrate endemism IVEIare 
derived by averaging the figures for all classes, and 
estimates for combined richness fWand combined 
endemism /flare derived for each country by 
averaging figures for vertebrates and plants. Inspection 
of the data shows that PR and W correlate quite 
closely, while Pf tends to be approximately half VE; 
where estimates of PPand Pf are missing the 
appropriate vertebrate-based value has been inserted 
before calculating Rand £ 

An overall diversity index IDII is calculated for each 
country as the mean of R and £ This treats richness 
and endemism equally and so makes fewest 
assumptions about their relative significance in terms of 
overall biodiversity, but the 0/ could be weighted to give 
greater importance to either. Because species richness 
tends to increase with area, and with proximity to the 
humid tropics, 0/ is strongly affected by country area 



296 WORLD ATLAS OF BIODIVERSITY 



and geographical position, but it also takes account of 
levels of endemism, which are shaped by several 
factors, including topography, geographic isolation 
and tectonic history. 

Because area is an important determinant of 
species number, there is much interest in evaluating 



relative levels of biodiversity per unit area, i.e. how 
much more or less rich in species is any given area 
lor country). This may be addressed by the Arrhenius 
equation describing the species-area relationship 
Hog S = g + z log A], where S = number of species, 
A = area, z is the slope of the line, and g another 





^flW^ 


^^^5!^^ 


^^!P^ 


^mSnfWS^ 




. 


km' 






total 


endemic 


no. theatened 


Afghanistan 


652 225 


0.063 


-0.296 


119 


2 


13 


Albania 


28 750 


0.035 


-0.019 


68 





3 


Algeria 


2 381 7^5 


0.045 


- 1.003 


92 


2 


13 


American Samoa 


197 






3 





3 


Andorra 


465 






44 





1 


Angola 


1 246 700 


0.176 


0.544 


276 


7 


18 


Anguilla 


91 






3 








Antigua and Barbuda 


M2 






7 








Argentina 


2 777 815 


0.196 


0.423 


320 


49 


32 


Armenia 


29 800 


0.042 


0.153 


84 


3 


7 


Aruba 


193 






- 





1 


Australia 


7 682 300 


0.608 


1.268 


252 


206 


63 


Austria 


83 855 


D.036 


-0.293 


83 





9 


Azerbaiian 


86 600 


0.05 


0.027 


99 





13 


Bahamas 


13 865 


0.017 


- 0.503 


12 


3 


5 ■ 


Bahrain 


661 






17 





1 


Bangladesh 


laooo 


0.059 


0.058 


125 





21 


Barbados 


430 






6 





i 


Belarus 


207 600 


0.029 


-0.771 


74 





5 


Belgium 


30 520 


0.023 


- 0.441 


58 





11 : 


Belize 


22 965 


0.056 


0.526 


125 





4 


Benin 


112 620 


0.08 


0.437 


188 





7 


Bermuda 


54 






3 





2 


Bhutan 


46 620 


0.058 


0.366 


160 





20 


Bolivia 


1 098 575 


0.239 


0.882 


316 


16 


23 1 


Bosnia and Herzegovina 


51 129 


0.034 


-0.200 


72 





10 1 


Botswana 


575 000 


0.062 


-0.287 


164 





' 1 


Brazil 


8 511 965 


0.74 


1.436 


394 


119 


79 1 


British Ind. Oc. Terr. 








- 





1 


Brunei ^^|||||||^ 


5 765 


0.071 


1.145 


157 





' 1 


Bulgaria ^^^^^ 


110910 


0.044 


-0.167 


81 





15 1 


Burkina Faso 


274 122 


0.068 


0.011 


147 





' i 


Burundi 


27 835 


0.072 


0.723 


107 





5 


Cambodia 


181 000 


0.059 


0.001 


123 





21 


Cameroon 


475 500 


0.167 


0.762 


409 


14 


37 


Canada 


9 922 385 


0.067 


- 1.014 


193 


7 


14 


Cape Verde 


4 035 






5 





3 


Cayman Islands 


259 






8 








Central African Republic 


624 975 


0.08 


- 0.058 


209 


2 


12 

















APPENDIX 5 297 



constant. If it is assumed, given its area dependence, 
that Dl IS scaled in the same way as 5, a regression 
analysis of log Dl and log A allows the constants g and z 
to be calculated. The regression line establishes the 
expected biodiversity value of each country for its area, 
and the distance of each country point from the line 



gives a measure M//of how much more Itvel or less 
l-vel diverse is a given country than expected. Al 
attempts to assess diversity per unit area, rather than 
overall biodiversity value per country, and it is noticeable 
that several smaller mainland countries and island 
states move up in rank order, as would be expected. 



Hwa^R^^ 




^^ira^^ 






Hlant^^ 


^^linl^^ 


% threatened 


breeding total 


endemic 


no. threatened 


% threatened 


total 


endemic , 


11 


235 





11 


5 


4 000 


800 


A 


230 





3 


1 


3 031 


24 


U 


192 


1 


6 


3 


3 164 


250 


100 


34 





2 



6 


471 


15 


2 


113 








1 350 


- 


7 


765 


12 


15 


2 


5 185 


1 260 





- 










321 


1 





49 





1 


2 


1 158 


22 


10 


897 
242 


19 



38 
4 


4 
2 


9 372 


1 100 


8 


3 553 


- 




48 











460 


25 


25 


649 


350 


32 


5 


15 638 


14 074 


11 


213 





3 


1 


3 100 


35 


13 


248 





8 


3 


4 300 


240 
118 


42 


88 


3 


4 


5 


1 111 


6 


28 





6 


21 


195 


- 


17 


295 





23 


8 


5 000 


- 





24 





1 


4 


572 


3 


7 


221 






3 


1 


2 100 


_ 


19 


180 


2 


1 


1 550 


1 


3 


356 





2 


1 


2 894 


150 


i 


307 





2 


1 


2 500 


- 


67 


8 


1 


2 


25 


167 


15 


13 


448 





12 


3 


5 468 


75 


7 


18 


27 




17 367 


4 000 


U 


218 





3 


1 


- 


- 


3 


386 


1 


7 


2 


2 151 


17 


20 


1 492 


185 


113 


8 


56 215 


- 




14 











101 


- 


6 


359 





15 


4 


6 000 


7 


19 


240 





10 


4 


3 572 


320 


5 


335 





2 


1 


1 100 


- 


5 


451 





7 


2 


2 500 


- 


17 


307 



8 


19 
15 


6 


- 


- 


9 


690 


2 


8 260 


156 


7 


426 


5 


8 


2 


3 270 


147 


60 


38 


4 


2 


5 


774 


86 





45 





1 


2 


539 


19 


6 


537 


1 


3 


1 


3 602 


100 

















= zero: - = no data. 



298 WORLD ATLAS OF BIODIVERSITY 





Area 


°'jl 


^^^^ 


J^mmals 


Mammals 


Mammals 




; km' 


1 


im^i 


^nal 


endemic 


no. theatened 


Chad 


1 2U 000 


0.049 


- 0.739 


134 


1 


17 


Chile 


751 625 


0.112 


0.229 


91 


16 


21 


China 


9 597 000 


0.392 


0.767 


394 


83 


76 


Colombia 


1 138 915 


0.538 


1.685 


359 


34 


36 


Comoros 


1 860 






12 


2 


2 


Congo, Dem. Rep. 


2 345i10 


0.218 


0.579 


450 


28 


40 


Congo. Republic 


342 000 


0.128 


0.589 


200 


2 


12 


Cook Islands 


233 






1 





1 


Costa Rica 


50 900 


0.162 


1.358 


205 


7 


14 


Cote d'lvoire 


322 i65 


0.116 


0.507 


230 





17 


Croatia 


56 538 


0.036 


-0.169 


76 





9 


Cuba 


1U525 


0.12 


0.829 


31 


12 


n J 


Cyprus 


9 250 


0.017 


- 0.429 


21 


1 


3 1 


Czech Republic 


78 864 


D.033 


-0.356 


81 





8 1 


Denmark 


43 075 


0.021 


- 0.643 


43 





5 1 


Djibouti 


23 000 


0.02 


-0.528 


61 





^ J 


Dominica 


751 






12 





' 1 


Dominican Republic 


48 440 


0.076 


0.625 


20 





' 1 


Ecuador 


461 475 


0.353 


1.519 


302 


25 


31 1 


Egypt 


1 000 250 


0.038 


- 0.936 


98 


7 


12 1 


El Salvador 


21 395 


0.048 


0.393 


135 





2 


Equatorial Guinea 


28 050 


0.084 


0.869 


184 


1 


15 


Eritrea 


117 600 


0.057 


0.088 


112 





12 


Estonia 


45 100 


0.025 


- 0.483 


65 





5 


Ethiopia 


1 104 300 


0.145 


0.383 


277 


31 


3^ -- 


Falkland Islands 


15 931 


0.004 


- 2.040 








4 jl 


Faroe Islands 








- 





3 jl 


Fiji 


18 330 


0.028 


-0.100 


4 


1 


5 S 


Finland 


337 030 


0.023 


- 1.145 


60 





6 fl 


France 


543 965 


0.051 


- 0.473 


93 





18 H 


French Guiana 


91 000 


0.079 


0.483 


150 


3 


9 M 


French Polynesia 


3 940 












3 H 


French S. and Antarctic Te 


rr 7 241 


0.001 


-3.261 


- 





;fl 


Gabon 


267 665 


0.116 


0.56 


190 


3 


15 fl 


Gambia 


10 690 


0.036 


0.308 


117 





3 S 


Georgia 


69 700 


0.051 


0.111 


107 


2 


U H 


Germany 


356 840 


0.033 


-0.770 


76 





12 S 


Ghana 


238 305 


0.114 


0.571 


222 


1 


13 9 


Gibraltar 


7 






7 





S 


Greece 


131 985 


0.062 


0.129 


95 


3 


14 fl 


Greenland 


2 175 600 


0.007 


-2.821 


9 





7 ^1 


Grenada 


345 






15 





S 


Guadeloupe 


1 780 






11 


4 


5 S 


Guam 


450 






2 





2 fl 


Guatemala 


108 890 


0.142 


1.014 


250 


3 


6 S 


Guinea 


245 855 


0.094 


0.373 


190 


1 


11 fl 


Guinea-Bissau 


36 125 


0.05 


0.289 


108 





2 B 



APPENDIX 5 299 



Birds Birds Birds Birds Plants Plants 

breeding total endemic no. threatened % threatened total endemic 



J 



= zero; - = no data. 



13 


370 





5 


1 


1 600 


_ 


23 


296 


16 


15 


5 


5 284 


2 698 


19 


1 100 


70 


73 


7 


32 200 


18 000 


10 


1 695 


67 


77 


5 


51 220 


15 000 


17 


50 


14 


9 


18 


721 


136 


9 


929 


24 


28 


3 


11 007 


1 100 


6 


449 





3 


1 


6 000 


1 200 


100 


27 


6 


7 


26 


284 


3 


7 


600 


6 


13 


2 


12 119 


950 


7 


535 


2 


12 


2 


3 660 


62 


12 


224 


Q 


4 


2 


4 288 


35 


137 


21 


18 


13 


6 522 


3 229 


U 


79 


2 


3 


4 


1 682 


- 


10 


199 





2 


1 


1 900 


- 


12 


196 



1 


1 
5 


1 

4 


1450 
826 


1 
6 


7 


126 


8 


52 


2 


3 


6 


1 228 


11 


25 


136 





15 


11 


5 657 


1 800 


10 


1 388 


37 


60 


4 


19 362 


4 000 


12 


153 





7 


5 


2 076 


70 


1 


251 











2911 


17 


8 


273 


3 


5 


2 


3 250 


66 


11 


319 





7 


2 


- 


- 


8 


213 





3 


1 


1 630 


- 


12 


626 
64 


28 


16 


3 


6 603 


1 000 




4 


3 


5 


165 


14 




71 











236 


1 




74 


24 


12 


16 


1 518 


760 


10 


248 





3 


1 


1 102 


- 


19 


269 


1 
1 


5 




2 


4 630 


133 


6 


- 


5 625 


144 




60 


25 


22 


37 


959 


560 




48 


3 


3 


6 


- 


- 


8 


466 


1 


5 


1 


6 651 


- 


3 


280 





2 


1 


974 


- 


13 





3 




4 350 


380 


16 


239 





5 


2 


2 682 


6 


6 


529 





8 


2 


3 725 


43 





34 





1 


3 


600 


- 


15 


251 
62 






7 


3 


4 992 


742 


78 








529 


15 





50 


1 


1 


2 


1 068 


4 


ib 


52 


2 


1 


2 


1 400 


26 


100 


18 


2 


2 


11 


330 


69 


2 


458 
409 


1 


6 


1 


8 681 


1 171 


6 





10 


2 


3 000 


88 


2 


243 











1 000 


12 



300 WORLD ATLAS OF BIODIVERSITY 



■Countri^HH^^^I 




■ 




■Mammals 

' total 


Mammals 
endemic 


Mammals 
no. theatened 


Guyana 


2U970 


0.133 


0.758 


193 


1 


9 


Haiti 


27 750 


0.071 


0.71 


20 





4 


Honduras 


112 085 


0.094 


0.597 


173 


2 


9 


Hungary 


93 030 


0.031 


- 0.457 


83 





9 


Iceland 


102 820 


0.006 


- 2.080 


11 



44 


6 
86 


India 


3 166 830 


0.326 


0.896 


390 


Indonesia 


1 919^5 


0.731 


1.844 


515 


222 


140 


Iran 


1 648 000 


0.091 


-0.194 


140 


6 


23 


Iraq ^^^^^^^^ 


438 445 


0.041 


-0.629 


81 


2 


10 


^^l^^^^^l 


68 895 


0.013 


- 1.248 


25 





5 
14 


^^^|H^H 


• 20 770 


0.043 


0.285 


116 


4 


Italy ^^^^^B 


301 245 


0.065 


-0.056 


90 


3 


14 


Jamaica 


11 425 


0.051 


0.619 


24 


2 


5 


Japan 


369 700 


0.124 


0.536 


188 


42 


37 


Jordan 


96 000 


0.036 


-0.310 


71 





8 


Kazakhstan 


2 717 300 


0.071 


-0.581 


178 


4 


18 


Kenya 


582 645 


0.145 


0.56 


359 


23 


51 


Kiribati 


684 






- 








Korea, DPR 


122 310 


0.025 


-0.775 


- 





13 . 


Korea, Republic 


98 445 


0.03 


-0.518 


49 





13 1 


Kuwait 


24 280 


0.007 


- 1.564 


21 





1 


Kyrgyzstan 


198 500 


0.036 


-0.537 


83 


1 


7 


Lao PDR 


236 725 


0.081 


0.229 


172 





27 


Latvia 


63 700 


0.025 


- 0.553 


83 





5 


Lebanon 


10 400 


0.031 


0.145 


57 





6 


Lesotho 


30 345 


0.025 


- 0.354 


33 





3 


Liberia 


111 370 


0.059 


0.132 


193 





16 


Libya 


1 759 540 


0.029 


-1.343 


76 


5 


9 ■ 


Liechtenstein 


160 






64 





3 ; 


Lithuania 


65 200 


0.026 


- 0.544 


68 





5 
6 


Luxembourg 


2 585 






55 





Macedonia, FYR 


25 713 


0.037 


0.077 


78 





11 1 


Madagascar 


594 180 


0.298 


1.277 


141 


93 


50 1 


Malawi 


94 080 


0.079 


0,473 


195 





' 1 


Malaysia 


332 965 


0.254 


1.28 


300 


36 


47 1 
fl 


Maldives 


298 






3 





Mall 


1 240 140 


0.053 


- 0.658 


137 





13 M 


Malta 


316 






22 





m 


Marshall Islands 


181 












I 


Martinique 


1 079 






9 





■■ 


Mauritania 


1 030 700 


0.041 


-0.856 


61 


1 


10 ■ 


Mauritius 


1 865 






4 


1 


3 M 


Mayotte 


376 






- 





S 


Mexico 


1 972 545 


0.589 


1.621 


491 


140 


69 ^ 


Micronesia, Fed. States 


702 






6 


3 


6 ■ 


Moldova 


33 700 


0.025 


- 0.396 


68 





3 j 


Monaco 


2 






- 





9 



APPENDIX 5 301 



Birds 
breeding total 



Birds 
endemic 



Birds 
no. threatened 



Birds 
% threatened 




5 


678 





2 





6 409 






75 


1 


14 


19 


5 242 


1 623 


5 


422 


1 


5 


1 


5 680 


148 


11 


205 





8 


4 


2214 


38 


55 


88 
923 



58 



70 





377 


1 


22 


8 


18 664 


5 000 


27 


1 519 


408 


113 


7 


29 375 


17 500 


16 


323 


1 


13 


4 


8 000 


- 


12 


172 


1 


11 


6 


- 


- 


20 


U2 





1 
12 


1 
7 


950 


- 


12 


180 





2317 


- 


16 


234 





5 


2 


5 599 


712 


21 


113 


26 


12 


11 


3 308 


923 


20 


250 


21 


32 


13 


5 565 


2 000 


11 


U1 
396 





8 


6 
4 


2 100 
6 000 


- 


10 





15 


U 


844 


9 


24 


3 


6 506 


265 




26 


1 


3 


12 


60 


2 




115 


1 


19 


17 


2 898 


107 


27 


112 





25 


22 

35 


2 898 


224 


5 


20 





7 


234 


- 


8 


- 





4 




4 500 


- 


16 


487 


1 


19 


4 


8 286 


- 


6 


217 





3 


1 


1 153 


- 


11 


154 





7 


5 

12 


3 000 
1 591 


2 


9 


58 





7 


8 


372 


1 


11 


3 


2 200 


103 


12 


91 





1 


1 


1 825 


134 


5 


124 





1 


1 


1 410 


- 


7 


202 





4 


2 


1 796 


- 


11 


126 





1 


1 


1 246 


U 


210 





3 


1 


3 500 


- 


35 


202 


105 


27 


13 


9 505 


6 500 


4 


521 





11 


2 


3 765 


49 


16 


501 
23 


18 



37 
1 


7 
4 


15 500 


3 600 





583 


- 


9 


397 





4 


1 


1 741 


11 


U 


26 





1 


4 


914 


5 




17 





1 


6 


100 


5 





52 


1 


2 


4 


1 287 


30 


16 


273 





2 


1 


1 100 


75 


27 


8 


9 


33 


750 


325 




27 


2 


3 


11 


500 


- 


U 


769 


92 


38 


5 


26 071 


12 500 


100 


40 


18 


5 


13 


1 194 


293 


U 


177 





5 


3 


1 752 


- 




- 










- 


- 



= zero; - = no data. 



302 WORLD ATLAS OF BIODIVERSITY 



Countr^^^^^^^H 


Arefll 


^^■j 


1 " 


Mammals 


Mammals 


Mammals 




km'^" 




E 


total 


endemic 


no. theatened 


Mongolia 


1 565 000 


0.051 


-0.767 


133 





12 


Montserrat 


104 






7 





1 


Morocco 


458 730 


0.057 


- 0.304 


105 


4 


16 


Mozambique 


lU 755 


0.09 


0.005 


179 


2 


15 


Myanmar 


678 030 


0.141 


0.493 


300 


6 


36 


Namibia 


824 295 


0.102 


0.116 


250 


3 


14 


Nauru 








- 








Nepal 


U1 415 


0.096 


0.549 


181 


2 


27 


Netherlands 


41 160 


0.022 


-0.599 


55 





11 


Netherlands Antilles 


800 






- 





3 


New Caledonia 


19 105 


0.078 


0.904 


11 


3 


6 


New Zealand 


265 150 


0.065 


-0.017 


2 


2 


8 


Nicaragua 


148 000 


0.098 


0.555 


200 


2 


6 


Niger 


1 186 410 


0.061 


-0.512 


131 





11 


Nigeria 


923 850 


0.107 


0.131 


274 


4 


25 


Niue 


259 






1 








Northern Marianas 


477 






- 





2 


Norway 


386 325 


0.024 


- 1.107 


54 





10 


Oman 


271 950 


0.03 


-0.812 


56 


2 


9 


Pakistan 


803 940 


0.08 


-0.121 


188 


4 


18 


Patau 


492 






2 





3 


Panama 


78 515 


0.162 


1.236 


218 


16 


20 


Papua New Guinea 


462 840 


0.271 


1,254 


214 


65 


58 J 


Paraguay 


406 750 


0.115 


0.429 


305 


2 


' 1 


Peru 


1 285 215 


0.396 


1.344 


460 


49 


47 1 


Philippines 


300 000 


0.225 


1.188 


153 


102 


50 1 


Pitcairn Islands 














1 


Poland 


312 685 


0.032 


-0.761 


84 





15 


Portugal 


92 390 


0.045 


- 0.088 


63 


1 


17 
2 1 


Puerto Rico 


8 960 


0.033 


0.259 


16 





Qatar 


11 435 


0.005 


- 1.770 


11 





,■■ 


Reunion 


2510 






2 





3 H 


Romania 


237 500 


0.039 


- 0.490 


84 





17 .^1 


Russia 


17 075 400 


0.179 


-0.179 


269 


22 


42 "9 


Rwanda 


26 328 


0.087 


0.925 


151 





B m 


San Marino 








13 





1 ^ 


Sao Tome and Principe 


964 






8 


4 


3 '9 


Saudi Arabia 


2 400 900 


0.04 


- 1.129 


77 





7 S 


Senegal 


196 720 


0.057 


-0.065 


192 





11 ^1 


Seychelles 


404 






6 


2 


4 ^1 


Sierra Leone 


72 325 


0.083 


0.588 


147 





11 ^1 


Singapore 


616 






85 


1 


3 H 


Slovakia 


14 035 


0.037 


0.252 


85 





9 S 


Slovenia 


20 251 


0.036 


0.106 


75 





9 ^1 


Solomon Islands 


29 790 


0.049 


0.316 


53 


21 


21 |H 


Somalia 


630 000 


0.087 


0.025 


171 


12 


19 ^H 


South Africa 


1 184 825 


0.252 


0.915 


247 


35 


41 H 



APPENDIX 5 303 



^^^nmals 


Birds 


Bird^^l 


^^^M 


Birds 


Plants 


Plants 


^threatened 


breeding total 


endemic iliSiW^Shed 


% threatened 


total 


endemic^H 


9 


426 





16 


4 


2 823 


229 


U 


37 


1 


2 


5 


671 


2 


15 


210 





9 


4 


3 675 


625 


8 


498 





16 


3 


5 692 


219 


12 


867 


4 


35 
9 


4 


7 000 


1 071 


6 


469 


3 


2 


3 174 


687 




9 


1 


2 


22 


50 


1 


15 


611 


2 


26 


4 


6 973 


315 


20 


191 





4 


2 


1 221 


- 




77 





1 


1 


- 


- 


55 


107 


22 


9 


8 


3 250 


3 200 




150 


74 


49 


33 


2 382 


1 942 


3 


482 





5 


1 


7 590 


40 


8 


299 





3 


1 


1 460 


- 


9 


681 
15 


2 



9 
1 


1 


4715 


205 





7 


178 


1 




28 


2 


8 


29 


315 


81 


19 


243 





2 


1 


1 715 


1 


16 


107 





10 


9 


1 204 


73 


10 


375 





17 


5 


4 950 


372 




45 


10 


2 


4 


- 


- 


9 


732 


9 


16 


2 


9915 


1 222 


27 


644 


94 


32 


5 


11 544 


- 


3 


556 





26 


5 


7 851 


- 


10 


1 538 


112 
186 


71 

67 


5 


17 144 


5 356 


33 


196 


34 


8 931 


3 500 




19 


5 


8 


42 


76 


14 


18 


227 





4 


2 


2 450 


3 


27 


207 


2 


7 


3 


5 050 


150 


13 


105 


12 


8 
6 


8 


2 493 


235 





23 





26 


355 


- 




18 


4 


5 


28 


546 


165 


20 


247 





8 


3 


3 400 


41 


16 


628 


13 


38 


6 


11 400 


- 


5 


513 





9 


2 


2 288 


26 


8 


- 










- 


- 


38 


63 


25 


9 


14 


895 


134 


9 


155 





15 


10 


2 028 




6 


384 





4 


1 


2 086 


26 


67 


38 


11 


10 


26 


250 


182 


7 


466 


1 


10 


2 


2 090 


74 


/I 


118 





7 


6 


2 282 


2 


11 


209 





4 


2 


3 124 


92 


12 


207 





1 





3 200 


22 


iO 


163 
422 


43 
11 


23 


14 


3 172 


30 


11 


10 


2 


3 028 


500 


17 


596 


8 


20 


3 


23 420 


- 



= zero; - = no data. 



304 WORLD ATLAS OF BIODIVERSITY 



Country 


Area 


Dl 


Al 


Mammals 


Mammals 


Mammals 


^ 


km' 






total 


endemic 


no. theatened 


Spain 


504 880 


0.067 


-0.172 


82 


4 


24 


Sri Lanka 


65 610 


0.082 


0.606 


88 


15 


20 


St Helena and dep. 


411 






2 





1 


St Kittsand Nevis 


261 






7 








St Lucia 


619 






9 





1 


St Vincent 


389 






8 


1 


2 


Sudan 


2 505 815 


0.137 


0.093 


267 


11 


24 


Suriname 


163 820 


0.092 


0.471 


180 


2 


11 


Swaziland 


17 365 


0.044 


0.353 


47 





4 


Sweden 


440 940 


0.026 


-1.067 


60 





8 


Switzerland 


41 285 


0.033 


-0.173 


75 





6 


Syria 


185 680 


0.046 


- 0.265 


63 


2 


4 


Taiwan 


36 960 


0.058 


0.418 


63 


11 


13 


Taj il<i Stan 


143 100 


0.033 


- 0.536 


84 


1 


9 


Tanzania 


939 760 


0.189 


0.693 


316 


15 


43 


Thailand 


514 000 


0.162 


0.709 


265 


7 


34 


Togo 


56 785 


0.094 


0.781 


196 





9 


Tokelau 

















Tonga 


699 






2 





1 


Trinidad and Tobago 


5 130 


0.045 


0.729 


100 


1 


1 


Tunisia 


164 150 


0.033 


- 0.572 


78 


1 


11 


Turkey 


779 450 


0.114 


0.237 


116 


2 


17 


Turkmenistan 


488 100 


0.044 


-0.572 


103 





13 


Turks and Caicos Islands 


430 






- 








Tuvalu 


25 






- 








Uganda 


236 580 


0.12 


0.624 


345 


6 


19 


Ukraine 


603 700 


0.05 


- 0.509 


108 


1 


17 


United Arab Emirates 


75 150 


0.019 


- 0.883 


25 





3 


United Kingdom 


244 880 


0.024 


-1.003 


50 





12 
37 


United States of America 


9 372 614 


0.342 


0.638 


428 


105 


Uruguay 


186 925 


0.05 


-0.186 


81 


1 


6 


US Pacific Islands 


658 






- 








Uzbekistan 


447 400 


0.051 


-0.413 


97 





9 


Vanuatu 


14 765 


0.014 


-0.728 


11 


2 


4 


Venezuela 


912 045 


0.379 


1.398 


323 


19 


25 


Viet Nam 


329 565 


0.147 


0.737 


213 


9 


37 J 


Virgin Islands (British! 


153 






3 





° 1 


Virgin Islands (US) 


352 






- 





' 1 


Wallis and Futuna 


255 






1 





° 1 


Western Sahara 








32 


1 


3 .1 


Western Samoa 


2 840 






3 





' i 


Yemen 


477 530 


0.041 


-0.654 


66 


1 


' J 


Yugoslavia 


102 173 


0.046 


- 0.086 


96 





11 M 


Zambia 


752 615 


0.096 


0.074 


233 


3 


12 m 


Zimbabwe 


390 310 


0.099 


0.298 


270 





1 



APPENDIX 5 305 



nmals 


Birds 


Birds 


Birds 


Birds 


Plants 


Plafl| 


eatened 


breeding total 


endemic 


no. threatened 


% threatened 


total 


endei^ 


29 


278 


5 


7 


3 


5 050 


941 


23 


250 


24 


14 


6 


3314 


890 


50 


53 


9 


9 


17 


165 


50 





32 





1 


3 


659 


1 


11 


50 


4 


5 


10 


1 028 


11 


25 


108 


2 


2 


2 


1 166 


- 


9 


680 


1 


6 


1 


3 137 


50 


6 


603 





1 





5018 


- 


9 


364 





5 


1 


2715 


4 


13 


249 






2 
2 


1 
1 


1 750 
3 030 


1 


8 


193 


1 


6 


204 





8 


4 


3 000 


- 


21 


160 


u 


20 


13 


3 568 


- 


11 


- 





7 




5 000 


- 


U 


822 


24 
2 


33 
37 


4 
6 


10 008 

11 625 


1 122 


13 


616 


- 


5 


391 











3 085 


- 




5 





1 


20 


26 


- 


50 


37 


2 


3 


8 


463 


25 


1 


260 


1 


1 





2 259 


236 


U 


173 





5 


3 


2 196 


- 


15 


302 





11 


4 


8 650 


2 675 


13 


- 





6 




- 


- 




42 





3 


7 


448 


9 




9 





1 


11 
2 


4 900 


- 


6 


830 


3 


13 


- 


16 


263 





8 


3 


5 100 


- 


12 


67 





8 


12 


- 


- 


Ik 


230 


1 


2 


1 


1 623 


16 


9 


650 
237 


67 


54 


8 


19 473 


4 036 


7 







11 
-1 


5 


2 278 


40 


9 


_ 


1 
9 




4 800 


400 


36 


76 


9 


7 


9 


870 


150 


8 


1 340 


40 


24 


2 


21 073 


8 000 


17 


535 


10 


35 


7 


10 500 


1 260 





70 





2 


3 


- 


- 




70 





2 


3 


- 


- 





25 





1 


4 


475 


7 


9 


60 
40 



8 




6 





330 


- 


67 


15 


737 


- 


6 


U3 


8 


12 


8 


1 650 


135 


11 


224 





5 


2 


4 082 


- 


5 


605 


2 


11 


2 


4 747 


211 


4 


532 





10 


o 


4 440 


95 



= zero; - = no data. 



306 WORLD ATLAS OF BIODIVERSITY 



APPENDIX 6: 
IMPORTANT AREAS FOR 
FRESHWATER BIODIVERSITY 

This table presents information on areas 
identified as of special importance for diversity 
(species richness, and/or endemism) in the 
inland water groups treated (fishes, mollusks 
['MolL' in table], crabs, crayfish, fairy shrimps). 
This is a preliminary synthesis, designed to 
represent expert opinion on relative levels of 



Source: This information was prepared for WCMC on 
the basis of data kindly provided by a number of expert 
ichthyologists and members of the lUCN/SSC Specialist 
Groups for Inland Water Crustaceans and Molluscs. The 
particular source of information is indicated by letters in 
square brackets in the Remarks text. Please note that 
numerical estimates and other information may have 
been superceded by later survey and taxonomic work. 

GA Gerald Allen, in litt. March 1998. 

MK Maurice Kottelat, report compiled for 

WCMC^'. 
SK Sven Kullander. report compiled for 

WCMC'^ 
M5G Adapted from a report by lUCN/SSC Mollusc 

Specialist Group, primarily by Philippe 

Bouchet and Olivier Gargominy, and also by 

Arthur Bogan and Winston Ponder 
KC Keith Crandall, information on 

distribution of crayfish genera and species. 
DB Denton Belk, summary of fairy shrimp 

distribution patterns. 
NC/RvS Neil Cumberlidge and R. von Sternberg. 

report compiled for WCfvlC . 



diversity for each taxon at continent level. In 
the absence of global criteria for relative 
importance, areas on different continents do not 
represent strictly equivalent levels of diversity. 

The rows of data in this list are sorted 
first by continent, and secondly by the taxon 
concerned. 



For North American and African fishes, where source is 
UNEP-WCMC, information has been extracted from 
available literature, with additional data and advice for 
Africa from Christian Leveque and Guy Teugels. 

In most instances [MK, SK, MSG, NC/RvS, DB| 
contributors indicated the approximate location of the 
important areas concerned on a series of A3-5ized base 
maps provided. These areas, and those identified from 
literature by UNEP-WCfvlC. were digitized 
for purposes of presentation [Maps 7.2. 7.3, 7.U\ 
and analysis. 



APPENDIX 6 307 



i 


^rea name ^^^ 


^iUfe 


Remarks ^^^^^^^^^^^^^B 






^^BSRSSBST 




1 

■ 


L Tanganyika 


Crabs 


L, Tanganyika is Ihe only East African great lake where 
endemic species of freshwater crabs occur: of the 9 species 
and 2 genera present, 1 genus and 7 species are endemic. 
iNC/RvSl 


w 


Lower Congo 


Crabs 


Diversity is marked in the Congo R. basin, but appears highest 
in 2 areas, the lower parts oi the basin (including Congo, 
Cabinda and DR Congo (former Zaire]] and the upper reaches 
(including Rwanda/Burundi and parts of DR Congo]. [NC/RvS! 


3 


Madagascar 


Crabs 


4 genera and 10 species of freshwater crabs, all endemic, 
occur in Ivladagascar [NC/RvSl 


4 


Niger-Gabon 


Crabs 


Southeast Nigeria, southern Cameroon and Gabon: 3 endemic 
genera and more than 10 endemic species of freshwater 
crabs'. [NC/RvS] 


5 


Upper Congo 


Crabs 


Diversity is marked in the Congo R. basin, but appears highest 
in 2 areas, the lower parts of the basin (including Congo, 
Cabinda and DR Congo (former Zaire]] and the upper reaches 
(including Rwanda/Burundi and parts of DR Congo]. (NC/RvS] 




Upper Guinea 


Crabs 


Upper Guinean rainforest, centered on Guinea, Sierra Leone, 
Liberia, and western Cote d'ivoire (including Mount Nimbal: 2 


, 






endemic genera and 5 endemic species of gecarcinucids"'". 


E 






(NC/RvS] 


' 7 


Southern Africa 


Fairy shrimp 


2 endemic genera, 45 species, 38 endemic. South Africa 
proper: 34 species, 22 endemic. [DB] 


8 


Cape rivers 


Fishes 


With 4 families and 33 species the fish fauna of southern Africa 
IS rather poor in comparison with most other parts of the 


1 


L 




continent; most species are cyprinids. However, there is 
marked local endemism; most rivers in the southern Cape 
region have 3 or 4 native endemics (several species are 
threatened]. [UNEP-WCMC] 

m 



AFRICA 



I 



308 WORLD ATLAS OF BIODIVERSITY 



AFRICA 



w- 



9 Congo IZairel basin 



10 Congo 'Cuvette 
Centra le' 



11 Congo rapids 



12 Cross R. 



13 L Barombi-Mbo 



U L Bernnin 



15 L Malawi 



Fishes General region of very high richness; second only to the 

Amazon basin in species richness. 25 families and 686 species 
have been reliably reported from the Congo/Zaire basin, j 

excluding L, Tanganyika and L. Moero*. Around 548 of the 
species present lea 70%] are endemic to this basin. The basin 
can be divided into i sections; Upper Lualaba, Cuvette 
Centrale, Luapula-M»(eru and the rapids. [UNEP-WCMC] 

Fishes High richness plus marked endemism. Around 690 species 

occur in the Congo system; the Cuvette Centrale section 
possibly has the highest species richness owing to the great 
diversity of freshwater habitats available. lUNEP-WCMCl 

Fishes High richness plus marked endemism. The rapids between 

Kinshasa and the sea have a high concentration of fish species 
(150 species!. 34 of which are endemic to this section. The 
caves near Thysville are fed by the Congo system and support 
one of Africa's few true hypogean fishes Caecobarbus geertsi. 
Caecomastacembelus brichardi and Gymnanatlabes tihoni, not 
strictly cave fishes, have been collected in the Stanley Pool in 
riffles under flagstones or in crevices. [UNEP-WCMC! 

Fishes Nigeria-Cameroon. 42 families, 166 species'. Very high species 

diversity compared to the relatively modest catchment area, 
and marked endemism. Transitional ichthyofauna between 
the Nile-Sudan province and the Lower Guinea province. 
(UNEP-WCMC! 

Fishes This small (ca 4.5 km'! crater lake in Cameroon has 15 species 

(plus another 2 present in the inflow stream, not the lake 
proper!. At least 12 of the species are endemic, notably the 11 
cichlids that form 1 of the 2 recorded 'species flocks' in West 
Africa. 4 of the 5 cichlid genera are endemic: Konia, Myaka, 
Pungu and Stomatepia. This very important site is at risk 
from overfishing, the effects of introduced crustaceans and 
fishes, siltation from local deforestation and water pollution. 
(UNEP-WCMC! 

Fishes A small (ca 0.5 km'! crater lake in southwest Cameroon with 2 

non-endemic fishes and a remarkable species flock of 9 
tilapiine cichlids. The cichlids are very small and not exploited; 
they are at some risk because of the small distribution and 
deforestation in the surrounding area. (UNEP-WCMC! 

Fishes 30 800 km'. 1 2 families, more than 845 species, most of them 

endemic to the lake. Rich species flocks among Cichlidae, and 
a small species flock of Clariidae. (UNEP-WCMC! 



APPENDIX 6 309 



16 L. Tana 



17 L.Tanganyika 



18 L. Turkana 



19 L.Victoria 



20 Madagascar 



21 Niger basin 



22 NlemR. 



Fishes The lish fauna olthis targe 13 150 kml lake includes 21 

species in k families and is dominated by lake endemic 
cyprinids. The large Sarbus cyprinids form 1 of 2 recorded 
cypnnid species flocks Ithe other being that of L Lanao 
in the Philippines, many species of which are severely 
threatened!. (UNEP-WCMC] 

Fishes 32 000 km'. In the lake itself, 16 families, around 250 cichlid 

species and 72 non-cichlid species. Several species flocks 
are present not only in Cichlidae, but also among Clariidae, 
Bagridae, Mochokidae, Centropomidae. Ivlastacembelidae. 

Fishes 6 750 km'. 51 species, 35 genera, 17 families. High family and 

generic diversity, many of the species are lake endemic; 
cyprinids form the most diverse family [UNEP-WCIvlCl 

Fishes 68 800 km\ 12 families, around 545 species (many 

undescribedl. High species diversity dominated by cichlids. The 
majority of species are lake endemic. [UNEP-WCIvlCl 

Fishes Around UO fish species have been recorded from the brackish 

and freshwaters of Ivladagascar'; although species richness is 
not remarkable, endemism is high. Two endemic families 
IBedotiidae and Anchariidael have been recognized in 
Madagascar, as well as 13 endemic genera and 43 endemic 
species. Most endemic species are restricted to freshwater 
habitats, mainly in eastern forested regions. About one quarter 
of endemic species are known only from the type of locality. 
Blind cave fishes have been described from Madagascar: 
the gobiid Glossogobius ankaranensis and the elotrids 
Typheleotns madagascarensis and T. pauliani. [UNEP-WCMCl 

Fishes General region of high richness. 36 families, ca 243 species, 

with 225 primary freshwater species'. Endemism moderate: 20 
species endemic to Niger The basin includes 1 1 of the 13 
primary freshwater families that are endemic to Africa. 164 
primary freshwater fishes reported' from the Niger delta in 
Nigeria, based on reference specimens for each species; the 
high diversity 173% of the freshwater species in the entire 
basinl in this area is seriously threatened by oil pollution. 
[UNEP-WCMC) 

Fishes Cameroon. High richness for area, plus marked endemism. 16 

families. 94 species, 8 endemic. lUNEP-VCMCl 



AFRICA 



310 WORLD ATLAS OF BIODIVERSITY 



AFRICA 




23 Ogooue lOgowel R. 



2i Sanaga R. 



25 Upper Guinea rivers 



26 Volta basin 



27 L. Malawi 



28 L. Tanganyika 



29 L, Victoria 



Fishes Gabon. High richness for area, plus marked endemism. 23 

fan^ilies, 185 species, 48 species endemic (o Ogowe. A 
relatively small drainage basin wiith a very high concentration 
of species. Ivlany of the families represented are endemic to 
Africa. Available data certainly underestimate actual diversity 
(several new species are now being described, resulting from a 
project of Tervuren Ivluseum, the American Museum of Natural 
History and Cornell Universityl. (UNEP-WCMCl 

Fishes Cameroon. 21 families; high concentration of species in a 

small river basin; probably at least 135 land this figure is 
believed to be a significant underestimate!'. Between 10 and 18 
species endemic to the Sanaga. [UNEP-WCMC) 1 

Fishes High richness for area, plus marked endemism. The Upper • 

Guinea province includes coastal rivers from south of the ,' 

Kogon R. in Guinea to Liberia, and has faunal affinities with the ' 
lower Guinea province and the Congo/Zaire. The fauna includes 
many taxa endemic to the area"", fvlany small river basins, 
many of Ihem still poorly investigated. Konkoure R. (Guinea!: 
19 families, 85 species, at least 10 endemic. Kolente or Great 
Scarcies R. (Gumea-Sierra Leone!: 19 families, 68 species. 
Jong R. (Sierra Leonel: 20 families, 94 species. Saint-Paul R. 
(Liberia!: 19 families, 76 species. Cess-Nipoue R. (Liberia-Cote 
divoire!: 20 families, 61 species. (UNEP-WCMCl 

Fishes General region of high richness. 27 families, about 139 

species, 8 endemic to Volta basin. High species richness, with ,i 
9 of the 13 African endemic primary freshwater fish families 
represented". [UNEP-WCfvlC! 

MolL Gastropods: 28 species, 16 endemic. Bivalves: 9 species, 1 

endemic. (MSGI 

fvlolL Gastropods: 68 species, 45 endemic. Bivalves: 1 5 species, 8 

endemic. (MSGl 

MolL Gastropods: 28 species. 13 endemic. Bivalves: 18 species, 9 

endemic. (MSGl 



APPENDIX 6 311 




30 Lower Congo basin 



Moll. 



31 Madagascar 



32 Western lowland forest 
and VoKa basin 



33 The Mollucas, New Guinea 
and northerm Australia 

34 Southeast Australia 



35 Southwest Australia 

36 Fly R., Papua New Guinea 



37 Kikori R., L Kutubu, 
Papua New Guinea 



t 



38 Kimberley District, 
Western Australia 



39 Aikwa llwakal R., Irian Jaya 



40 Southeast Australia 



41 Southwest 

Western Australia 



Moll 



Moll. 



Crabs 



Crayfish 



The region downstream of Kinshasa in Congo and OR Congo 
(former Zairel. Gastropods: 96 species, 24 endemic. Endemic 
gastropods are almost all prosobranchs; 5 endemic 
'rheophilous' {specialized for life in the rapids] genera, 
belonging to the Bithyniidae ICongodoma. Liminitesta] and 
Assimineldae [Pseudogibbula. Septanellina, Valvatorbis]. 
Bivalves: no data. [MSGl 

Gastropods: 30 species, 12 endemic. Genus Melanatha 
endemic. Bivalves: no data. IMSGI 

Upper Guinea region in Ghana, Cote d'lvoire. Sierra Leone, 
Liberia, Guinea. Around 28 gastropod species of which 1 9 
endemic (and 9 near-endemicl. Bivalves: no data. [MSGl 

More than 30 species of freshwater crabs belonging to 5 
genera, all in Parathelphusidae. [NC/RvS] 



Large area of high richness and endemism, centered on 
Victoria, 35 species, and Tasmania, 1 9 species. IKCl 



Fairy shrimp 19 species, 12 endemic, [DB] 
Fishes 



Ih 



Fishes 



Fishes 



Fishes 



Fishes 



Fishes 




High species richness, 103 species in Fly proper, and high 
local endemism, 12 endemics in system. [GAl 

i 

Headwaters of Kikori and Purari systems, with L. Kutubu. 
High richness, 103 species and high endemism, 16 species in 
Kikori: plus 14 species in L Kutubu. [GA] 

14 endemic species (a density second only in Australia to 
Tasmania and equal to southwest Western Australia], including 
5 species within Prince Regent Reserve and 4 in the Drysdale 
R. area; and 47 species in total. (GA] j 

Near Timiki, Irian Jaya. High species richness: ca 78 species. 
IGA] 

I 
1 1 endemic species occur in coastal southeast Australia, a 

lower count per area than the other 3 areas cited here, and 42 

species in total, [GA] 

9 endemic species [i.e. density similar to the Kimberleysl, and 
14 species intotaL [GA] 



AFRICA 



AUSTRALASIA 



312 WORLD ATLAS OF BIODIVERSITY 



AUSTRALASIA 



EURASIA 



kl Tasmania 



43 Vogelkop. Irian Jaya 

kk Great Artesian basin, 
Australia 

45 l^ew Caledonia 



Fishes 



Fishes 



Moll, 



MolL 



46 Western Tasmania, Australia MolL 



47 Indonesia 



Crabs 



48 Myanmar-Malaysia 



Crabs 



49 South China 



50 South India 



Crabs 



Crabs 



51 Sri Lanka 



Crabs 



12 endemic species, a greater number per area than anywhere 
else in Australia, including 6 concentrated in the Central 
Plateau area; and 24 species in total. [GA] 

Moderate richness with high local endemism, ca 14 endemic 
species, including Triton and Etna Bay lakes, IGAl 



Springs and underground aquifers. Important area of 
gastropod diversity. Bivalves: no data, [MSGl 



<l 



Springs and underground aquifers. Gastropods: 81 species, 65 
endemic. Bivalves: no data. [MSGl , 

Springs and underground aquifers. Important area of j 

gastropod diversity Bivalves: no data. [MSGl 

The area containing Sumatra. Java, Borneo, Sulawesi and the 
southern Philippines has the greatest freshwater crab diversity 
in Indo-Australia, with representatives of the Parathelphusidae 
(10 genera and 71 speciesi and the Gecarcinucidae 15 genera 
and 21 speciesi. [NC/RvSl 

Northeast India lAssaml, Myanmar, Thailand, the Mekong 
basin in southern Indochina, to the Malaysian peninsula and 
Singapore. In this region there are an estimated 30 genera 
and more than 100 species of freshwater crabs in 3 families, 
the Potamidae, the Parathelphusidae and the 
Gecarcinucidae'"'. [NC/RvSl 

Only the Potamidae occur in China, but more than 160 species 
and subspecies in 22 genera are present, most of which are 
endemic. The southern provinces of China represent the 
hotspot of biodiversity for this country""'". [NC/RvSl 

The freshwater crabs of the Indian peninsula south of the 
Ganges basin are all endemic to the subcontinent and belong 
to 2 families, the Gecarcinucidae and the Parathelphusidae"". 
The west coast of the peninsula and the south show most 
diversity: an estimated 7 endemic genera and about 20 ■ 

endemic species in 2 families Ithe Parathelphusidae and 
Gecarcinucidael. A third freshwater crab family, the Potamidae, 
is found only in northern India but is not represented in the 
Indian peninsula. [NC/RvS] 

Sri Lanka has some 16 endemic species of freshwater crabs 
belonging to 3 genera, 1 of which ISpirabthelphusa] is 
endemic to the island'' ". iNC/RvS] 



APPENDIX 6 313 



52 Italy 

53 Borneo highlands 



54 Caspian Sea 



55 Central Anatolia 



56 Coastal peat swamps and 
swamp forests of Malaysia. 
Sumatra and Borneo 



57 Coastal rainforest 
of Southeast Asia 



58 High Asia 



Fairy shrimp 16 species, 7 endemic. IDBI 

Fishes The fish fauna of the highlands of Borneo seems to be poor 

in absolute number of species, but many of them have 
developed specialization for hill-stream habitats and are 
endemic to single basins. The area is still largely unsurveyed. 
About 50 known endemic species, but actual figure might be 
over 200" IMK] 

Fishes Moderate species richness. Although many species are shared 

with the Black Sea region, and/or the Aral basin, there is 
marked endemism, including the monotypic lamprey 
Caspiomyzon. ca 12 gobies, including monotypic genera Asra 
and Analirostrum. also 3 Absa. [UNEP-WCMCl ,: 

Fishes An and plateau with several endorheic lakes. About 20 

endemic species, apparently underestimated by inadequate 
taxonomy. Adjacent areas also have a number of endemics. In 
urgent need of critical reassessment: probably one of the most 
poorly known fish faunas in Eurasia. [MK] 

Fishes Includes Bangka island. Extent along eastern coast of Borneo 

not known. Probably formerly present on Java but apparently 
cleared. About 100 endemic species in peat swamp forests, a 
habitat type often restricted to a narrow fringe along the 
coasts, still largely unsurveyed. Although peat swamps are 
traditionally considered as a habitat with poor diversity, good 
data for limited areas in Malay peninsula and Borneo indicate 
that up to 50 species may be found within a small area (less 
than 1 km'l, about half of them endemic and stenotypic. Most 
species have small distribution ranges Isome possibly only a 
fewkm'PMMK] 

Fishes Thailand, Cambodia and southern Viet Nam. Southern extent 

not known accurately This habitat is largely destroyed in 
Thailand, and virtually unsurveyed in Cambodia and Viet Nam. 
Endemic species expected in peat swamp forests"'". [MK] 

Fishes Boundaries not known with accuracy; includes the Tibetan 

plateau and probably parts of Chinese Turkestan. Distribution 
and ecological data are sparse outside the Chinese literature. 
About 150 known fish species, about half of Ihem endemic to 
this area". Survey probably still superficial as a result of 
difficulties of access. [MKl 



EURASIA 



3U WORLD ATLAS OF BIODIVERSITY 



EURASIA 



^^^^^^I^^^^^^^^^B 


59 


Karstic basins of Yunnan, 


Fishes 


Boundary not known with accuracy. About U known species of 




Guizhou and Guangxi 




cave fishes. Survey is still superficial and numerous additional 
species are expected''. [MKl 


60 


L. Baikal, Siberia 


Fishes 


A species flock of 36 species of the family Cottidae Isculpins] 
(including the endemic family Comephoridael, 4 'ecologically 
differentiated stocks' Imany probably endemic species using 
Western concepts! of Coregonus. 2 of Thymallus, 2 of Lota". 
Endemic mollusks, gammarids, sponges and Baikal seal. [MKl 


61 


L. Biwa, Japan 


Fishes 


Reportedly 4 endemic species". [MKl ■1 


62 


L. El'gygytgyn. Siberia 


Fishes 


An old lake formed on the site of a meteorite crater 113 kml 
Total fish diversity: 5 species, including an endemic genus and 
species ISalvethymus svetovidovi], an endemic species 
[Salvelinus elgyticus], and 1 species endemic to eastern 
Siberia [Salvelinus boganidael. Endemic diatom species and 
apparently endemic invertebrate[sl*''. IMKl 


63 


L Inle, Myanmar 


Fishes 


About 25 native fish species, ca 10 of them endemic, including 
3 endemic genera"". [MK] 


64 


L Lindu, Sulawesi 


Fishes 


Very limited information. One native and endemic species; ,1 
others might be expected". (MKl 


65 


L. Poso, Sulawesi 


Fishes 


1 
10 native and endemic species, 2 endemic genera [both ' 

extinct?! and with L. Lindu comprises the entire known 

distribution of the subfamily Adrianichthyinae"". Additional 

species might still be expected, [MKl 


66 


L. Thingvalla, 


Fishes 


5 native hsh species, including 3 endemic Salvelinus [recent 
summary in Kottelat"!. [MKl 


67 


Lakes of Isles 


Fishes 


A number of lakes host 1 or 2 species of British Salvelinus, 
although information on individual lakes is usually inadequate. 
At the beginning of the century up to U species were ^ 
recognized; although generally not accepted under later ' 
systematic concepts, recent work suggests that this figure may 
be underestimated. Also at least 5 endemic Coregonus. 
1 endemic Clupeidae and potential for endemic Salmo [recent 
summary in Kottelat"!. [MK] . 



APPENDIX 6 315 




Lakes of Central Yunnan. Fishes 

China 



69 Lough Melvin, Ireland 



70 Lower Danube 



71 Mainland Southeast 
Asian hills 



Fishes 



Fishes 



Fishes 



Lakes Dianchi. Fuxian, Er Hai, Yangling, Yangzong, Xingyun, 
etc., have a distinctive fauna; despite the lakes being now in 
different river basins IMekong. Yangtze. NanpangjiangI, they 
have similar fauna, characterized by numerous endemic 
species in the genera Cypnnus, Schizothorax, Anabantius and 
Yunnanilus. Exact up-to-date figures of the number of species 
are difficult to extract from the Chinese literature, but vne have 
the following data: Oianchi: 25 native species. 1 1 endemic of 
which apparently all but 2 are extinct. The lake basin has 2 
other endemics'': Fuxian: 25 native species. 12 endemic plus 2 
endemic shared only with Xingyun"; Er Hai: 17 native species, 
9 endemic, several apparently extinct"; Yangzong has Ihad) at 
least 2 endemics; Yangling and Xingyun at least 1 each. [MKl 

Three endemic species of Salmo Irecent summary in 
Kottelat'l [MK] 

The lower Danube basin has a relatively richer fauna 
(especially more diverse communities! than any other 
European river. Endemics: about 6. possibly underestimated 
(counted in Kottelat 1997-1, [MK( 

Northern boundary not clear as published data on fish 
distribution land ground surveys! in southern China are too 
scanty Could be subdivided into a! upper Song Hong Imcludes 
hills of Hainan and southern Nanpang Jiang!; bl Annamite 
Cordillera; c! upper Mekong, ChaoPhraya and t^ae Khlong 
basins; dl Salween, upper Irrawaddy and southeastern Assam 
(including Tenassenm!. Recorded fish fauna estimated to be 
over 1 000 species Iwith an estimated 200-500 species still 
awaiting discovery!, 500 endemic to this area. Includes 
ca 400 known species endemic to headwaters of individual 
sub-basins. The fauna of the lower reaches of the main rivers 
(excluded from this polygon! is richer (in terms of the number 
of species that can be observed at a given locality! but most 
have wide distributions crossing several river basins-'"'". IMK] 



EURASIA 



316 WORLD ATLAS OF BIODIVERSITY 



EURASIA 



72 Malili Lakes, Sulawesi Fishes 



73 Maros karst, Sulawesi Fishes 



74 Mindanao, Philippines Fishes 



75 Northwest Mediterranean Fishes 

drainage 



76 Palawan, Philippines Fishes 



Most important single site for aquatic biodiversity In Asia, A 
complex of 5 lakes ITowuti, Matano, Mahalona, Wawontoa, 
Masapil with endemic radiations of fishes of the families 
Telmatherinidae 13 genera, 15 species, all but 1 endemici, 
Hemiramphidae 13 endemic speciesi, Oryziidae 13 endemic 
species!, Gobiidae lat least 8, all but 1 endemic], prawns lea 12 
species?], crabs 14 species?], mollusks lea 60 endemic 
species], etc. The distribution of the fishes is not uniform 
within the lakes, all but 1 of the species of L. Matano are 
endemic, while the others land 2 genera] are endemic to 
Towuti. Mahalona, Wawontoa. Masapi has not yet been 
surveyed. Only 2 species of the Telmatherinidae are known 
outside this area. [MK] 

One endemic genus ]possibly an artifact of limited collection; 
more surveys might show it to be present outside this area] 
and about 6 endemic species, including a cave species" [MKl 

About 30 endemic species of cyprinid fishes, including about 
18 endemic species of Puntius in L. Lanao lall but 2 or 3 
reportedly extinct]"" Cyprinids are fishes which live only in 
freshwater and cannot disperse in marine environments; 
several other families also occur in the island's freshwaters, 
but all are able to disperse through the seas. [MK] 

Includes Spam, Portugal, southern France and northern Italy. 
The total diversity in the whole area is quite low, the 
communities are quite poor, but this area holds 55 endemics, 
many with small distribution ranges. 3 of the Rhone endemics 
extend almost to the northern extremity of the basin. 
Endemics;! Petromyzonidae, 1 Acipenseridae, 1 Clupeidae. 
34 Cyprinidae. 5 Cobitidae. 6 Salmonidae, 1 Valenciidae, 
1 Cyprinodontidae, 2 Cottidae. 1 Percidae and 4 Gobiidae 
(counted in Kottelafl. [MK] 



About 10 recorded species of cyprinid fishes, actual figure 
probably higher [MKl 



d 



APPENDIX 6 317 




77 Southwest Balkans 



Fishes 



78 Southwest Sri Lanka 



79 Subalpine lakes 



Fishes 



Fishes 



Sundaic foothills 
and floodplains 



81 Western Ghats, India 



Fishes 



Fishes 



82 Balkans region 
{Former Yugoslavia, 
Austria, Bulgaria, Greece! 

83 ChilkaL 



84 L. Baikal 



Moll. 



Moll 



Moll 



The total diversity in the whole area is quite low, the 
communities are quite poor, but the area holds 84 endemics, 
most of them with restricted or very restricted distribution 
ranges: 1 Petromyzonidae. 2 Clupeidae, 48 Cyprinidae, 8 
Cobitidae, 1 Balitondae, 1 Siluridae, ISSalmonidae, 1 
Valenciidae, 1 Gasterosteidae, 1 Percidae and 7 Gobiidae. 
The systematics of many groups is still very poorly known 
and more species will be recognized or even discovered in 
the future (possibly 10-201. Noteworthy are L. Ohrid with 
apparently 4 endemic Salmo. L. Prespa with apparently 
7 endennic species and the Vardar basin with at least 8 
endemic species". [MK] 

28 of the 91 native fish species of Sri Lanka are endemic to 
this area. Several of the species traditionally given the same 
name as Indian species are being revised and turn out to be 
specifically distinct, so that the figure will rise'"". [MKl 

Stretches from L. Bourget in the west to Traunsee in the east. 
Numerous endemic Coregonus (possibly 27, several already 
extinctl, at least 2 endemic Salvelinus and possibly some 
endemic Salmo. Some lakes have more complex communities, 
e.g. L. Konstanz with 4 Coregonus, 2 Salvelinus. 1 Salmo and 
several other species (recent summary in Kottelaf. ||v1K] 

About 400 known species. Most of the floodplain species are 
widely distributed over the whole area, while those of foothill 
streams have more localized distributions and are of greater 
interest in terms of endemicity. Northern limit: Tapi basin in 
peninsular Thailand '"'". (MK( 

About 100 endemic fish species (estimated from Talwar and 
Jhingran*'; Pethiyagoda and Kottelat'^l. Difficult to give 
accurate figures. Many wide-ranging 'species' of fishes in 
South Asia are in fact complexes of species, so that the actual 
number of species is likely to increase significantly after 
adequate systematic revision. [MK( 

Springs and underground aquifers. Gastropods: ca 190 species, 
some 180 endemic. Bivalves: no data. (MSGj 



Brackish water Gastropods: 28 species, ca 1 1 endemic. 
Bivalves: 43 species, 25 endemic. (MSGl 

Gastropods: 147 species, 114 endemic. Bivalves: 13 species. 
3 endemic. (MSGl 



EURASIA 



318 WORLD ATLAS OF BIODIVERSITY 



EURASIA 



NORTH AMERICA 



85 L. Biwa 



L, Inle 



Moll, 



Moll. 



87 L, Ohrid and Ohnd basin Moll. 

88 L Poso and Malili Moll. 
Lakes system 

89 Lower Mekong R., Thailand, Moll. 
Laos, Cambodia 



90 North Western Ghats 



Moll. 



91 Zrmanja R., Croatia 

92 Southeast USA 

93 Western USA 

94 Bear L. 



Moll. 



Crayfish 



Gastropods: 38 species, 19 endemic. Bivalves: 16 species, 
9 endemic. [MSGl 

Gastropods: 25 species, 9 endemic. Bivalves: 4 species, 
2 endemic. IMSG] 

Gastropods: 72 species, 55 endemic. Bivalves: no data. [MSG] 



Sulawesi. Gastropods: ca 50 species, ca 40 endemic. 
Bivalves: 5 species. 2 endemic. [MSG] 



^ 



River habitat. Only ca 500 km of the lower 

Mekong mam course (with the tributary Mun R.I 

has been well studied. Gastropods: 121 species, 111 endemic. 

Two rissoacean groups dominate this entirely prosobranch 

assemblage of 120 plus species, the pomatiopsid Triculinae 192 

endemic species, 11 endemic general and the Stenothyridae 

119 endemic species). Bivalves: 39 species, 5 endemic. [MSG] 

River habitat. Gastropods: ca 60 species, 10 endemic. 2 
endemic genera Turbinicota, Cremnoconchus. The succineid 
genus Lithotis is known from 2 species: L. tumida not collected 
since its description in 1870. and L wpicola only known from a 
single locaUty. The highly localized genus Cremnoconchus is 
the only littorinid living in a freshwater/terrestrial environment. 
Bivalves: 1 1 species, 5 endemic. [MSGl 

Gastropods: all are hydrobioid snails, 11 species, 5 endemic. 
Bivalves: no data. [MSG] 

Large area of high richness and endemism at generic and 
species level; including the eastern and southern Mississippi 
drainage (Ohio R., Tennessee R., to Ozark and Ouachita 
mountains!; 72 species in Alabama, 71 in Tennessee. [KC] 



Fairy shrimp 26 species, 13 endemic. [DB] 

I 
Fishes This lake is part of the Bonneville R. basin and contains 1 

local endemic \Prosopium gemmiferum] and 2 species that are 
now restricted to this site [Prosopium spibnotus and 
P. 3bys5icot3l-\ [UNEP-WCMC] 




APPENDIX 6 319 




95 Colorado basin 



96 



Cumberland Plateau 
(Cumberland and 
Tennessee rivers! 



97 Death Valley region 




Eastern USA 




The largest basin of the western USA, this has high species 
richness and endemism, including 5 endemic genera of which 
only Plagopterus is monotypic''. About one third of the 
ichthyofauna of the Colorado is threatened, endangered or 
extinct due to dams and introduced species*'. lUNEP-WCt^Cl 

Fishes This area has the highest species richness and local 

endemism in North America. It is part of the highly diverse 
Ivlississippi basin, vjith ca 240 species in total. 160 present in 
both the Tennessee and Cumberland drainages, U endemic to 
the 2 basins. Of these U. 10 are darters, 3 are minnowis and 1 
IS a topminnow. The Tennessee has the greatest species 
diversity with 224 species including 25 endemics las well as 64 
not found in the Cumberland). The Cumberland has 176 native 
species, including 9 endemics and 16 species not shared with 
the Tennessee"'. [UNEP-WCMCl 

Fishes There is a high level of local endemism associated with the 

dispersed pattern of springs and marshes. 4 families are 
present ICypnnidae. Catostomidae. Cyprinodontidae and 
Goodeidael with 9 species including an endemic species of 
Catostomidae''. Several are globally threatened, including 
2 of the 5 Cypnnodon species ICypnnodon radiosus and j 

C, disbolisi lUNEP-WCMC) | 

Fishes This is a general area of high species richness and endemism 

which, with the possible exception of the incompletely known 
East Asian fish species, represents the most diverse of all the 
freshwater faunas of the temperate zone". This includes a) the 
Ozark Plateau, bl the Ouachita Mountains, cl the South Atlantic 
Central Plain and dl the Tennessee-Cumberland Plateau. 
[UNEP-WCMCl 



Klamath-upper Sacramento Fishes 



100 Ouachita Mountains 



The Klamath R. basin contains 28 species in total with 
relatively high endemism. The 6 endemic species include 2 
Catostomus. 1 Chasmistes and 1 Gila^. The ichthyofauna of the 
Sacramento differs from that of the Klamath and contains 4 
genera that are confined to this river and a few neighboring 
drainages'^IUNEP-WCMCl 



Fishes This area includes parts of the lower Red and Ouachita rivers, 

each containing 133 species. The Ouachita and the Red river 
systems both contain 18 endemic species^. lUNEP-WCMC] 



NORTH AMERICA 



320 WORLD ATLAS OF BIODIVERSITY 



NORTH AMERICA 



^pm^^^m^mm^^^^^g^ 


101 


Ozark Plateau 


Fishes 


The Ozark Plateau is an area of high species diversity and 
particularly high local endemism in the southeast USA; it 
represents a concentration of the species-rich southwestern 
Mississippi drainage (more than 30 endemic fish speciesP. 
[UNEP-WCMCl 


102 


Rio Grande-Pecos confluence 


Fishes 


The Rio Grande basin overall has more than 60 endemic 
species" and the Pecos, a tributary, has 5 " Many of the ' 
endemics occur at the confluence of the 2 rivers, and many 
are globally threatened. [UNEP-WCMCl 


103 


Southern Oregon-California 
rivers 


Fishes 


These rivers share few family similarities with the eastern 
USA and have about 25% of the number of species, but the 
region is high in local endemism. [UNEP-WCMC] 


m 


Southern Atlantic 
coastal plain 


Fishes 


This includes the Alabama-Tombigbee basin with a species- 
rich fauna, including about iO endemic taxa" This region 
also contains the Pearl R., with 106 species" and the species- 
rich lower Mississippi. lUNEP-WCMC] 


105 


And/semi-arid western USA 


Moll. 


Springs and underground aquifers. Gastropods: all are 
hydrobioid snails, ca 100 species, at least 58 endemic. 
Great radiation in genus Pyrgubpsis. 3 extinct species, and 
all others are candidates for listing by US Fish and Wildlife 
Sereice. Bivalves: no data. [MSGl 


106 


Cuatro Cienegas basin, 
Mexico 


Moll. 


Springs and underground aquifers. Gastropods: all are 
hydrobiids; 12 species, more than 9 endemic. 5 genera 
[Nymphophilus. Coabuilix, Paludiscala, Mexithauma, 
Mexipyrgus] are endemic to this small area of 30 x 40 km. 
Bivalves: no data. [d^SGl 


107 


Florida, USA 


Moll. 


Springs and underground aquifers. Gastropods: mostly 
hydrobiid snails. 84 species, ca 43 endemic. No bivalves. [MSGl 


108 


Mobile Bay basin 


Moll. 


Tombigbee-Alabama rivers. River habitat. Gastropods: 118 
species, 1 10 endemic; 6 endemic genera; greatest species 
richness (76 speciesl in Pleurocercidae. 38 of the gastropod 
species believed extinct. 70 candidates for listing by US Fish 
and Wildlife Service. Bivalves: 74 species, 40 endemic. 
25 extinct. [MSGl 


109 


Ohio-Tennessee rivers 


Moll. 


Eastern Mississippi drainage. River habitat. High species 
richness and endemism. [MSG] 



APPENDIX 6 321 




10 Central America 




111 South America 



112 Southern South America 

113 Altiplano of the Andes 

lU Amazon R, basin 




115 Aripuana R.. a tributary 
of the t^adeira 




Crabs The freshwater crabs of Central America belong to exclusively 

neotropical families, Pseudothelphusidae and Trichodactylidae. 
Central America from Ivlexico to Panama, including some of 
the Caribbean islands, holds at least 22 genera and over 
80 species of pseudothelphusid crabs, and 4 genera and 
ca 10 species of trichodactylids. The Isthmus of Tehuantepec in 
central Mexico is a hotspot of biodiversity for freshwater crabs 
in Central America'"", and richness declines toward to the 
south and north. The 7 species of freshwater crab belonging to 
1 genus found in Cuba are all endemic to that island" 
iNC/RvSl 

Crabs 2 freshwater crab families iPseudothelphusidae, 

Trichodactylidael endemic to the neotropics occur here. 
Freshwater crabs do not extend to southern Chile or 
southern Argentina, There are an estimated 17 genera and 
over 90 species of pseudothelphusids, found mainly in the 
highland regions of Peru, Ecuador. Colombia, Venezuela and 
the Guianas, and on the islands of the southern Caribbean, 
and 12 genera and over 40 species of trichodactylids in the 
Amazon basin. The Cordilleras of Colombia"", coastal 
Venezuela and the Guianas'"^ and the highland areas of 
Ecuador and Peru" are all diversity hotspots for freshwater 
crabs. The Amazon basin is rich in species*'", but most are 
widespread in the basin, and it is not possible yet to delimit 
special areas, [NC/RvSl 

Fairy shrimp 18 species, 14 endemic. (DBl 

Fishes Species flock of Orestias with 43 or more species, representing 

an endemic subfamily. Orestiinae. of the Cyprinodontidae. [SKI 

Fishes The Amazon Kvith adjacent TocantinsI basin probably has 

about 3 000 species, and is one gigantic hotspot. The Amazon 
fauna eguals or exceeds other continental faunas in species 
richness. Endemism in tributaries and subtributaries makes 
up most of the overall diversity, rather than the main Amazon 
itself. Only a few of the constituent rivers have been studied in 
any detail, [SK] 

Fishes Known to have a highly endemic but still little-studied fauna 

upstream of the lowermost falls, with at least 10 endemic 
species, some restricted to rapids. [SKI 




CENTRAL AND 
SOUTH AMERICA 



322 WORLD ATLAS OF BIODIVERSITY 



CENTRAL AND 
SOUTH AMERICA 



116 Central America 
between the Isttimus of 
Tetiuantepec and 

the Isthmus of Panama 

117 Iguacu R. 



Fishes 



Fishes 




La Plata basin: Uruguay, Fishes 

Paraguay and Parana rivers 




119 L Titicaca and smaller lakes Fishes 
of the Altiplano extending 
from Chile to Peru 




280 freshwater hsh species, all endemic. [SKI 



On the border between Argentina and Brazil, tributary to the 
Parana R. Its fish fauna is separated from the Parana by the 
Iguacu falls, which do not permit any migration; highly 
endemic, with ca 50 endemic species out of a total of 65 
species lea 80%l. There are considerable difficulties with the 
nomenclature and systematic status of the Iguacu fish species, 
most belonging to groups that have never been revised. 
Nonetheless, endemism will probably remain above 50%. The 
endemic fauna, mainly a running water one, is highly 
endangered by hydroelectric power projects, pollution and 
Introduced species. The fauna is not protected. [SK] 

fiiarked by numerous waterfalls providing isolation. 
Mainly endemic species, including numerous local endemics. 
Number of species unknown, but estimated fewer than 1 000, 
possibly ca 600. The tributaries of the Parana down to about 
Encarnacion have a very high number of local endemics, 
often restricted to a single river, mostly separated from the 
Parana by one or more waterfalls near the mouth. Many of 
these have not been described or examined by a specialist, 
but are known only from occasional collections made before 
the Italpu. Acaray and Yacyreta dams were constructed. 
Unfortunately, environmental impact assessment for those 
dams did not result in any significant collections to show what 
species were in the area before the dams were built. A lesser 
collection of pre-dam fishes is available In Museo Naclonal de 
HIstorIa Natural del Paraguay and the Museum d'histoire 
naturelle de Geneve, ISKl 

These lakes hold a large number of species of the genus 
Orestias ICyprinodontidael, 23 are endemic to Titicaca. 
The genus, with a total of 43 species, has a narrow range from 
northern Chile to southern Peru. The lake species flocks may 
not be monophyletic", but the group certainly attained its 
present species richness in the area. The sister group is the 
North American Cyprinodontidae. Other highland Andean fish 
families include the Astroblepidae, ranging from Bolivia across 
Peru and Ecuador into Colombia, and many trichomycterid 
fishes ITrichomycterldae] occur L. Titicaca with its Orestias 
fauna Is the only identifiable hotspot. ISKl 



APPENDIX 6 323 



^l^g^j^^m^^jj^^^^ 


120 


Marowijne/Maroni R. drainage, 


Fishes 


Known to have many endemic species above the falls, with 




Guyana and Suriname 




the same genera as in the rest of the Guianas area. [SKI 


121 


Mata Atlantica 

HP 


Fishes 


Numerous endemic species in small mountain streams or in 
the few major river systems, most incompletely known. The 
Ribeira R, has 77 species, and similar numbers appear to be 
in the other rivers. The Jequitinhonha is notable for several 
endemic species, including 1 of Rbamdia, which is othenwise 
represented by only a few widespread species in South 
America. The Mata Atlantica fauna extends to eastern Uruguay 
and southeastern Paraguay as numerous fragmented habitat 
patches, and although not high in species richness Iperhaps 
ca 1501. has a large number of locally restricted species, with 
related species replacing each other from one river to another. 


■ 






ISKl 


122 


Mazaruni and Potaro rivers, 


Fishes 


Separated from the rest of the Essequibo system by falls, with 




Guyana highlands 




several endemic species, but little explored. [SK] 


123 


Mesa Central, Mexico 


Fishes 


Endemic subfamily Goodeinae of family Goodeidae with about 


i 






36 species. [SK) 


124 


Mexican Plateau 


Fishes 


il 


125 


Negro R. and upper 


Fishes 


At least 700 species, probably nearer to 1 000. many of which 




Orinoco R., Brazil, Colombia 




are endemic to the clear and black waters distinguishing 




and Venezuela 




the basin. [SK] 


126 

1 


Nicaraguan lakes 

i 


Fishes 


The Nicaraguan great lakes [Nicaragua and Managua! in the 
San Juan basin do not have great numbers of species (about 
16 cichlidsl. but endemism is high [9 endemic species. 
2 endemic general. [SK] 


m 


Orinoco R. basin 


Fishes 


More than 1 000 species, most of which may be endemic. 
There is much local endemism as habitats vary considerably, 
including lowland inundation savannahs, fast-flowing mountain 
rivers, etc. Includes thus different biogeographic regions. [SK] 


128 


Oyapock R., Brazil and 


Fishes 


Known to have many endemic species, especially rheophilic. 




French Guiana 




from the lowermost falls upstream. Still little studied. [SK] 


129 


Pacific coast of Colombia 


Fishes 


Although a high-rainfall region there are few large rivers. 


k 


and Ecuador 




The fauna is poor, but species are highly endemic to the region 
and to particular rivers or portions of rivers. In particular, the 
Baudo R. and San Juan R. in Colombia seem to have 
numerous endemics. Possibly the Atrato R. should be 
included. KK^^^^^^ ^^ 



CENTRAL AND 
SOUTH AMERICA 



324 WORLD ATLAS OF BIODIVERSITY 



CENTRAL AND 
SOUTH AMERICA 



130 Patagonia lArgentma and 
Chile, from around 
the R. Negro southward - 
except the most and areas] 



Fishes 



131 Panuco R. basin 



132 Upper Uruguay R. 



Fishes 



Fishes 



133 Western Amazonia 



Fishes 



13^1 L, Titicaca 



MolL 



Low diversity but endemic relict fauna of more general 
southern hemisphere type, with families such as Geotriidae 
and Galaxiidae, and also the endemic catfish family 
Diplomystidae with 6 species, the monotypic catfish family 
Nematogenyidae. and k species of the percoid family 
Percicihtyidae. This is not a hotspot of species richness, 
but a region of considerable local endemism, and a fauna 
completely different from that of the rest of South America. 
The Nematogenyidae are related to the Loricariidae and 
Trichomycteridae of northern South America (the Brazilian 
faunal. but the Diplomystidae are the most primitive living 
catfish family The scaleless characid Gymnocharacinus bergii 
represents the Brazilian fauna, but lives isolated in one 
Patagonian locality on the Sumuncura Mountain which 
maintains about 22.5°C water temperature year-round. [SKI 

A small drainage with ca 75 species, ca 30% endemic, 
including several closely related species of Henchthys 
ICichlidael, Eastern Mexico, ca 25 endemic of ca 75 known 
species. ISKl 

The river is relatively well known from collections made by 
teams of the Museu de Zoologia of the Pontificia Universidade 
Catolica do Rio Grande do Sul during environmental impact 
assessment in the area, which is destined for numerous 
hydroelectric power plants. The collections concern the 
middle and upper portions, located in Brazil. More than 130 
species of fish are recorded from the middle and upper 
Uruguay, and the number is likely to rise to over 150 at least. 
About half of those may be endemic. Lucena and Kullander" 
described 1 1 species of Crenicichla from the Uruguay R.. and 
noted that this is double the number of a similar Amazonian 
river. 6 of the species form a species flock originating on site 
and diversifying by trophic adaptation similar to cichlids of 
East African lakes. The lower Uruguay R., along the 
Argentinian-Uruguayan border, is very little studied and 
may have fewer endemics, (SK] 

Lowland Amazonian Peru. Ecuador and Colombia, and parts of 
Brazil, representing a large expanse of lowland Amazonia, very 
rich in species, but not well studied. Work in Peru and Ecuador 
suggest that there may be at least 1 000 species in the area. 
and at least half may be endemic. [SK] 

Gastropods: 24 species. 15 endemic. Bivalves: no data. IMSGI 



I 



APPENDIX 6 325 



HH^ipH^mi^^^m^^B 


135 Lower Uruguay R and 


Moll 




Gastropods; 5i species, 26 endemic. Bivalves: 39 species, 


Rio de la Plata, Argentina, 






8 endemic. IMSGI 


Uruguay. Brazil 








136 Parana R. 


Moll. 




More than 7 species, 7 endemic, of vnhich 3 are extinct in the 






3KS0II 


wild. Bivalves: no data, IMSGI 



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41 Kottelat, M. 1997. European freshwater fishes. An heuristic checklist of the freshwater fishes 
of Europe (exclusive of fornner USSRl. with an introduction for non-systematists and 
comments on nomenclature and conservation. Biologia, Bratislava, Sect. Zool. 52 (suppl. 51: 
1-271. 

42 Kottelat, M. and Chu, X.-L. 1988. Revision of Kunnan/7us with descriptions of a miniature 
species flock and six new species from China (Cyprlnlformes: Homalopteridae]. Env. Biol. 
F/s/i. 23(1-21: 65-93, updated. 

43 Yang, J.-X. and Chen, Y.-R. 1995. [The biology and resource utilization of the fishes of Fuxian 
Lake, Yunnan]. Yunnan Science and Technology Press, Kunming [Chinese, English 
summary]. 

44 Li, S.-Z. 1982. [Fish fauna and its differentiation in the upland lakes of Yunnan). Acta Zool. 
Sinica 28(21: 169-176, updated [Chinese, English abstract]. 

45 Kottelat, M. 1990. Indochinese nemacheilines. A revision of nemacheiline loaches IPisces: 
CypriniformesI of Thailand, Burma, Laos, Cambodia and southern Viet Nam. Pfell, Munich. 

46 Kottelat, M. 1998. Fishes of the Nam Theun and Xe Bangfai basins ILaosj, with diagnoses of 
a new genus and twenty new species (Cyprinidae, Balltorldae, Cobltldae, Coiidae and 
Eleotrldidae). Ichthyol. Explor Freshwat. 9(1]: 1-128. 

47 Myers, G.S. 1960. The endemic fish fauna of Lake Lanao, and the evolution of higher 
taxonomic categories. Evolution 14: 323-333. 

48 Kornfleld, I.L. and Carpenter, K.E. 1984. The cyprinids of Lake Lanao, Philippines: 
Taxonomic validity, evolutionary rates and speciatlon scenarios. In: Echelle, A. A. and 
Kornfleld, I.L. (edsl. Evolution of fish species flocks, pp. 69-84. University of Maine at Orono 
Press, Orono. 

49 Pethiyagoda, R. 1991. Freshwater fishes of Sri Lanka. Wildlife Heritage Trust of Sri Lanka, 
Colombo. 

50 Pethiyagoda, R. 1994. Threats to the indigenous freshwater fishes of Sri Lanka and remarks 
on their conservation. Hydrobiologia 28b: 189-201. 

51 Kottelat, M. 1995. The fishes of the Mahakam River, East Borneo: An example of the 
limitations of zoogeographic analyses and the need for extensive fish surveys In Indonesia. 
Trop. Biodiv. 2(31:401-426. 

52 Talwar, P.K. and Jhlngran, A.G. 1991. Inland fishes of India and adjacent countries. 2 vols, 
Oxford and IBH Publishing Company, New Delhi, updated. 

53 Pethiyagoda, R. and Kottelat, M. 1994. Three new species of fishes of the genera 
Osteochilichthys (Cyprinldael, Travancoria (Balitoridael and Horabagrus (Bagrldae) from the 
Chalakudy River, Kerala, India. J. South Asian Nat. Hist. 1(1]: 97-116. 

54 Minckley et ai 1986. In: Hocutt, C.H. and Wiley, E.O. (eds]. The zoogeography of North 
American freshwater fishes. John Wiley and Sons. 

55 Banarescu. P. 1991. Zoogeography of freshwaters. 1. General distribution and dispersal of 
freshwater animals. Aula Verlag, Wiesbaden. 

56 Carlson, C.A. and Muth, R.T 1989. In: Dodge, D.P (ed.] Proceedings of the International 
Large River Symposium. Can. Spec. Publ. Fish. Aquat. Sci. 106. 



328 WORLD ATLAS OF BIODIVERSITY 



57 Starnes and Etnier. 1986. In: Hocutt, C.H. and Wiley, E,0. ledsl. The zoogeography of North 
American freshwater fishes. John Wiley and Sons. 

58 Cross, F.B. et at. 1986. In: Hocutt, C.H. and Wiley, E.G. (edsl. The zoogeography of North 
American freshwater fishes. John Wiley and Sons. 

59 Smith, M.L. and Rush Miller, R. 1986. In: Hocutt, C.H. and Wiley, E.G. (eds). The 
zoogeography of North American freshwater fishes. John Wiley and Sons. 

60 Swift, CO. etal. 1986. In: Hocutt, C.H. and Wiley, E.G. ledsl. The zoogeography of North 
American freshwater fishes. John Wiley and Sons. 

61 Alvarez, F. and Villalobos, J.L. 1991. A new genus and two new species of freshwater crabs 
from Mexico, Odontothelphusa toninae and Stygiothetphusa lopezformenti [Crustacea: 
Brachyura: Pseudothelphusidael. Proceedings of the Biological Society of Washington 
10^121:288-29^. 

62 Rodriguez, G, 1986. Centers of radiation of fresh-water crabs in the neotropics. In: Gore, R.H. 
and Heck, K.L. ledsl. Biogeography of the Crustacea. Crustacean Issues 4: 51-67. 

63 Chace, F.A. and Hobbs, H.H. 1969. The freshwater and terrestrial decapod crustaceans of the 
West Indies with special reference to Dominica. United States National Museum Bulletin 
292: 1-258. 

bU Rodriguez, G. and Campos, M.R. 1989. The cladistic relationships of the freshwater crabs of 
the tribe Strengerianmi ICrustacea, Decapoda, Pseudothelphusidael from the northern 
Andes, with comments on their biogeography and descriptions of new species. Journal of 
Crustacean Biology9: Hl-156. 

65 Rodriguez, G. and Pereira, G. 1992. New species, cladistic relationships and biogeography of 
the genus Fredius (Decapoda: Brachyura: Pseudothelphusidael from South America. 
Journal of Crustacean Biology 1 2: 298-31 1 . 

66 Rodriguez, G. and von Sternberg, R. 1998. A revision of the freshwater crabs of the family 
Pseudothelphusidae (Decapoda: Brachyura] from Ecuador Proceedings of the Biological 
Society of Washington 111. 

67 Rodriguez, G. 1982. Les crabes d'eau douces d'Amerique. Famille des pseudothelphusidae. 
Faune Tropicale 22, ORSTOM, Paris. 

68 Rodriguez, G. 1992. The freshwater crabs of America. Family Trichodactylidae and 
supplement to the family Pseudothelphisidae. Faune Tropicale 31 , ORSTOM, Paris. 

69 Magalhaes, C. and Tiirkay, M. 1996. Taxonomy of the neotropical freshwater crab family 
Trichodactylidae I. The generic system with description of some new genera (Crustacea: 
Decapoda: Brachyura). Senckenbergiana biologica 75(1-21: 63-95. 

70 Magalhaes, C. and TiJrkay, M. 1996. Taxonomy of the neotropical freshwater crab family 
Trichodactylidae II. The genera Forsteria, Melocarcinus, Sylviocarcinus and Zilchiopsis 
(Crustacea: Decapoda: Brachyura). Senckenbergiana biologica 75(1-2): 97-130. 

71 Magalhaes, C. and TiJrkay, M. 1996. Taxonomy of the neotropical freshwater crab family 
Trichodactylidae III. The genera Fredilocarcinus and Goyazana (Crustacea: Decapoda: 
Brachyura). Senckenbergiana biologica 75U-2\: 131-1/12. 

72 Parenti, L.R. 1984. A taxonomic revision of the Andean killifish genus Orestias 
ICypnnodontiformes, Cypridontidae). Bull. Amer. Mus. Nat. Hist. 178: 107-2U. 

73 Lucena, C.A.S. de and Kullander, S.O. 1992. The Crenicichia species of the Uruguai River 
drainage in Brazil. Ichthyol. Explor. Freshwat. 3: 97-160. 

74 WCMC 1998. Freshwater biodiversity: A preliminary global assessment. Groombridge, B. and 
Jenkins, M. World Conservation Press, Cambridge. 



INDEX 329 



Index 



abyssal plains 119, 142 
Aceraceae 83 
Acheulian tools 37, 39 
acid deposition 99, 185 
acritarclis 25 
Acrochordidae 126-7. 174 
Acrochordus granulatus 126-7 
Addax nasomaculatus 1 1 1 
advanced very high resolution 

radiometer lAVHRR] 

satellite sensors 73, 74-5 
Aepyornis maximus 53 
Africa 

dry forests 92. 94 

Great Lal<es 166. 181, 185, 

186, ]87. 307-10 

human origins 36, 37 

inland water biodiversity 

307-7 7 

mountain regions 101 

rain forests 86, 87, 88 

savannah 96, 104 

see also named countries 
Agenda 21 195 
agriculture 48 

crop biodiversity 40 

land devoted to 48-9, 104-5 

origins 33, 39-40 

traditional systems 45-6 

water consumption 178 
air 72 

Alaska pollock 145, 146 
algae 7, 1 1 

aquacuUure 149, 150 

marine/coastal 122-3, 124- 

5, 133, 138 
alien species 

deserts 1 1 1 

inland waters 185-6 

marine and coastal 

ecosystems 152-3 

Mediterranean-type 

ecosystems 107 



and recent extinctions 64-5 
almond 264 
alpaca 275 
alpha-diversity 78 
alpine tundra 99-102 
Alps 83, 101 
Amami rabbit 85 
Amazon basin 87, 88-9, 165, 

168, 182 
Amazonian varzea forests 87 
Amblyrhynchus cristatus 127 
Ambyostoma mexicanum 174 
Americas 

first human settlement 

36-7 

see also Central America; 

North America; South 

America 
amphibians 88, 174 

aquatic 174, 175 

areas of diversity 204-5 

recent extinctions 288 

threatened species 61, 188 
anadromous species 126, 172 
Anatidae 128, 175. 176 
anchoveta 144, 145, 146 
Andes 101 
Angiosperms see flowering 

plants 
Anguillidae 126. 172-3 
animal domestication 39-40 
Animalia 

estimated total species 18 

key features 20 

major extinctions 28-9 

marine phyla 122 

phyla 228-35 
Anopheles 172 
Antarctic krill 46, 47 
antelope 

Addax 111 

saiga 104 
Anthocerophyta 236 
Anthophyta see flowering 

plants 
ants 89. 107 



apple 263 
apricot 264 
aquaculture 

inland 179 

marine and coastal 148-51. 

153 
aquarium trade 150-1. 182 
Arabian Sea 140-1. 145 
arable lands see croplands 
Aral Sea 184 
Archaea 18-19. 122. 225 

described species 18 

key features 20 
arctic tundra 99-102 
arid environments 72. 73. 

108-9. 109-11 

wetland losses 754 
aroid. edible 183 
Arrhenius relationship 76-8 
artichoke 260 
Asia 

coral reefs 138 

forests 89, 92, 97 

human origins 38 

marine aquacuUure 150 

see a(so individual 

countries/regions 
ass 276 

Atlantic Ocean 125, 138. 139 
Atlantic salmon 153 
atmosphere 5. 6 
Australasia 

inland water biodiversity 

311-12 

see also individual 

countries 
Australia 

extinctions 52-3. 55 

forests 84. 85 

heath habitats 105-6, 107 

human origins 36 

open woodlands 96 
Australian Heritage 

Commission 188-9 
Australopithecus 36, 38 
autotrophs, defined 7 



Note: Page numbers in bold 
refer to figures or maps in 
the text; those m italics to 
boxed material or tables in 
the text or Appendices. 



330 WORLD ATLAS OF BIODIVERSITY 



AVHRR see advanced very 
high resolution radiometer 
(AVHRR) satellite sensors 

avocado 264 

Azollaceae 170 

Azraq oasis 184 



B 



bacteria 18-19 
classification 15-16, 226-7 
described species 18 
global biomass 46, kl 
key features 20 

Bactrian camel 274 

Baektu Mountain, China- 
Korea 82 

Bali cattle 273 

banana 254 

barley 245 

bathyal zone 1 19 

beans 250. 258-9 

beavers 178 

beeches 83. 84 

below-ground productivity 10, 
104 

beta-diversity 78 

beverage crops 255, 266-7 

Biaolowieza forest, Poland 50 

biodiversity change 195-6 
future scenarios 217-21 
responses 196-7 

biodiversity conservation 
defining priorities 199-204 
ecosystem maintenance 
198-9 

international agreements 
212-16 

protected areas 198 
restoration and 
reintroduction 210-12 
species protection 197 
systematic planning 203 

biodiversity indicators 201-2 

biodiversity information 17, 79 

biodiversity measures 76-8 

biodiversity-related 
conventions 213, 214-15 
see also individual 
conventions 

biogeography 79-80 

biomass 

below-ground 10, 104 
burning 38 

forests 84, 85, 89-90, 94 
global estimates 10-1 1 
grasslands 104 



human and livestock 46. 47, 

50 

marine 140-1. 154 

selected organisms 46 
biosphere 

defined 3 

extent 3-5 

human impacts 11-12 
biosphere reserves 210-11, 

216 
BirdLife International 63, 64- 

5, /?7, 201, 206-7 
birds 

biodiversity by country 295- 

305 

conse^^/atlon priorities 201, 

206-7 

global biomass 46 

grasslands 104 

inland waters 175, 176-7. 

189 

marine species 128-9, 147, 

155, 157 

recent extinctions 56-60, 

282-7 

shrublands 108 

temperate broadleaved 

forests 85 

temperate needleleaf 

forests 82 

threatened species 61-3. 

64-5. 189, 295-305 

see also individual groups 

and species 
bison 

American 49-50 

European 85 
Bison fa/son 49-50 
Bison bonasus 85 
Black Dragon fire 82 
blackcurrants 265 
BLI see BirdLife International 
'blitzkrieg' hypothesis 53 
bogs 167 

Boiga irregularis 64-5 
Bolivia 93 

Bonn Convention 214 
bonobo 89 
boreal forests 81-3. 84, 

106-7 
Sos primigenius 49 
bovids 

domestic 49-50. 272-4 

new species 16 
Brazil 36. 65. 96 
brazil nuts 91. 270 
broad bean 259 
Bromeliaceae 87 



brown algae (Phaeophytaj 

122-3. 124. 242 
brown tree snake 64-5 
Bryophytes 169-70, 236 
buffalo, water 273 
Bufo periglenes 99 
Bunolagus monitcularis 1 08 
bushmeat 42 
bycatch 147-8, 148-9, 158 



cabbage 249 

Californian shrublands 105, 

107 
Cambrian period 25, 26 
camel 

Bactrian 110, 274 

dromedary 275 
Camelus bactrianus 1 10 
Cams lupus 39-40 
Canis lupus baileyi 2M 
Cape flora 106-7 
capybara 178 
carbon cycle 7-12 

global budget 10-11 

oceans 154 

primary production 5-7 
carbon dioxide, atmospheric 

12 
carbon storage 

grasslands 104 

temperate forests 83, 84. 

85 

tropical dry forests 94 

tropical moist forests 89-90 

tundra 102 

see also biomass 
Carboniferous period 26, 27 
cardamom 268 
Caretta caretta 127 
Caribbean islands 53 
Caribbean sea 132 
carp 173. 174, 180, 188 
carrot 257 
cassava 248 
Castor spp. see beavers 
cat species 93, 107 
catadromous species 126. 

172-3 
Catagonus wagneri 93 
catchment basins 164-5. 168. 

185 

dams/reservoirs 184 

global high-priority 190-1 

indicators of habitat 

condition 188-90 



INDEX 331 



management 204-5 
catfishes 173. 174, 188 
cattle, domestic 49-50, 272-3 
CBD see Convention on 

Biological Diversity 
Central America 321 
Centres of Plant Diversity 

project 200 
Ceratopteris 1 70 
cereal crops 

major importance 24^-7 

secondary/local importance 

256 
cerrado 96 

Cetacea 130, 175, 177 
Chaco forests, Bolivia 93 
Chacoan peccary 93 
Chamela dry forest, Mexico 

93 
cfiaparral 105, 107, 108 
Cfiaraciformes 173. 174, 188 
charcoal production 80, 98 
cfiarophytes (stonewortsl 169 
Cfielonia 127-8, 135, 174, 175. 

182, 186 
Chelonia mydas 127-8, 135 
chemoautotrophs 142 
cfierry 26i 

Chesowanja, Kenya 38 
cfiickpeas 258 
Cfiile36, 84, 105-6 
cfiili pepper 268 
China 

aquaculture 150 

forest fires 82 

human origins 38 
chlorophyll 

ocean mapping 120 

terrestrial mapping 73, 

74-5 
Chlorophyta 138, 169 
Chromista 19 
cichlid fishes 185, 186, 187, 

278, 291-3 
Cisticota haesitatus 1 08 
CITES see Convention on 

International Trade in 

Endangered Species of 

Wild Fauna and Flora 
citrus fruits 262 
civil society 196, 197 
climate change 

arctic tundra 102 

and extinctions 30. 31, 54-5 

and fire 65, 82 

forests 8Z 99 

future scenarios 217 

human role 12 



marine and coastal impacts 

153-4 

Pleistocene history 33-5 

UN Framework Convention 

215. 217 
cloud forests 86, 91, 99 
Clovis hunting culture 36-7 
club mosses iLycophytal 170, 

237 
coastal regions 119, 125, 143 

alien species 152-3 

assessing health of 158 

climate change 154 

mangroves 88, 119, 125, 

132-3, 134-5 

pollution 151-2 

rocky shores 133-4 

seagrasses 134-6, 136-7 

threatened species 155-6 
cocoa 255 
coconut 251 
coelacanth 51 
coffee 267 
Committee on Recently 

Extinct Organisms (CREOl 

278 
community ecology 80 
condor, California 212 
Congo River basin 307 
conservation see biodiversity 

conservation 
Conservation International 

200 
continental shelf HS, 119 
Convention on Biological 

Diversity ICBD) 1, 196, 210- 

11,213,2?4 

ecosystems approach 198-9 

Global Taxonomic Initiative 

17 
Convention Concerning the 

Protection of the World 

Cultural and Natural 

Heritage see World 

Heritage Convention 
Convention on the 

Conservation of Migratory 

Species of Wild Animals 

[Bonn Convention! 214 
Convention on International 

Trade In Endangered 

Species of Wild Fauna and 

Flora (CITES) 8b. 214 
Convention on Wetlands of 

International Importance 

see Ramsar Convention 
coral reefs 119, 120. 122, 

136-8 



algal flora 125, 138 

climate change 153 

deep-water 138-9 

diversity 138, 140-1 

global area 137-8, 139 
hotspots' 126-7 
corals 

extinctions 30. 31 

live trade 151 
cotton seed 252 
country-based biodiversity 

90-1, 199-200, 295-305 
crabs 168. 182-3, 311. 312 

321 
cranlates see vertebrates 
crayfish 168. 182-3, 311. 318 
CREO see Committee on 

Recently Extinct Organisms 
Cretaceous period 26-7 

extinction event 29, 30. 31 
crocodile, estuarine 127 
Crocodilla 127, 174, 175 182 
Crocodylus porosus 127 
croplands 

expansion of 48-9, 97 

global distribution 76-7, 

108-9 
crops 

biodiversity 40 

major economic 

importance 41, 244-55 

origins 40-1 

secondary/local importance 

256-70 

wetlands 182-3 

see also individual crops 
crustaceans 230 

aquaculture 150 

areas of biodiversity 

importance 307. 311. 312- 

13. 321 

described and estimated 

total species 18 

inland waters 168. 172, 

182-3 

marine 123 

threatened species 61 
cucumber 266 
Curcubita see squash 
curlew, Eskimo 102 
cyanobacteria 3, 7, 11, 121, 

226 
Cyanospitta spixii 93 
Cynomys mexicanus 110-11 
Cyprmiformes 173. 174, 188 
Cyprinus carpio 1 80 
Cyrtosperma chamissonis 

183 



Note: Page numbers in bold 
refer to figures or maps In 
the text; those in italics to 
boxed material or tables in 
the text or Appendices. 



332 WORLD ATLAS OF BIODIVERSITY 



D 



dams 184 

date 26i 

deep-sea communities 141-3 

deforestation 185 

Dendroica kirtlandii 82 

Dermochelys coriacea 1 27 

desertification 1 1 1 

UN Convention 111, 215 
deserts 108-9, 109-11 
Devonian period 25, 26, 27, 

28, 30 
Diceros bicornis 96 
diet see human diet 
diet classes 44-5, 48-9 
Diptera 172 

dipterocarps 90, 93, 98 
dog 277 

domestication 39-40 

Mexican prairie 1 10-1 1 
dolphins 147 
domestication 

animals 39-40 

plants 40-1 
donkey 276 
dromedary 275 
drylands 72, 73, 108-9, 109-11 

wetland losses 184 
dugon 130, 155 



eagle 
Philippine 89 

white-tailed 212 
Earth 3 

age 24 

early life history 24-5 

land and water 71 

physical geography 4-5 

see also biosphere 
Earth Summit 1, 195, 196, 199 
East Pacific Rise 142 
EBAs see endemic bird areas 
Echlnodermata 122, 230 
ecoregions 80, 202 

Inland waters 168-9 

marine 120 
ecosystem, defined 74 
ecosystem diversity 74 
ecosystem maintenance 

198-9 
ecosystem maps 75, 106-7 
Ediacaran fauna 25, 28 
eels 126, 172-3 



EEZs, see exclusive economic 

zones 
eggplant 266 
eider, spectacled 102 
El Nino Southern Oscillation 

lENSOl events 65, 146, 147, 

253 
elasmobranchs 125, 128-9 
elements, essential 6-7 
elephant bird 53 
Elopiformes 173 
end-Permian mass extinction 

26, 28-9, 30, 31 
endangered species