Fundamentals of Biogeography (Routledge Fundamentals of Physical Geography)

TA

F BIOGEO

Fzlndamen~als of ~ ~ o g e o g ~presents a ~ ~ y an engaging and comprehensive introduction to biogeography, explaining the ecology, geography, and history of animals and plants. Defining and explaining the nature of populations, communities and ecosystems, the book examines where different animals and plants live and how they came to be living there; investigates how populations grow, interact, and survive, and how communities are formed and change; and predicts the shape of communities in the twenty-first century. Illustrated throughout with informative diagrams and attractive photos (many in colour), and including guides to further reading, chapter summaries, and an extensive glossary of key terms, Fzlndamentals of ~ ~ o g e o g ~ a ~ ~ explains y clearly key concepts, life systems, and interactions. The book also tackles the most topical and controversial environmental and ethical concerns including: animal rights, species exploitation, habitat fragmentation, biodiversity, metapopulations, patchy landscapes, and chaos.

F,mdamentals o f ~ ~ o g e o g presents ~ a ~ ~ y an appealing introduction for students and all those interested in gaining a deeper understanding of the key topics and debates within the fields of biogeography, ecology and the environment. Revealing how life has and is adapting to its biological and physical surroundings, Huggett stresses the role of ecological, geographical, historical and human factors in fashioning animal and plant distributions and raises important questions concerning how humans have altered Nature, and how biogeography can affect conservation practice.

Richard John Huggett is a Senior Lecturer in Geography at the University of Manchester

ROUTLEDGEFUNDAMENTALS OF PHYSICAL GEOGRAPHY SERIES Series Editor: John Gerrard Thls new series of focused, introductory textbooks presents comprehensive, up-to-date introductlons to the fundamental concepts, natural processes and humadenvironmental impacts wlthin each of the core physical geography sub-disciplines: Biogeography, Climatology, Hydrology, Geomorphology and Soils. Uniformlydesigned, each volumeincludesstudent-friendly features: plentifulillustratlons, studies, key concepts and summaries, further reading guides and a glossary.

boxed case

FUNDAMENTALS OF BIOGEOGRAPHY

Richard John Huggett

Routledge Fundamentals of Physical Geography

London and New York

First published 1998 by Routledge 11 New Fetter Lane. London EC4P 4EE Simultaneously published In the USA and Canada by Routledge 29 West 35th Street, New York, NY 10001

0 1998 Richard John Huggett The right of Richard John Huggett to be identified as the Author of thls Work has been asserted by him In accordance wlth the Copyrlght, Deslgns and Patents Act 1988 Typeset In Garamond by Keystroke, Jacaranda Lodge, Wolverhampton Prinred and bound in Great Brltaln by The Bath Press All rights reserved. N o part of thls book may be reprlnted or reproduced or utilized in any form or by any electronic, mechanical, or other means, now known or hereafter Invented, including photocopying and recording, or In any Information storage or retrieval system, without permmion in wrltlng from the publishers. Brrtrrh

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IS

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111 Prrbiirutron Data Huggett, Rlchard J. Fundamentals of hiogeographyiRlchard John Huggetr p. cm. - (Routledge Fundamentals of Physlcal Geography Serles) Includes bibliographical references and index 1. Biogeography. I. Title. 11. Series Q H 8 4 . H 8 4 1998 5 7 8 ' . 0 9 4 c 2 1 97-38998

ISBN 0-415-15498-7 (hbk) ISBN 0-41 5-1 5499-5 (pbk)

For m y family

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vii

CONTENTS

Serzes editor's pre/ke Lzst o f plates Lzst of jgures List of tables List o f Boxes Author's pr&z

INTRODUCTION: STUDYING BIOGEOGRAPHY 1 W H A T IS BIOGEOGRAPHY?

ix xi ... x111 xvi xvii xix

1

4

2 LIFE AND THE ENVIRONMENT

13

3 THE DISTRIBUTION OF ORGANISMS

43

4 POPULATIONS

84

5 INTERACTINGPOPULATIONS

109

6 COMMUNlTIES

138

7 COMMUNITY CHANGE

171

8 LIFE, HUMANS, AND MORALITY

209 226 235 240 254

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S E R I E S EDITOR’S PREFACE

W e are presently livlng In a time of unparalleled change and when concern for the envlronment has never been greater.Globalwarmingandclimatechange, possible rlsing sea levels, deforestatlon, desertlfication and wldespread soil erosion are just some of the issues of current concern. Although 1 t is the role of human activity in such issues that is of most concern, this actlvlty affects the operation of the natural processes that occur wlthin the physical environment. Most of these processes and their effects are taught and researched within the academic disclpline of physlcal geography. A knowledge and understandingof physlcal geography, and all it entails, is virally Important. It IS the aim of this Fvndamentuls of Physzrul Geogrupby Serm to provide, In five volumes, the fundamental nature of the physical processes that act on or just above the surface of the earth. The volumes In the series are Climatology, Geomorphology,Biogeography, Hydrology and Soi1.r. The topics are treated in sufficient breadth and depth to provide the coverage expected in a Fundunmtuls series. Each volume leads Into the toplc by outlining the approach adopted. Thls is important because there may be several ways of approaching lndivldual topics. Although each volume is complete initself, there are many explicit and implicit references to the toplcs covered in the other volumes. Thus, the five volumes together provide a comprehenslve inslght into the totality that is Physlcal Geography. The flexibility provided by separate volumes has been deslgned to meet the demand created by the variety of courses currently operating In hlgher educatlon instltutlons. The advent of modular courses has meant that physical geography I S now rarely taught, in its entirety, in an ‘all-embraclng’ course but I S generally split into its main components. This is also the case with many Advanced Level syllabuses. Thus students and teachers are being frustrated lncreaslngly by lack of suitable books and are having to recommend texts of which only a small part mlght be relevant to their needs. Such texts also tend to lack the detail required. I t is the aim of this series to provide lndivldual volumes of sufficient breadth and depth to fulfil new demands. The volumes should also be of use to sixth form teachers where modular syllabuses are also becomlng common. Each volume has been written by hlgher educatlon teachers wlth a wealth of experlence In all aspects of the toplcs they cover and a proven ability in presentlng information in a lively and lnterestlng way. Each volume provldes a comprehensive coverage of the sub~ect matter using clear text divided into easily accessible sectlons and subsectlons. Tables, figures and photographs are used where appropriate as well as boxed case

X

SERIES EDITOR'S PREFACE studies and summary notes. References to important prevlous studies and results are included but are used sparingly to avoid overloading the text. Suggestlons for further reading are also provided. The m a n target readership is introductory level undergraduatestudents of physical geography or envlronmental science, but there will be much of Interest to students from other disclplines and I t is also hoped that slxch form teachers will be able to use the lnformatlon that I S provided In each volume. John Gerrard, 1997

PLATES

COLOUR All colour plates appear in two plate sections between pp. 60-1 and pp. 156-7 1 Yellow-footed rock wallaby (Petrogale xanthrop/r.r) 2 Puma o r mountain lion (Felis concolor) 3 A cycad (Zatttia iitdenii) In Ecuador 4 Common cassowary (Casuurirrscas/urru.r) 5 Common wallaroo or euro (Macropus robustus) 6 Leadbeater’s possum (Gyrttnobelideus feadbeateri) 7 Honey possum or noolbender (Tarszpes rostratm) 8 Cattle egrets ( B / h I c u s ibis) 9 (a) Canadian beaver (Cartor tanadetrsr.r)(b) Beaver dam and lodge, Rocky Mountains Natlonal Park, Colorado 10 (a) Red squlrrel (Scrurns zwlgarrs) (b) Grey squmel (Scrwrds rarofinenszs) 1 1 Sklpper butterfly (He.rperra conmu), on the North Downs, Surrey 12 (a) Retlculated velvet gecko (Oedrrra retzcxfrta) (b) Glmlet gum (Eucalyptrrs .ralrrbris) in a wheat crop, near Kellerberrin, Western Austkalia 13 Golden lion tamarin (Leontidens rosalia)

BLACK A N D WHITE 1.1 Central Americm or Baird’s taplr (Tdprr1r.r bairdi), Belize 1.2 Nestor parrots (a) Kaka (Ne.rtor v/Nrrzdiorrulz.r)(b) Kea (Nestor notabi1i.r) 2.1 Splnifex hopplng mouse (Notoray.r rrlexis) - world champion urlne concentrator 3.1Tuatara (Sphenotlvnpzrnctatrrs)

8

9 26 50

xii

PLATES 3.2 Brown or spotted kiwi (Aptwix awtralis) 3.3 A relict Antillean insectivore (Solenodon) 3.4 Muntjac deer (Muntzacus reevesi) 5.1 Ratel or honey badger (Mellivma capensis) 7.1 Plant succession at Glacier Bay, Alaska (a) Upper Muir inlet (b) Goose Cove (c) Muir Pomt (d) York Creek 7.2 Secondary succession in Great Basm ghost towns, United States (a) Terrill (b) The sole remaining building (c) Wonder (d) The foundanon at Wonder 7.3 Successional pathways on Rakata (a) Casualina equiserifolta woodland (b) Lowland Neonauclea calycina forest ( c ) Lowland Ficus pubinwvis-Neonauclea calycma forest (d) Upland forest 7.4 Semi-permanent prairie wetland, Stutsrnan County, North Dakota

72 74 80 111 176 180

183 2 00

FIGURES

1.1 1.2 1.3 2.1 2.2 2.3 2.4

2.5 2.6 2.7 2.8 2.9 2.10 2.1 1 3.1

3.2 3.3 3.4 3.5 3.6

3.7 3.8

3.9 3.10 3.1 1

Breeding distribution of the ring ouzel Tapirs: their origin, spread and present distribution (a) Sclater and Wallace classification of faunal regions (b) Numerical classification of mammal distributions Tolerance range and l i m m Ecological valency, showmg the amplitude and position of the optimum Temperature control in poikilotherms and homeotherms Temperatures at an Esklmo dog's extremities Lethal ambient temperatures for four populations of woodrats, western United States Examples and explanation of climate diagram Ecozones of the world Soil and vegetation toposequences on Hodnet Heath, Shropshire Plant life-forms Proportion of plant life-forms in various ecozones Autoecological accounts for the bluebell in the Sheffield region Romanian hamster: a species with an endemlc and restricted distribution Two restricted and endemic plant families: Degeneriaceae and Leitneriaceae Two widespread plants: sunflower and grass families Zonal climatlc distributions of four plant families Protea, banksia, and grevillea family, and magnolia and tulip tree family Alpine marmot: a climatic relict (a) Scattergraph of the greatest north-south versus the greatest east-west dimension of North American snake specles ranges (b) Some shapes and sizes of ranges Home ranges (a) Land Iguana, Sante Fe (b) Red fox, Unlversity of Wisconsin arboretum Boreal forest in Canada (a) Predicted forest types (b) Observed forest types Predicted and observed distributions of longleaf pine and Florida poison tree Winter distributlon and abundance of the eastern phoebe

7

9 11 16 17 19 20

21 24 29 34 38 40 41 44 46 46 47 48 49

52 54 57 58 59

xiv

FIGURES 3.12 3.13 3.14 3.15 3.16 3.17 3.18 3.19 3.20 3.21 3.22 3.23 3.24 3.25 3.26 3.27 4.1 4.2 4.3 4.4

How spermatophyte flora reached Rakata, Krakatau Islands, 1883-1989 Widest ocean gaps crossed by terrestrlal animals Bird abundance on various islands between New Gulnea and New Brltaln Westward spread of the European starling in North Amerlca History of South American mammals Yapok or water opossum Convergent evolutlon of the marsuplal sabre-tooth and placental sabre-tooth Glyptodont: a tanklike Pleistocene herblvore descended from old xenarthran Invaders Unlque South Amerlcan mammals that evolved from Late Cretaceous condylarthran Invaders Bolivar Trough and Panamanlan Isthmus Distribution of Me.ra.ra//r//.r in the Permlan period Dlstributlon of Lystrusu~/r~~.r In the Early Triassic perlod Geological history of the Caribbean region over the last 100 million years Dlstributlon of Amerlcan mlnk and munt~ac deerIn Britain Effect of the Indian mongoose on Pac~ficisland lizards Effect of the chestnut blight in Watershed 41, western North Carolina Mam areas occupled by the red deer populatlon, Lyme Park, Cheshlre Exponential growth of a hypothetlcal population Logistlc growth of a hypothetical population Internally driven populatlon dynamlcs: stable points, stable cycles and chaos 4.5 Population crash in tlcklegrass: hlgh seed and low seed denslty monocultures 4.6 Survivorshlp curves of Dall mountaln sheep, Amerlcan robin and Brltish robin 4.7 Granville fritillary population in &and, Finland 4.8 Ecologlcal strategles in plants: competitor, stress tolerator and ruderal 4.9 The great auk's former distribution 4.10 Shrlnklng range of the Amerlcan blson 5.1 Two unpalatable Trinidadian butterflies resemble one another: Mullermn mimics 5.2 Distributions of native red squirrel and introduced grey squirrel in Britain 5.3 Vertical distributlon of larvae and adult acorn barnacles on intertldal-zone rocks, Millport, Scotland 5.4 Warblers in spruce forests, Maine, United States 5.5 Beak size of ground finches on the Galipagos Islands 5.6 Herbivore-plant interactions 5.7 Ant-eatlng mammals populatlon Canadian snowshoe inand density 129 lynx of hare 5.8 Cycles Ala~okl, prey, western predators Finland 130 and 5.9 Observed cycles In laboratory experments wlth protozoans as prey 5.10 Outcomes of Gause's and predator 131 5.11 Population cyclesin a laboratoryexperiment:apredatorymite feeds uponanothermite132 5.12 Populationchanges inalaboratoryexperlment:azukibean weevil andtwoparasitic wasps 6.1 Hertfordshlre Wood, Great Northaw ( a ) Geology (b) Soils 6.2 distributlon Tree Hertfordshire Wood, InGreat Northaw 6.3 Productlon zones, consumption zones and blogeochemlcal cycles In ecosystems biomes different in of blomass 6.4 Distribution net prlmary production for the biospherecomponent its and biomes 145 6.5 Mean patternWorldprlmary net of terrestrlal production 6.6

60

61 62

64 65 66 67 68 68

69 70 71 75 79 81

82 85 87 87 90 91 93

96 102 105 106 112 115 117 118 119 126 127

133 139 1.40

143 144 146

FIGURES 6.7 6.8 6.9 6.10 6.1 1 6.12 6.13

Grazing and detrital feeding relations in an ecosystem, showmg commlnutlon spirals (a) Carbon cycle (b) Nitrogen cycle (c) Phosphorus cycle (d) Sulphur cycle Carbon stored in blomass, litter, humus and charcoal in the major ecozones A food web, Wytham Wood, Oxfordshire Ecological pyramids: primary producers and a plant-parasite-hyperparasite food chain Sea otter distribution What holds communitles together? (a) 'The world is green' hypothesls (b) 'The world is prickly and tastes bad' hypothesis ( c ) 'The world I S whlte, yellow and green' hypothesis 6.14 Simple and complex food webs: repercusslons of removlng trophic levels 6.15 DDT biomagnlfication in the Long Island estuary food web, New York 6.16 Specles-area curve for Hertfordshire plants 6.17 (a) Herpetofauna (amphibians plus reptiles) diversity on West Indian islands (b) Bird species on Scottish ~slands 6.18 Spec~es diverslty,island area and habltat diverslty, Shetland 6.19 Latitudinal diversity gradient mammal species richness in the Amerlcds 6.20 Controls on biotic diversity In polar, temperate and tropical zones 6.2 1 Latitudinal variations in abiotic pressure and biotic pressure actlng on spec~esof three hypothetical body plans 7.1 Types of climax communities 7.2 Positlons of glacier t e r m m and Fastie's study sltes, Glacier Bay, Alaska 7.3 Location of the Krakatau island group 7.4 Secondary succession in Cedar Creek Natural History Area, Minnesota 7.5 Indivldualistic response of small North American mammals smce Late Quaternary 7.6 Land-cover transformatlon, A D 900-1977 7.7 Skipper butterfly decline in England during the twentieth century 7.8 Location of the Santee River and proposed river diversion 7.9 Bottomland forest community subjected to annual flood durations 7.10 Habitat changes in the Santee River study area 7.1 1 Atchafalaya Delta and Terrebonne Parish marshes study area, southern Louisiana 7.12 Observed distribution of habitats in the Atchafalaya-Terrebonne study area 7.13 Location of North American pralrie wetlands 7.14 Changes in biome area predicted by the MAPPS model under various scenarios 7.15 Geographical shift In blomes induced by global warming 7.16 Ecotone changes induced by global warming 7.17 Simulations of forest biomass dynamics over one millennium in response to climatic change induced by increasing levels of carbon dioxide in eastern North Amerlca 7.18 Simulated changes in species composltion of forests In eastern North America 8.1 Integration of ecologlcal, economlc and social needs in a decision-analysls model

148 150 152 153 153 155 157 158 160 161 162 164 165 167 168 172 175 177 179 184 187 188 190 192 193 194 195 199 201 202 203 204 207 220

xv

TABLES

1.1 2.1 2.2 3.1

3.2 4.1 4.2 4.3 4.4 4.5 5.1

5.2

5.3 6.1 6.2

6.3 6.4 6.5 7.1 8.1 8.2 8.3

Blogeographical regions and subregions, as defined by Alfred Russel Wallace Habitat scales Temperature tolerance in plants Species classed according to range size and cosmopolitanism Mean geographlcal ranges of Central and North American mammal species grouped by order Life table of a hypothetical population Parameters for a cohort-survival model of the blue whale (Balaenoptwa musculus) population Demographic parameters for a wandering albatross (Diomedza exdans) population Environmental contingencies and ecological strategies in plants A key for identifylng ecological strategies of herbaceous plants Interactions between population pairs Herbivorous tetrapods Main pest control technlques Mammals In Northaw Great Wood, Hertfordshlre Human appropriatlon of net prlmary production in the 1980s The diversity of livmg thmgs Habitat classification used in Shetland study Some factors thought to Influence species diverslty gradients Area occupied by each habitat type for three years for which data are available Brands of environmentalism and their characteristics Slogans for deep and shallow ecology Different attitudes of deep and shallow ecologrsts

12 14 22

45 51 92 94 95 100 101 110 122-3 134 141 147

161 164 166 196 214 216 2 17

BOXES

1.1 What's in a name? Classifylng organlsms 2.1 The electromagnetic spectrum emltted by the Sun 2.2Reptiles In deserts 2.3Mammals in deserts 2.4' Plant life-forms 3.1Accounting for regularities inrange size 3.2AcurlousSouthAmericanmarsupial 4.1 The red deer population In Lyme Park, Cheshlre, England 5.1 The hawthorn and the American robln: a frult-frugwore system 5.2 Populatlon models of herblvore-plantsystems 6.1 Removingtrophic levels - three ideas 6.2 Latitudinal diversity gradients, In genera, fiamilies and orders 7.1 Key features of FORFLO 7.2Amphibiansand global warmlng 8.1Deep ecology 8.2 Religlon, Nature and the World Wildlife Fund

6 18

25 26 38-9 53

66 85-6

124 I 26-7 157

168-9 191

198 2 16-17 218

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AUTHOR'S PREFACE

Biogeographymeansdifferentthingstodifferentpeople. To biologists, it is traditionallythehlstoryand geography of animals (zoogeography) and plants (phytogeography). Thls historical biogeography explores the long-term evolution of life and the Influence ofcontlnental drift, global climatic change, and other large-scale environmental factors. Its origins lie In seventeenth-century attempts to explain how the world was restocked by animalsdisembarklngfromNoah'sark.Itsmodernfoundatlonswere l a l d by CharlesDarwlnand Alfred Russel Wallace in the second half of the nineteenth century. The sclence of ecology, which studies communities and ecosystems, emerged as an dependent study in the late nineteenth century. An ecolog~cal element then crept into traditlonal biogeography. It led to analytical and ecologlcal biogeography. Analytlcal biogeography considers where organisms live today and how they disperse. Ecological biogeography looks at the relations between life and the environmental complex. It used to consider mainly present-day conditlons, but has edged backwards into the Holocene and Plelstocene. Physlcal geographers have a keeninterestinbiogeography.Indeed,somearespeclalistteachersinthat field. Biogeography courses have been popular for many decades. They have no common focus, their content varyingenormouslyaccordingtotheparticularinterests of the teacher.However,many courses show a preference for analytical and ecological biogeography, and many include human impacts as a major element. Biogeography is alsobecominganImportantelementinthegrowingnumberofdegreeprogrammes in environmental sclence.Biogeography courses In geographyandenvironmental science departments are supported by agood range offine textbooks.Popularworksinclude B~ogeograph~': N'ztwal nnd C h d (Simmons 1079),B L ~ . Biot/ts.rd/{hris)and salmon gum ( E N ( U / ~ / I ~ N .rise I duringthe twenty-firstcentury also poses a .w/t~/r)ttr~pbioiu). Much of the woodland has become m a p r threat t o coastal wetlands. The following two fragmented by clearing and remalns as Islands wlthin cases illustrate the complexity of changes in wetland a sea ofwheat (Colour plate 12b).For example, before habitats. land clearance, about 40 per cent of the Kellerberrin regmn contamed smooth-barked eucalypt woodland Bottomland forests in the Santee River suitable for reticulatedvelvetgeckopopulatlons. floodplain The same habltat now occup~es~ 1 s 2t per cent of the reglon. Only fourremnants,includingthreenature The coastal plain dramage of the Santee River, South reserves, contalnstands of morethan 570 gimlet Carolina,UnltedStates, was drastically modified gum and salmon gum trees. Demographic character- in 194 I . Almost 90 percent of theSanteeRlver istics of nine &dwu populatlonsthatcurrently discharge was diverted into the Cooper River as part survive In Kellerberrln eucalypt woodland remnants of a hydroelectric power pro~ect (Figure 7 . 8 ) . Howwere assessed and compared with Oec//~r'irpopulatlons ever,thls cliverslonof water has led tosilting In In three nature reserves. The woodland remnants vary Charleston Harbor, the main navigatlon channels I n in arm from 0.37 to 5.40 ha, but are slmilar in age, which requlre year-round dredglng. To allcvlate this

An Australian gecko

189

190

COMMUNITY CHANGE

20 km

0

Georgetown

VY

d

Figure 7.8 Location of the Santee River and proposed river diversion. Pearlstine et al. (1985)

Sourre: After

Aforestgrowthmodel,calledFORFLO, was costly problem, authorization was granted in 1968 to of thechanged redivert most of the water back into the Santee Rlverdevelopedtoquantifytheeffects regime on the bottomland forests throughan 18.5 kmlongcanalfeeding off Lake hydrological (Pearlstine et al. 1985) (Box 7.1).I t was used to simMoultrie(Figure 7.8). Anadditionalhydroelectric power plant wouldbe constructed on this canal.Fears ulate the effect of the proposed river rediversion, and a modified version of it, along a 25 km reach of the have been voiced that this rediversion would inunthe rediversion site date much of the Santee floodplain and cause a sub- Santeeforestedfloodplainfrom downstreamtoJamestown(Figure 7.8). Inthe stantial decline in the bottomland forest. After the diverslon, the Santee River would return to 80 per proposed rediversion, flow from Lake Marion to the cent of its pre-diversion flow rate with seasonal spates Santee Rlver staysthe same, flow to the Cooper River IS reducedto 85 m3/s(the level whlchprevents offloods and stages of low flows. A crucial change silting in Charleston Harbor), increasing the annual would be a reductlon of flow early in the growing flow to the Santee River vla the rediversion canal to season.

COMMUNITY CHANGE

Box 7.1

KEY FEATURES OF F O R F L O

is the duration of the annual flood. Each species has a tolerance to flooding, and will survive if its range of tolerance should fall within theflood duration for The FORFLO model allows hydrological variables to influence tree species composition through seed the plot. Third, the optimum growth of trees was reducedby, amongotherthings,awater-table germination, tree growth, and tree mortality. Key features of the model are the assumed relationships functionthatmodelsfloodplainconditions.The function modifies the tree-growth betweenfloodingandvariousaspects of forest water-table growth succession.First,for alltreespecies, save equation to account for the tolerance of species to the level of water on the plot during the growing black willow (Salix n i p ) and eastern cottonwood of water (Populus deltoides),seeds will not germinate when the season. The model computes the height ground is flooded.If the plot should be continu- for each half month during the growing season. It fail to grow during the ously flooded during that period of the year when a is assumed that all trees will species would germinate, then the germination of half months when they are more than three-quarters submerged by flood water. At lowerlevels of that species fails. Black willow and eastern cottonwood can germinate whether the land is flooded or submergence, tree growthwas related to water level by a curvilinearfunctioninwhichtheoptimum not. Second, after having germinated, the survival water-tabledepth foreachspeciesis takeninto of seedlings depends on environmental conditions. A notable determinant of the seedling survival rate account. L

The effects of theproposedrediversionandthe 413 m3/s. The modified rediversion was the same as modifiedrediverslonversionon the frequency of the proposed rediversion, except that during early the habitat types are indicated in Figure 7.10. In both growing season(April toJuly), flow throughthe cases there is a large loss of bottomland forest: a 97 rediversioncanalwouldbekepttoalevelwhich could be handled by just one of the three turbinesat per cent loss in the case of the proposed rediversion and a 94 per cent loss in the modified rediversion. the power station. Although this would mean that The saving grace of the modified rediversion plan is the Cooper River would exceed the critical flowof 85 m3/s for the four months of the growing season, that a forest cover is maintained bottomland forest changes to cypress-tupelo forest, rather than to open and thus cause some silting at the coast, it might water. If these predictions of sweeping changes In promote the preservation of the bottomland forest. The simulated responses of the forests to the re- the bottomland forest along the Santee River should be advisable diversion are shown in Figures 7.9a and b. The annual be trustworthy, then plainly it would duration of flooding is a crucial factor in determining to rethink the plans for rediverting the flow from the Cooper River. the course of vegetational change. With an annual flood duration of more than 30-35 per cent (Figure 7.9a),bottomlandhardwoodforestis replaced by A coastal ecosystem in southern Louisiana cypress-tupeloforest,baldcypress (TaxodiumdisEcosystems occupyingcoastallocationsareunder tzchum) and water tupelo (Nyssa uquatzcu) being the only species that would manage to regenerate. When threat from a variety of human activities: gas and oil exploration, urban growth, sediment diversion, and annual flood duration was above 65-70 per cent, no specles was able to survive and the forest was replaced greenhouse-induced sea-level rise, to name but afew. by a non-forest habitat; this happened more rapidly inTo protect and preserve these ecosystems, it is valuable to know what the effects of proposedhuman the subcanopy.

19 1

192

COMMUNITY CHANGE

(b)

Canopy Bald cypress

/

Water tupelo

‘1,

i 1

Bald Mockernut hickory

5

X

“E

e “t

-

0 lo

I

1 Sub-canopy

0

60

1

120

180

0

I

I

120

180

Sub-canopy

60

Time (years) Fzxrrre 7.9 Results of simulations. (a) Bottomland forest community subjected to an annual flood duration of 45 * 4 per cent. (b) Bottomland forest community subjected to an annual flood duration of 72 t 5 per cent. The flood duration is the percentage of a year during which a plot is flooded. The plant species are bald cypress (Taxodiut~rdfJftfbttirJl), water tupelo ( N ~ J Jayrrarrfu), U and sassafras ( S a ~ ~ a f a/brdtm) va~ Sorwe: After Pearlstine et al. (1985)

activities are likely to be, and how these effects differ from natural changes. These questions were addressed in the Atchafalaya Deltaand adjacent Terrebonne Parish marshes in southern Loulsiana, Unlted States, using mathematical a model to simulate likely changes(Figure 7.11) (Costanza et al. 1990). This landscape, part of the Mississippi River distributary system, is one of themost rapidly changingland-

scapes in the world. The Atchafalaya River is one of thetwoprincipaldistributaries of the Mississlppi River. It carrles about 30 per cent of the Misslssippi dischargetotheGulf of Mexico. Since themid1940s, sedimentstransported by the Atchafalaya River have been lald down in the bay area. In consequence, the bay area has gradually become filled In, and in 1973 a new subaerial delta appeared that has

COMMUNITY CHANGE scenarlos consider what the system would have been like had not the environment been altered by human actlon, and if climatic conditions had been different. Boundary scenarios delve Into the potentla1 impacts of naturalandhuman-inducedvariationsinthe boundary conditions of the system, such as sea-level rise. The management and boundary scenarios were run by restartlng the model with the actual habitat map for 1983, rather than the predicted habitat map for that year, toaddmorerealism.Climateand historical scenarios were run startlng in 1956,so that the full impacts of climate and historical varlations Present After proposed After modified could be assessed. After 1978, the rate of canal and rediverslon rediverslon levee construction for oil and gas exploration slowed Figure 7.10 Habitat changes in theSantee Rwer study appreclably compared with the 1956-78 period, so area. all scenarios assumed that no canals nor levees were Sourre: After Pearlstine et al. (1985) built after 1978, except those specifically mentloned In the scenarlos. smcegrowntoabout SO km2.Otherchangesare It is clear from Table 7.1 that the assumptlons taklngplaceinthe area. The westernTerrebonne made about climatic change have a big influence on marshesarebecoming lesssalty,whiletheeastern part of the area is becoming more salty:the boundary habitatdistributlon by the year 2033.Themean between fresh and brackish marsheshas shifted closer climate scenario, wherein the long-term average for to the Gulf in the western marshes, and farther inland each varlablewas used for all weeks,produced a modin theeast (Figure 7.12).As a whole,the study region est loss in land area. On the other hand, the weekly is losing wetland, but rate of loss in the Terrebonne average climate scenario, wherein each weekly value marshes has slowed and reversed since the mid-1970s of each climatic variable for the entire run from 1956 owing to m e r deposition. The hydrology of the area to 2033 was set to the average value for that week has been greatly altered by dredging of waterways in the 1956-83 data, produced a drastic loss of land flood andthediggingof access canals for petroleum area. Thisfindingindicatedthattheannual cycle and other annualcycles In climatic variables are exploration. importanttothelandbuilding process, butthat Adynamlcspatlalmodel was developedand chance events, such as major storms and floods, tend christened the Coastal Ecological Landscape Spatial the Simulation model(CELSS model for short).It divided to have a net erosional effect on marshland. If globalclimateshouldbecome less predictablein the marsh-estuary complex Into 2,479 square gridcells, each with an area of 1 km’.The model was the future as a result of global warming, then the stability of coastal marshes may be in jeopardy. used to predict changes of habitats under a range Several management scenarlos were run. As the data of climatic,management,hlstorical,andboundary In Table 7.1 Indicate, the largest loss of land would scenarios. The results of severalscenarloanalyses, which predicted changes to the year 2033, are sum- arise from the full six-reach levee extension scheme that hadbeenconsidered atonetlme.Withthis marized in Table 7.1. Climate scenarios take ‘climate’ scheme, 48 km2 of brackish marsh and 6 km2 of fresh to mean all the driving variables including rainfall, by 2033,largely because the AtchafalayaRivet flow, wind,and sea-level. They marshwouldbelost address the impact of climatic changes on the study extended levees prevent sediment-laden water reacharea. Management scenarios lookat the effect of spe- ing the brackish marsh bordering Four League Bay, cific human manipulations of the system. Historical wheremost of the loss occurs.Boundaryscenarios 8.000

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COMMUNITY CHANGE

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considered the effects of projected rates of sea-level rise, both high and low projections, on the area. The results were unexpected. Surprisingly, doubling the rate of eustatic sea-level rise from 0.23 to 0.46 cmlyr caused a net gain in land area of 10 km2 relative to the base case. This was probably because, so long as sediment loads are high, healthy marshes can keep pace wlth moderate ratesof sea-level me. Two historicalscenariosweretested. The first

probed the changes in the system that might have taken place had not the original Avoca Island levee been built. The second considered the changes that might have ensued had not the Avoca levee nor any of the post-1956 canals been constructed. The results shown in Table 7.1 suggest that the original levee and the canals had a major influence on the development of the system, causing a far greater lossof land than would have occurred in their absence.

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GETTINGWARMER:COMMUNITIES IN T H E T W E N T Y - F I R S T C E N T U R Y Individuals and communities, by processes of natural selection, become adaptedtotheenvironmentin which they live. If that environment should change, life systems must adapt, move elsewhere, or perish.

Species under pressure The fate of many species in the twenty-first century, as the world warms and habitats shrink or vanlsh, is worrying (Peters 1992a, b). It seems that extinctlon will continue apace and global biodiversity will drop. But which species are the most vulnerable to global warming? Thesafest species are mobile birds, insects, and mammals who can track their preferred climatic zone. In the Brltlsh Isles, the white admiral butterfly (Ludoga ranriiia) and the comma butterfly (Polygonia c-album) expanded their ranges over the past century as temperatures rose by about0.5"C (Ford 1982). However, even mobile specieswould need a food source, andtheir favourlte menuitemmight be extinct or have wandered elsewhere. Five kinds of species appear to be most at risk: 1 Peripheralspecies.Populatlons of anlmals or plants that are at the contractlng edgeof a species range are vulnerable to extinction. 2 Geographicallylocalizedspecies. Many currently endangered species live in alarmlngly limited habltats. The golden lion tamarin (Leontidexs vosaiicr) I S a small primate that lives in lowland Atlantic forest, Brazil (Colour plate 13). Lowland Atlantic forest is one of the most endangered rain forests in theworld. I t once covered 1 million km' along the Brazilian coastline. Now, a mere 7 per cent of the original forest remains, putting the golden lion tamarin under enormous threat (but see p. 212). 3 Highly specialized species. Many species have a close assoclatlon with only one other specles. An example is the Everglade (or snail)kite (Rostrhamus sociabiiis) that feeds exclusively on the large and colourf~~l Ponzuceu snail inFlorida wetland.

Swamp drainage has already robbed the kite of its prey in some areas. 4 Poor dispersers. Manytrees have heavy seeds that are not broadcast far. Plants with limited dispersal abilities have knock-on effects alongthe food chain. Some birds, mammals, and Insects are closely associated with specific forest trees, which cannotmigraterapidly.These species wouldbe hard pressed to survive. TheendangeredKirtland's warbler (Dendrozca kirtiandii), for example, breeds in and nests only upon the ground under jack-pme (Pinus banksiana) forest on well-dralned sandy soil in north-central Michigan, United States. But notany jack-pine forest will do - it has tobeyoung, secondary growth forest that emerges In the aftermath of a forest fire. As the globalthermometer rises, the jack pines may move northwards onto less well-dramed soils, and theKirtland's warbler may find Itself without any suitablenestingsites(Botkin et ai. 1991). This specles could be the first casualty of global warming. Climatically 5 sensitive species. Specles in climatlcally sensitive communities are vulnerable toglobalwarming.Communities in thls class includewetlands,montaneandalplne biomes, Arctic biomes, and coastal biomes. Wetlands will dry out, wlth grave consequences for amphibians (Box 7.2), mountainswill become warmer towards their tops, tundrareglons will warmup, andcoastal biomes will be flooded. Climaticwarmmg IS predicted to be greatest at high latitudes. Anlmals and plants In these reglons will have to cope with themost rapidchanges. Tundra vegetation may be pushed northwards as much as 4 degrees of latitude.This could mean that, for aclimatic warmlng of 3"C, 37 per cent of present tundra will become forest. The same degree of climatic warmingwould fuel local temperature increases on mountains. Animals and plants would need to shift thelr ranges upwards by about 500 rn. Animal and plant populat~ons on mountains may retreat upwards as climate warms, but eventually some of them will have nowhere to go. Thls would probably happen to the pikas (Othotona spp.) living on

197

198

COMMUNITY CHANGE Canadian prairie provinces (Figure 7.13) (Poiani and alpinemeadowsonmountainsofthewestern Johnson1991).Itisthemainbreedingarea for United States. Pikas feed on grasses and do not range into boreal forests. Asthe boreal forest habi- waterfowlon theNorthAmericancontinent.The tat advances up the mountainsides, so the pikas habitat consrstsof relatively shallow-water temporary for a few weeksinspring), will retreat into an ever-decreasing area that may ponds(holdingwater early one day vanish. This processisalready happen- seasonal ponds (holding water from spring until seml-permanent ponds (holding water ingtoEdith’scheckerspotbutterfly (Euphydryas summer), throughoutmuch of thegrowing season inmost western North America editha), which lives 7.4). For (Parmesan 1996). Records of this species suggest years), andlargepermanentlakes(Plate that its range is shifting upwards and northwards, breedingpurposes, the waterfowlrequireamixof open water and emergent vegetation. The temporary and extinctions have occurred at some sites. and seasonal pondsprovidearich foodsourcein the spring and are used by dabbling ducks. The semiCommunities under pressure permanent ponds provide food and nesting areas for Community composition and structure change in the birds as seasonal wetlands dry. As temperatures rise face of environmentalperturbations.Durmgthe over the next decades, water depth and the numbers twenty-first century, many communities will be perof seasonal ponds in these habltats should decrease. turbed by global warming. They will affect the geoThe lower water levels wouldfavourthegrowth graphical location of blomes and the species of emergent vegetation and reduce the amount of composition of communities. openwater.Waterfowl may respond by migrating to differentgeographicallocations,relyingmore Prairie wetlands in North America uponsemi-permanentwetlandsbutnotbreeding, as theydoatpresentduring Prairiewetlandhabltatoccupiesabroadswath of ot failingtore-nest droughts. landacross theAmericanMidwestandsouthern

Pox 7.2 frogslivinginwetstream-bankpatches.These amphibians retreat into crevices or gather in damp pockets during the dry season. Four years later, the golden toad and the harlequin frog had vanished. 4mphibians need water. They must live in it, near Rainfall during May 1987 was 64 per cent less than t, or else in very humid conditions. Warmer and average, the lowest ever recorded for that month. jrierconditionscould bedisastrousforthem.In Normally, early rains in May and June helpthe ?laces whereglobalwarming is associatedwith amphibians to recover from the November and Increasing aridity, they are likely to suffer. Evidence December dry season. The 1987 dry season was af this comes fromendemicgoldentoad (Bnfo kiglenes) and harlequin frog (Ateiopus uavius) popu- extradry.Duringthedrought,watertablesfell, springsallbutdriedup,andstreams fedby iationsintheMonteverdeCloud ForestPreserve, Costa Rica(Pounds1990;Poundsand C m p aquifers, which would usually give enough water to keep the mossy riverbanks wet, dwindled. As the 1994). In May and April 1987, there were thoube that sands of golden toads thriving along streams in the amphibiansdisappearedsuddenly,itmay theextremedroughtkilledtheadultsandtheir reserve. The maleslivein undergroundburrows tadpolesoreggs.Thiscatastropheaugurs illfor near thewatertable;theyemergeonlytomate. There was also a considerable number of harlequin amphibians during the twenty-first century.

4MPHIBIANS AND GLOBAL WARMING

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Mires in the Prince Edward Islands TemperaturesinthePrinceEdwardIslands have Increased by approximately1°C since the early 1950s, while precipitation has decreased (Chown and Smith 1993). The changing water balance has led to a reduction in the peat-moisture content of mires and hrgher growingseason 'warmth'. In consequence, the temperature-sensitive and moisture-sensitive sedge, Uncinra compacta, has increased its aerial cover on Prince Edward Island. Harvestingof seeds by feral mice (Mus musculus), whichcanstrip areas bare, has prevented an increase in sedge cover on Marion Island. Such extensive use of resources suggests that prey switching may be taking place at Marion Island. Mice not only are eating ectemnorhinine weevils to agreaterextentthanfound rn previousstudies of

populations at Marion Island, but also prefer larger weevils. A decrease in body size of preferred weevil prey specles (Bothrumtopusrandi and Ectemnorhinus similis) hastakenplaceonMarlonIsland (19861992), but not on Prince Edward Island. This appears to be aresult of increased predation on weevils. Adults of the prey species, Ectmnorhinus srmilis, are relatively more abundant on Prince Edward Island Ectemnorhinus thanadults of thesmallercongener m a n o m , and could not be found on Marion Island in the late southern summer of 1991. Results not only provlde support for previous hypotheses of the effect of globalwarmingonmouse-plant-invertebrate interactions on the Prince Edward Islands, but also provide limited evidencefor the first recorded case of predator-mediated speciation.

199

200

COMMUNITY CHANGE

PI7.4 Semi- nnanent prairie wetland, Stutsman County, North Dakotu. This wetland is part of the CoftonwooS(Cake site where long-term hydrological and vegetation data have been collected b theUnitedStatesGeologicalSurvey(WaterResourcesandBiologicalResourcesDivision).The prank in the foreground are cattails (Typha spp.) Photogroph by Karen A. Poiani

Biome and ecotone shifts

and 82 other. the cent per in Temperate forest area alters little. Tropical forests show a slight reduction Global warming fuelled by a doubling of carbon in area in all but one scenario, which predicts a slight dioxidelevelscouldproducelargeshiftsin the dis-increase. Maps of predicted change In the leafarea tribution of biomes. Onestudy,whichexplored five Index (which reflects themaximumrate of transdifferent scenarios forvegetationredistributionwithplration)implydrought-relatedbiomass lossesin a doubling of atmospheric carbon dioxide levels, pre- most forested regions, even In the tropics. The areas dicted large spatial shifts, especlally in extratropical most sensitive to drought-induced vegetation decline regions(Neilson1993a).Largespatialshifis In tem-areeasternNorthAmerlcaandeasternEuropeto western Russia. perate and boreal vegetatlon were predicted (Figure 7.14). Boreal biomes (talga and tundra) retreat north- In detail, spatial shifts of blomes should produce wards and decrease in slze by 62 per cent (the range discrete 'change' zones and 'no change' zones (Figure is 5 1 to 7 1 per cent, depending on the scenario used). 7.15) (Neilson 1993b). Such changes alter the pattern Boreal and temperate grasslandIncrease In area under of vegetatlon. Large, relatively uniform biomes are two scenarios. The increase IS 36 per cent one case reduced In m e , while new biomes, which comblne

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theoriginalbiomeandtheencroachingbiomes, emerge. Biomes and ecotones alike would probably affected by droughtfollowed by infestationsand fire if climate were to change rapidly. Ecotones would be especiallysensitive toclimaticchange,whether it wereslow or fast. The pattern of ecotonechange would be sensitiveto water stress (Figure7.16). With global warming unaccompanledby water stress, habltats should not fragment as under water stress, but should display a wave of high habitat variability as the ecotonegraduallytracks theclimatlcshifts. Under extreme drought stress, the entire landscape

shouldbecomefragmented as everyvariationin topography and soil become important to site waterbe balances and thesurvivorship of different organisms. The ecotone disappears for a while and reappears at a new location. It does not visibly shift geographically, as it would with no water stress, but disassembles and then re-establishes itself later. Forest change in eastern North America

It 1s probably common for species to movein the same general directlon. However, that does not mean

20 1

202

COMMUNITY CHANGE

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Fipre 7.15 Geographicalshift in blomes inducedby global warming. Discrete 'change' and 'no change' zones are produced.

Large, uniform biomes shrink. New biomes, which cornbme characterlstics of orlginal biomes and encroachlng biomes, emerge. Sorwce: After Neilson (1993b) thatentirecommunities move together.Quitethe contrary, species move at different rates in response to climatic change. The result is that communities disassemble, splittmg into their component species and losing some species that fail to move fast enough or cannot adapt. Mathematical models are helpful in understanding the possible changes in communities as the worldwarms up. Clearly, profoundchanges in the composition and geography of communities would occur. This is evidentinthe following case studies. An early model simulated forest growth at twentyone locations in eastern North America, as far west as a line joining Arkansas in the south to Baker Lake, North West Territories,in the north(Solomon 1986). All forests started growing on a clear plot and grew' undisturbed for 400 years under a modern climate. After the year 400, climate was changed to allow for awarmeratmosphere.Alinearchange of climate between the years 400 and 500 was assumed, the new climate at the year 500 corresponding to a doubling of atmospheric carbon dioxide levels. Climatic change continued to change linearlyafter the year 500, S O that, by the year 700, the new climate corresponded toaquadrupling of atmospheric carbon dioxide

levels. At the year 700,climatestabilizedandthe simulation went on for another 300 years. Simulation runsat each site were repeated tentimesandthe results averaged. Validation of the results was achieved by testing with independent forest composition data, as provided by pollen deposited over the last 10,000 years and during the last glacialstage, 16,000 years ago. Results for some of the sites are set out in Figure 7.17.Atthe tundra-forest border,thevegetation responds to climatic changein a relatively simple way (Figure 7.17a). With four times the carbon dioxide intheatmosphere,theclimatesupportsamuch increased biomass and,at least at Shefferville in Quebec, somebirches, balsam poplar, and aspens. The response of northern boreal forest to warming I S more complicated (Figure 7.17b).With a four-fold increase inatmospherlc carbon dioxide levels, theclimate promotesthe expansion of ashes, birches, northern oaks, maples, and other deciduoustrees at theexpense of birches, spruces, firs, balsam poplar, and aspens. In all northern boreal forest sites, the change In species composition is similar, but takesplace at different tlmes. This underscores the time-transgressive nature of vegetational response to climaticchange.The

C O M M U N I T YC H A N G E

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southern boreal forest and northern deciduous forest (Figures 7 . 1 7 ~andd) also have complicated a response toclimatlcchange.Inbothcommunities, species die back twice, the first time around and just after 500 years, and the second t m e from about 600 to 650 years. In the southernboreal forest, the change is from a forest dominated by conifers (spruces, firs, and pines) to a forest dominated initially by maples and basswoods, and later by northern oaks and hickories. At some sites in the northern deciduous forest, species appear as climate starts to change thenvanish once the stable climate associated with a quadrupled level of atmospheric carbon dioxide becomes established. Specles thatdothls Include thebutternut Vzqhluns c-znereu),black walnut (Jugluns nzgru), eastern hemlock (Tsrtgu cunrrdensrs), and several species o f northern oak. In western and eastern deciduous forests, the response of trees to climatic warming I S

remarkably uniform between sites (Figure 7.17e and 0. Biomass declineseverywhere,generally as soon as warming starts in the year 400. The drier sites in the west suffer the greatest losses of biomass, as well as a loss of species; this is to be expected as prairie vegetation would takeover. The declineof biomass in the eastern deciduous forest probably results chiefly from the increased moisture stress in soils associated withthe warmer climate.The increased moisture stress also gives a competitive edge to smaller and as the southern slower-growing species, such chlnklpin oak (Quereus muebhlenbergii),post oak (Quereus stellutu), and live oak (Quercus virginzutu), the black gum (Nyssu sylvutrca), sugarberry (Celtrs lumigata), and the American holly (Ihlex opacu), whlch come to domlnatetherapid-growlng species such as the American chestnut (Custuneu dentutu) and various northern oak species.

203

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Soil water regimes and forest change Forest growth is sensitive to changesinsoilwater regimes.Globalwarming is suretoalter water regimes and this will have an inevltable impact upon forests. A forest-growth model was used in conlunction with a soil model to assess the impact on forests of a doubling of atmospherlc carbon dioxide levels (Pastor and Post 1988).The modelwas run for several sites in the north-eastern United States, including a site in north-eastern Minnesota. The climate of this region is predicted to become warmer and drier. The major changes will occurat the boundarybetween the boreal forest and cool temperate forest. Forest growth was modelledon two soil types:soils withahigh water-holding capacity and soils with a low waterholding capacity. Thesimulations beganin 1751. Bare plots were 'sown' with seeds of the tree species common in the area. Themodel forest was then allowed to grow for 200 years under present climatlc conditlons. Next, carbon dioxide concentratlons were

Figure 7.17 Simulatlons of forestbiomassdynamlcs

gradually increased until, after century, a their present value had been doubled. After having reached that value, carbon dioxide levels were held constant for the next 200 years. Themodelpredictedthat on soils withahlgh water-holding capacity, where there will be enough water available to promote tree growth, productivlty and biomass will increase. O n soils with a low waterholding capacity, productivity and biomass will decrease in response todrierconditions.Inturn, raised productivity and biomass will up levels of soil nitrogen, whereas lowered productivity and blomass will down levels of soil nitrogen.Thevegetation changes are,therefore,self-relnforcing. O n waterretentive soils, global warming favours the expansion of northern hardwoods (maples, birches, basswoods) at the expense of conifers (spruces and firs); on welldrained, sandysoils, it favours the expansion of a stunted oak-pine forest, a relatively unproductive vegetatlon low in nitrogen, at the expense of spruce and fir.

over one millennium in response to climatic change induced by lncreaslng levels of carbon dioxlde In the atmosphere at SIX sltes In eastern North Amerlca. (a) Shefferville, Quebec (57"N, 67"W). (b) Kapuskaslng, Ontario(49"N, 83"W).(c) West upper Michigan (47"N, 88"W). (d) North central Wisconsin (45"N. 90'W). (e) South central Arkansas (34"N. 93"W). (0 Central Tennessee (36"N. 85"W). The tree specles are as follows: A. American beech (Fagrrs grundi/&a); R. American chestnut (Castanea denfafa);C . American holly (Ilex opacu); D. ashes: green ash (Fraxrnrrs penxrylc,un/ru), whlte ash (Fraxrnrrsutr,ertcatru), black ash (Fruxrtrusnrgra), blue ash (Fraxrnlrs (Tilia atuerrcanu) and whlte basswood (Tilia beteropbylla); I:. blrches: yrrudratrgrrlaru);E. basswoods:Amerlcanbasswood sweet birch (Betula letrtu), paper birch (Betula papyri/ira), yellow blrch (Bernla ullegbatrrensrs), and gray blrch (Betula poptrlijinlia); G . balsam poplar (Pop~lus ba/sut~~$eru). bigleaf aspen (Po/url/rs grundidetrtata),trembling aspen (Poprr1rr.rtremdozdes); 1 3 . black cherry (Prrrw.r .rero/nru);I . black g u m (Nyssa s y / r ~ / ~ c aJ. ) ; butternut (Jtrghns crtrereu) and black walnut ( J q l a n s nmgru);K . eastern hemlock (Tsfrga catrudetrrrs); L. elms: Amerlcan elm (U/?uwaruerrcanu) and wlnged elm (Ultmrs alutu); M. firs: balsam fir (Ab2e.r holsrrt~rea)and Fraser fir (Abiesfraseri);N . hlckories: bltternut hickory (Caryu c-ordiforttm).mockernut hickory (Carya tomntosa), plgnuc hlckory ( C a v a glabra), shagbark hickory ( C a v a n t ~ / r r ) shellbark , hickory (Carya lucrurosrr), andblackhickory (Carya t e x a m ) ; O. hornbeams:easternhornbeam (Ostrya zwgrmana) andAmericanhornbeam (Curprnrr.r cwo/inrn?ra);P. maples: sugar maple (Arer sacrburrrtn),red maple (Acer rubra),and silver maple ( A msaccharmum); Q. northern oaks: white oak (Qtrerurs all~u),scarlet oak (Qnerctrs c.occmnerr), chestnut oak (Qtrercusprrnus), northern red oak (Qrrrrc~rs rtdra), black oak (Quercx.r rrlrr/ma), bur oak (Qrrercrrs macrocurpa), gray oak (Qtren.rr.r borealis), and northern pln oak (Querc~rsellipso/dulis); R. northern whlte cedar (Tbtrp occtdenta/i.r), redcedar (Jlrnzpernsvrrgmzunu), and tamarack (Larmx larmtru); S. plnes: jack pme (Pitrzrs bnnkszrrna),red plne (Pinm resrwoso), shortleaf plne (Pintrs ec.brna/a),loblolly plne (Pinrrs iaedu), Virginla pine (Pinus t~rrgm~una), and p t c h plne (Pinus rmgtdu), and, 'I', white plne (Piwr.r s/roBtrs);IJ. yellow buckeye (Aewrlrrs ortundra); v spruces: black spruce (Pil-eu tmrrana), and red spruce (Picea r/rbens);and v l whlre spruce (Piceu glarrca); w. southern oaks: southern red oak (Qtrercxsfalcatrr), overcup oak (Qnere-nslyruta), blacklack oak (Qtrercmrsmarrlandia), chinkipln oak (QIMWS tmreblenbergii),Nuttall's oak (Qrrerc-usnuttallii), pin oak (Querempuhstrts), Shumard's red oak (Qtrwcxr shrrmardii,post oak (Qrwwr srellata), and live oak(Qmmws rwgmnzann);X. sugarberry (C'eltls lurt,rga/a);Y.sweetg u m (Lzqtrrdumbur sryracrfltra);Z. yellow poplar (Lrr/odendrot/ tdip$eru). Source: After Solomon (1986)

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COMMUNITY CHANGE Disturbance and forest dynamics

Global warming would favour a rise in the rate of forest disturbance owing to an increase in meteorological conditions likely to cause forest fires (drought, wind, and natural ignition sources),convective windsandthunderstorms, coastalflooding, and hurricanes. Simulationssuggestthatchanges in forest composition associated with global warming woulddependuponthedisturbanceregime(Overpeck et a/. 1990).Twosets of simulations were run.Inthe first set (‘step-function’experlments), simulated forest was grown from bare ground under present-day climate for 800 years. Thisenabled the natural variability of the simulated forest to be characterized. At year 800, a single climatlc variable was changed In a slngle step to a new mean value, which perturbed the forest. The simulation was then contmued for a further 400 years. In each perturbationexperiment,theprobability of acatastrophlc disturbance was changed from 0.00 to 0.01 at year 800. This is a realisticfrequency of aboutone plot-destroying fire every 115 years when a 20-year regeneration period(during whlch no further catastrophe takes place) of the trees in a plot is assumed. In each of the step-function simulations, three types of climaticchange(perturbation) were modelled: a 1OC increase in temperature; a 2°C increase in temperature; and a 15 per cent decrease in precipitation. Inthe second set of simulationruns(‘transient’ experiments), forest growth was, as in thestepfunction experiments, started from bare ground and allowed to run for 800 years under present climatic conditions.Then, from the years 800 to900,the mean climate,bothtemperatureandprecipitatlon, was changed linearly year by year tosmulatea twofold increase in the level of atmospheric carbon dioxide,untilthe year 1600 whenmean climate was again held constant. As In thestep-function experlments,theprobability of forest disturbance was changed from 0.00 to 0.01 at year 800. In all slmulationruns, a relatively drought-resistant soil was assumed,andthe resultswereaveragedfrom 40 random plots into a single time series for each model run.

Themodel was calibrated forselected sites in the mixed coniferous-hardwood forest of Wisconsin and the southern boreal forest of Quebec. Selected summary resultsare presented in Figure7.18.An increase in forest disturbance will probably create a climatically induced vegetation change that is equal to, or greaterthan,thesameclimaticallyinduced change of vegetation without forest disturbance. In many cases, thisenhancedchangeresulting from increased disturbance is created by rapid rises in the abundances of species associated with the early stages of forest successlon,which appear because of the Increased frequency of forest disturbance.Insome cases, as in Figures 7.17a, d, and e, a step-function change of climate by itself does not promote a significant change in forest blomass, but the same change workinghand-in-handwith Increased forest disturbance does have athoroughgoing effect on forest compositionand biomass. Interestingly, the altered regimes of forest disturbance, as well as causinga change in the composition of the forests, also boost the rate at which forests respond to climatic change. For instance, in the transient climate change experiments, where forest disturbance IS absent through the entire duration of the simulation period, vegetation change lags behind climatic change by about 50 to 100 years, andsimulatedvegetationtakesat least 200 to 250 years to attain a new equilibria1 state. In thesimulationruns where forest disturbance occurredfrom year 800 onward,thevegetatlonal change stays hard on the heels of climatic change and takes less than 180 years to reach a new equilibrium composition after the climatic perturbation at year 800. Ecological lessons from simulation models

Species show a wide range of responses to climate. In consequence, the response of different animal and plant species to climatic change will be varied. Thlsshould mean thatnaturalcommunities are likely to disassemble and that habitats restructure in transient, a non-equilibrium fashion as climatlc change unfolds. Validated models that help forecast theseeventsare needed to aid scientists in better

COMMUNITY CHANGE

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f 5

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c L

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.-m0

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Early successional trees

0 Hickories Maples

trees Other

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n hardwoods Other

F i p 7.18 ~ Simulated changes in species composition of forests at two sites investlgated in eastern North America. (a)

to (c) is a site in Wisconsin, and (d) to (0 is a site in southern Quebec. At both sites, experlments were run with an increase in disturbance at year 800 (top of figure for each slte) and without an increase In disturbance at year 800 (bottom figure for each site). Additlonally, three climatic change scenarios were slmulated: l0Catemperature Increase at year 800 (left-hand figures, a and d); a 15 per cent decrease In preclpltation (mlddle figures, b and e); and a translent change in which mean monthly preclpttatlon and temperature were changed linearly from year 800 to year 900 and thereafter held constant (nght-hand figures, c and f). Souwe: After Ovetpeck et al. (1990)

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COMMUNITYC H A N G E understandingthe e c o l o g d ramifications of global E S S A Y Q U E S T I O N S climatic change.Also, and perhaps more Important for conservationbiology,suchvalidatedmodels can help 1 Why hasClements‘sunidirectional view provide probabilities for the occurrence of these Of succession been revised? events, which will allowpolicy makers to make better, 2 Why does habitat fragmentation pose a informeddeclsions (T. L. RootandSchnelder 1093). threat to many species?

SUMMARY Communities change. The nature of this change I S debatable. Theclasslc view saw climatic climaxes and balanced ecosystems resulting from unldirectional successlon. Succession 1s a complex process and IS explained by at least three models - the facilitatlon model,the tolerance model, and the inhibltion model. It may also be drlven by fiactors external to the community (allogenic factors). Prlmary succession is the colonization of land or submarine surfaces that have never existed before. Secondary successlon is the Invasion of newly created surfaces resulting from removal of pre-existmg vegetation. A modern view of communitychange stresses thedisequilibrium behavlour of communitles. It sees succession going inmanypossible directions,and sees communities as temporary collections of specles that assemble and disassemble as the envlronment changes. Much communltychange has been caused by land cover transformation over the past two hundred years. Two Important aspects of this transformation are habitat fragmentation,and I t attendant effects onwildlife, andthe loss of wetlands.Globalwarmingduring the twenty-first century is likely to put peripheral, geographicallyrestrlcted,highly specialized, poorly disperslve, andclimatically sensitivespecies under intense pressure. It will also cause significant changes in many communitles. Wetlands, tundra, and alpine meadows are especially vulnerable.

3 What communitychanes are likely to occur as a result of gloI 2al warming?

F U R T H E RR E A D I N G Committee on Characterization of Wetlands, National Research Council (1995) Wetlands: Charactertstics and Boundaries, Washington, DC: National Academy Press. Gates, D. M. (1993) Climate Change and Its Biological Conseqwences, Sunderland, Massachusetts: Sinauer Associates. Huggett, R . J. (1993) Nodefling the Httnzan Impact on Nutwe: Systenu Analysis of Envtronnmtal Problems, Oxford: Oxford University Press. Huggett, R. J. (1997) EnvironmentalChange: EvofLhg Ecosphere, London: Routledge.

The

Jackson, A. R.W.andJackson,J.M. (1996) Environnrental Scrence: The Natltral Envtronment and Ht/n~anImpact, Harlow: Longman. Matthews, J. A. (1992) The Ecology of RerentlyDqlaciated Terrain: A Geoecologtcal Approarb t o Glacw Forelandr and Primary Succession, Cambridge: Cambridge Universlty Press. Peters, R. L. and Lovejoy, T. E. (eds) (1992) Global Warnmg and Bzological Dtversity, New Haven, Connecticut and London: Yale Unlversity Press. Schwarrz, M. (1997) Consewation In Highly Fragmented Landrtapes, New York: Chapman & Hall.

LIFE, H U M A N S ,A N DM O R A L I T Y Human attitudes toulards the living world are relevant to biogeographers involved with conservation a n d ecosystem management. This chapter covers: non-humanrights kinds of environmentalism biogeographicaltheoryandconservationpractice

Biogeographical and ecological prlnciplesnaturally Inform questions of envlronmental management. However, envlronmental management does not just have a ‘sclentific’ dimenslon. It has social and ethical dimensions, too. All dimensions must be consldered in discussing the relations between humans and other livlngthings.Thischapter will explore therlghts of animals and Nature, attitudes towards Nature, as seen in the different brands of envlronmentalism, and therelationships between biogeography, ecology, and conservation practlce.

BlORlGHTS Environmental ethics deals wlth value judgements aboutthe biologlcal and physicalworlds. I t was begun, at least in Its modern form, by philosophers during the 1970s. Environmental ethics is a diverse andimmense field of study. I t arose from three conslderatlons (Botkin and Keller 1995: 619). First, human actlons, aided by technologlcal developments, affect Nature deeply, so it seems necessary to examine the ethical consequences of these actions. Second, many human actions have world-wde consequences, and a global perspective ralses Its own moral questions. Third, moralconcerns have expanded so that

animals, and even trees and landscapes, possess moral and legal rights - civilization includes all Nature wlthln its ethicalsystems. This expansion of concerns demands a new environmental ethics.

Animal wrongs: do organisms have rights?

What are rights? Rights are a social contrivance that help people to live together. They are not some quality that indivrduals possess. Rather, they are a quality that members of a socletyare willing to pretend that individuals possess. Such apretence assures socially acceptable behaviour (most of the time). This idea is straightforward, though open to philosophical debate. The big questlon is whether the scope of ‘rights’ shouldbe extendedtoincludeother species, and even rocks, rlvers, and landscapes. There I S no convincmg reason why they should not be so extended. If rlghts should help humans to avoid conflict with one another, why should not animal rights, plant rlghts, and landscape rights help humans to protect thelr envlronment? An all-encompasslngdefinitlon of rlghts would helphumansto respect Nature.Butthematter is complicated by three preconditions, without whlch

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LIFE, HUMANS, AND MORALITY an objectcannot be grantedrights - commensurability, responsibility, and sensibility (Tudge 1997). Commensurability means that rightsare a reciprocal agreement - you respect my rights and I’ll respect yours. Unfortunately, this equitable arrangement is distorted by power structures. Some groups of people have less power to infringe the rights of others. More powerful groups (the richer for instance),therefore, tend to possess more rights than less powerful groups (the poorer for instance). As animals, plants, and landscapes have no effective power, why should they be grantedrights?Stated so bluntly,thlsargument smacks of insensitlvity. To sootheconsclences, a rider is added - to earn rights, people must accept responsibility. Animals, plants, and landscapes cannot accept responsibility, therefore they can have no rights. However, there IS no reason why rights and responsibilitles should be Inseparable. If animals are worthy of human respect, then they can be awarded rights, even though they should have nofeelings toward humans.Sensibility, or lack of It, inall species save humans, arose from a mechanlstic v ~ e wof animal behaviour instigated by Ivan Pavlov and hls dogs. To be brief,most scientists are now prepared to see otheranlmals as somethmgmorethanblochemicalmachlnes. I t wouldseem that ‘respectable science has caught up wlth theanlmal-lovers’ and I t IS no longer perverse ‘to perceive animals as sentient beings, who register their emotional responses with varylngdegrees of consciousness’ (Tudge 1997: 41; see also Regan 1983). So, if animals have rights, how should they be treated? This is a difficult Issue with many aspects. All the aspects stem from a sea change in human attitudes toward animals, at the root of which is a new-found sympathy towards the natural world, and towards anlmals in particular (e.g. Fisher 1987). This sympathy has drlven the forces of animal welfare over recent years. Theanimalliberationmovement, through films andothermedia, has shownpeople how many anlmals are forced to live in pounds, on factory farms, and In laboratories.People have been stirred into various klnds of actlon - boycotting pet shops, breakinginto laboratories to free animals, ceaslng to eat meat, spurnlng the wildlife trade, and

so on.However,thechange in humanattitudes toward animals does not express itself solely in militant action. There is a qulet slde to the change that is less evident but possibly more fundamental. Two examples from AustraliaandNew Zealand (O’Brlen 1990) should demonstrate this subtle and yet profound attitude change. In Australia, the VictorianAcclirnatlzation Society was formed in 1861. A central objective of this early wildlife management society was ‘the introductlon, acclimatizarlonanddomesticatlon of allinnoxious animals, birds, fishes, Insects and vegetables whether objective useful or ornamental’(Rolls1969).This was regarded as worthy and servlng the public interest by sclentists, natural historians, and landholders of the Soclety. InmodernAustralia,theacclimatization of exotic specles would be regarded by many as Irresponsible and bizarre, and is prevented by legislatlon. Attitudes towards ’pests’, some of which are the ancestors of the ‘innoxious’ animalsintroduced in the nineteenth century, have also shifted. Take the example of the rabbit. The huge Australian rabbltpopulation was controlled in the1950s by introduclng myxomatosls. But In 1987, when the same vlrus was consldered as a control agent for the wild rabbit population I n New Zealand, the welfare of wild rabblts was put first and the virus was not introduced. Three moral issues arising from thenotion of animal rlghts deserve closer scrutiny - culling animals, the wildlife trade, and keepmg animals in captivity.

Culling Controllinganimalpopulatlons may mean culling. Few people raise objectionstothe eradication of viruses, bacteria, or pestilential insects. Attempts to limit populations of organisms higher up the evolutionary ladder cause eyebrows to be raised. Mammal culling causes public outcry. The cullingof seal pups IS met by a vociferous and appalled response. Theshooting of feral horses from helicopters began In Australia in late1985. By 1988,the CommonwealthDepartment of PrimaryIndustries

LIFE, H U M A N SA, N D MORALITY was receiving more than 7,000 protest letters a year, and to ban trade in a further 800 specles threatened mostly ‘campaign’ mail from correspondents overseas with extinction. About a quarter of the wildlife trade is unlawful (O’Brien 1990). In the United States, feral horses are protected by law. The Wild Free-Roaming Horse and commerce in rare andendangered species. The unBurro Act (197 1)decrees that feral horses and burros fortunate animals are poached in the wild and smugmust be consldered as an integral part of the natural gled across frontiers. A typlcal year will see the sale systems in which they are found. The Actprovides for of 50,000 primates, tusk ivory from 70,000 African the Bureau of Land Management to set suitable levels elephants, 4 million live birds,10millionreptile for herd size, and for surplus animals to be removed skins, 15 million pelts, 350 million troplcal fish, and and offered for privatemamtenance by qualified about 1 million orchids (Lean and Hinrichsen 1992: individuals, or destroyed if they are old, sick, lame,or 145).Peoplecaught illegallytrafficking In wildlife private care cannot be found. In 1987, 8,000 horses are arrested, fined, and even jailed. Thegrowing willingnessanddeterminationtostopthewildlife were held in captivity awaitlng ‘adoption’. trade is anotherinstance of humans upholding the Many people accept thatsomehumaneculling rights of anmals. of largemammals is a necessary evil toprevent over-exploitation of vegetatlonandenvtronmental degradation. In Great Britain, for example, the Keeping animals in captivity culling of deer populations is probably necessary. are notwhat theywere. As With animal rights now to the fore, the method of Zoologicalgardens society’s attltudes toward animals have changed, zoos culling I S the subject of scientific study. On Exmoor and the Quantock Hills, In Somerset, red deer (CKWLS have responded by rethinkingthe reasons for their in the heyday of natural K ~ U ~ ~ are U J culled ) by rifle (stalking) and by hunting existence. Zoos began history, when almost every educatedVictorian was with hounds. A study carried out for the National an amateur natural hlstorlan. Interests atthetime Trust(the Bateson Report)concludedthathunts with hounds cause deer great distress. The exerttons centred around collectmg, identifying, and naming. of the chase change muscles and blood far beyond Zoos were places to display the ordinary and bizarre anychanges that would be expected In normal life. creatures found in far-flung parts of the globe. This Red deeraresedentary animals that escape natural role for zoos lingered well into the twentieth century. Even in 1970, the lion enclosure at Newquay Zoo, in predators (mainly the wolf, although there are none now in Britaln) by brlef sprints. Hunting deer with Cornwall,England, was designedto keeplionson hounds is not ‘natural’, because wolves would never display to the public for as long as possible. This was chase deer for longdistances. As a result of the achieved by keeping them in a relatively small space, even smaller sleepBateson Report, membersof the National Trustvoted and letting them return to their ing quarters only at nlght. In the 1990s at the zoo a to ban deer hunting with hounds on all its lands. largelionenclosureincorporatesmanyfeatures that resemble the lions’ natural savannah environment. The wildlife trade There is, for instance, a platform (simulating a small The wildlife trade is an international business knoll) and trees. Food is hidden around the enclosure worth about $15 billion a year. Much of it I S legal and the lions have to find It. And the lions are free to ‘go indoors’ when they wlsh. and controlled by national laws and an international A brochure for Newquay Zoo captures the splrlt treaty - CITES (Convention on International Trade inEndangered Specles of Wild Fauna and Flora). of the tlmes when i t says that good zoos are good CITES was drawn up in 1975 and 139 countries are for anlmals and good for people. They are good for now party to it. Collectlvely,these countrles act to anmals for one reason only - conservation. Five regulate and monltor trade In around 25,000 spec~es aspects of conservation are paramount:

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LIFE, H U M A N S , A N D M O R A L I T Y Breeding 1 programmes save animals from extinctlon. 2 Wildlife research helps to preserve habitatsand save animals. 3 Animal behaviour research assists their survival in the wild. 4 Providingsanctuaries for animalsthat have lost their habitat. 5 Provlding hospitals for sick and injured animals.

Does all Nature have rights?

A belated effect of the Darwinian revolution is that humansare now seen as part of Nature.Environmental philosophers now passionately prosecute this idea, but remain unclear as to its implications. What does it mean to be part of Nature? This philosophical issue is complexand beset by aterminological labyrinth (see Colwell 1987). But all commentators agree that ‘being part of Nature‘ dispenses with the Unless wild populations are supplemented by duality between humans and the natural world, provldes a moral justification for treating Nature more captive breeding, and culling populations that outhumanely, and gwes a philosophical ~ustification for grow their reserves, then most of the world’s largest seeing intrinsic valueinall things(e.g.Callicott terrestrlal mammals will vanish. Zoos make signifi1985; McDaniel 1986; Zimmerman 1988). cant contributions to conservatlon as genetic refuges An early and eloquent advocate of theintrinsic and reservoirs, especially for large vertebrate species worth of the natural world was Aldo Leopold (1049), threatened with extinction (Rabb 1994).Przewalski’s who advanced an all-embracing land ethic. Leopold J are believed to be extinct horses ( E ~ u przeutafskii) affirmed the right of all resources, including plants, In thewild;thecurrentknownpopulation of 797 animals, and earth materials, to continued existence, animals existsin zoos (Boyd 1991).It is hoped to and, at least In places, tocontinued existencein a remtroducethis species toits former Mongolian natural state. This land ethic assumes that humans habitat by the early years of the twenty-first century. are ethically responsibility tootherhumansand Zoo resources for conservation and research are human societies, and to the wider envlronment that limited, so zoos are encouraglngthedevelopment includes, animals, plants, landscapes, seascapes, and of criteria to help prioritize actions for conservatlon the air. Granting rlghts of survlval to animals does of biodiversity.NorthAmerican,European,and Australian zoos are assisting the developmentof tech- not necessarily mean that they cannot be eaten, but it does mean that endangered spectes should be cared nical capacltles among zoo counterparts, government for and helped to recuperate. A land ethic behoves agencies, andprotected areas in bothdevelopmg and developed countries of the world to further the humans to sustain Nature for the present generation and for future generatlons. conservation of biodiversity. The question of Nature’s rights arose as a legal A remarkablesuccess story (so far) I S the goldenlion issue in the 1970s. Mineral King Valley is a wilderZ ~ J (p.197).Thlssquirreltamarin ( L ~ O W Z ~ ~roJalia) ness area in the Sierra Nevada, California.Disney sized primate lives in themuch-reduced lowland Enterprises, Inc., wlshed to develop the valley as a Atlantlc forest, Brazil. Since the mid-l99Os, a comski resort withln multimilliondollar recreational binatlon of relocating five tamarin familiesfrom facilities. The Sierra Club (founded in 1892 to help vulnerable areas Intotheprotectlon of the POGO preserve the Yosemite Valley and other natural das Antas Biological Reserve, and relntroduclng 147 captive-bred tamarins into the wild, has put the wonders In California) objected that the development value and ecologlcal little prlmate on the road to recovery. For long-term would destroytheaesthetlc balance of the area, and brought a suit against the survival, the tamarm’s habitat area will need to be government. The Sierra Club could not claim direct doubled. harm from thedevelopment,andthe land was government owned and the government represented the people, so peopleingeneral were not wronged.

LIFE, H U M A N S , A N D M O R A L I T Y A lawyer, ChristopherD.Stone (1972), suggested that, as there were precedents for Inanimate objects such as shlps havlng legal standing, trees should also have legal standing. So the sult was made on behalf of thenon-human wilderness. The case was taken tothe US SupremeCourtbut was turneddown. However, in a famous dissenting statement, Justice William 0. Douglas proposed the establishment of a new federal rulethatwould allow ‘environmental issues t o be litlgated before federal agencies or federal courts in thename of theInanimateobjectabout to be despoiled, defaced, or invaded by roads and bulldozers and where injury is the subject of public outrage’. Although trees were not given legal rights in thls case, theethlcal values and legal rights for wilderness were discussed.

brands tend to polarize around, at one extreme, ecocentrism, with Its decidedly motherly attitude towards the planet; and, at the other extreme, technocentrism with a more luissez-farre, exploitative attitude (Table8.1). The ‘green’ endof this spectrum was Initiated by theatomlcbombandgalvanued into action by pesticide abuse. Each strand represents a system of beliefs, although no individual would necessarily believe in just one strand. Technocentrics (pale greens and very pale greens)

Technocentrics tend to have a human-centred vlew of the Earth coupled with a managerial approach to resource development and environmental protection. They favour maintaining the status quo in existing governmentstructures.Technocentricstendto be conservatlve politlclans of all political partles, leaders A T T I T U D E ST O W A R D S N A T U R E of industry, commerce andtrades’ unlons, and skilled The Age of Ecology opened on 16 J d y 1945 In the workers. In short, they are members of the producNew Mexican desert ‘wlth a dazzling fireball of light tive classes. They aspire to improve wealth for themand a swelling mushroom cloud of radioactive gases’ selves and for society at large. They en~oymaterlal (Worster 1994: 342). Observmg the scene, a phrase acquisition for own sake and for the status I t endows. politically and economlcally very from the Bbagazlarl-Gita came Intoproject leader And they are J. Robert Oppenhiemer’s mind: ‘I am become Death, powerful. There are two brandsof technocentrlc - the cornuthe shatterer of worlds’. For the first time in human history, a weapon of truly awesome power existed, a copians (or optimists) and the accommodationists weapon capable of destroying life on a planetary scale. (or environmental managers).Conservative technoare From under the chillingblack cloud of the atomic age centrics are cornucopians or optmlsts.They arose a new moral concern - envlronmentalism - that named after the legendary goat’s horn thatoverflowed with fruit, flowers, and corn, and signified prosperity. soughttotemperthemodern science-basedpower over Nature with ecological inslghts into the radia- They have faith in the applicatlon of sclence, market tion threat to the planet. The publication of Rachel forces, and managerial ingenuitytosustalngrowth and survlval. Carson’s Silenr S p r r q (1962), the first study to bring Liberal technocentrics are accommodationists. the lnsldious effects of DDT applicationtopublic They have falth in the adaptability of institutions and notice, was more grist for theenvlronmentalist’s mill. The dual threats of radiation and pesticldes set mechanisms of assessment and decision makingto accommodate envlronmental demands. They believe environmentalism moving. that adjustments by those In positions of power and significance can meet theenvlronmentalchallenge. Brands of environmentalism Theadjustments involve adaptlngtoandmouldenvironmental tmpxt There are several brands of environmentalism, each ing regulation (including assessment) and modifylng managerial and busmess of whlch promotes a different attitude towards the envlronment (see O’Rlordan 1988, 1996). These practices to reduce resource wastage and economlcally

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LIFE, H U M A N SA,N M D ORALITY Table 8. I

Brands of environmentalism and their characteristics

Environmental inclination

Type and colour

Characteristics

Ecocentrism (Naturecentred)

Deep ecologists or

Stress the intrinsic importance of Nature for humanity Believe that ecological (and other) laws should dictate human morality Ardently promote biorights - the right of endangered species, communities, and landscapes to remain unmolested

Gaians (dark greens)

Communalists or self-reliance soh-technologists (medium greens)

Technocentrism Accommodationists (humancentred) or environmental can continue managers (pale greens)

Emphasizethesmallness of scale ('small is beautiful') and hence community identity in settlement, work, and leisure Interpret concepts of workand leisure through a process of personal and communal improvement Stress the importance of participation in community affairs, and guarantee of the rights of minority interests Believe that economic growthand resource exploitation assuming: suitable economic adjustments to taxes, fees, etc.; improvements in the legal rights to a minimum level of environmental quality; compensation arrangements satisfactory to those who experience adverse environmental and social effects Accept new project appraisal techniques and decision reviews arrangements to allow for wider discussion or genuine search for consensus among representative groups of interested parties

Cornucopiansor Believe that humans canalwaysfind a way out of any optimists (very pale difficulty, beit political, scientific, ortechnological greens - dark blueAccept that prc-growth goalsdefine the rationalityof under the surface) project appraisal and policy formulation Optimistic about the human ability to improve the lot of the world's people Faith that scientific and technological expertise provides a firm foundation for advice on matters pertaining to economic growth, public health, and public safety Suspicious of attempts to widen basis for participation and lengthy discussion in project appraisal and policy review Believe that all impediments can be overcome given a will, ingenuity, and sufficient resources arising out of growth

LIFE, HUMANS, A N D MORALITY inconvenientpollutlon,without any fundamental shift in the distribution of political power. Pressure groups aligned to technocentrics tend to be NIMBY (not-in-my-back-yard)groups - theydonotmlnd development so long as ~t does nottake place near them - and private Interest groups (consumer groups, civil liberties groups, and so on). They are very vocal and command much media attentlon.

the essential co-evolution of humans and natural phenomena. Extreme ecocentrics believe that the moral basis for economic development must lie in the interconnections between natural and social rights: there is no purely anthropocentric ethic. Within thls group might be Included a powerfulreligiouslobby (Box 8.2).

Alignments of ecologists Ecocentrics (medium greens and dark greens)

Ecocentrics have a Nature-centred vlew of the Earth in which social relations cannot be divorced from people-mvironmentrelations.They hold a radical vlslon of how future soclety should be organized. They would give far more real power to communlties andto confederations of regionalInterests. Central control and natlonal hegemony, so treasured by technocentrlcs, are the very antithesls of ecocentrism. Ecocentrics tend to be found among those on the frlnge of modern economles, that is, the‘non-productlve’ classes - clerics, artists, teachers, students, and, to an ever increasing extent, women. Theyare characterized by willingness a to eschew materlal possessions for their own sake, and to concern themselves more with relationships; andby a lack of faith in large-scale modern technology and assoclated demands on elitlst expertise and central state authority. Pressure groups aligned to ecocentrics tend to be green groups and alternative subcultures (e.g. femlnists, the peace movement, religious movements). There are twobrands of ecocentrics. The more conservative ecocentrlcsare communalists or selfreliance, soft technologists. They believe in the cooperativecapabilities of societies to be collectlvely self-reliant using approprlate science and technology. That I S , they believe In theability of peopleto organlze theirown economles if glventherlght incentlves and freedoms. Thls vlew extends a ‘bottom LIP’ approach to the development of the Third World based on the application of indigenous customs and appropriate technical and economicassistance from western donors. Liberal ecocentrics are Gaians or deep ecologists (Box 8.1). They believe in the rights of Nature and

I t would be wrong, if excusable, to Imagine that all ecologists andenvironmentalscientists hold ecocentric views. Therelatlonship betweenecocentric environmentalists and ecologists has not always been as cosy as might be supposed: ecologists do not always say what ecocentrlcs wantto hear. Paul Colinvaux (1980: 105) wrote: ‘If the planners really get hold of us so that they can stamp out all individual liberty and do what they like wlth our land, they might declde that whole countles full of Inferior farms should be put back into forest’. His displeasure IS wlth land-use planningandenvironmentalism evident. Later In the book (Colinvaux 1980: 1lo), hls words smack of social Darwinism and ‘Nature red in tooth and claw’, when he talks of different specles ‘going about earnlng thelr livlngs as best they may, each In its own indivldual manner’, and what ‘look like community properties’ being ‘the summed results of all these blts of private enterprise’, though elsewhere he sees peaceful coexistence, not struggle, as the outcome of natural selection, the peace belng broken In upon by a devlant aggressor species such as Hotno .rappiens who seek to encroach on another’s niche (Colinvaux 1980: 131). Today, few ecologists wholeheartedlyacceptColinvaux’s viewpolnt,and many approach Naturegreen In head andheart.One of the many I S Danlel B. Botkin (1990) who advocated uslngmodern technology In a constructwe .and positive manner, a position that tries to brldge the mlddleground betweenecocentrlsm andtechnocentrlsm.

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2 17

LIFE, HUMANS,ANDMORALITY -

~~~~

Table 8.3 Different attitudes of deep and shallow ecologists Deep ecology

Environmental Shallow ecology problem Advocate a technology that seeks to purify the air and water and to spread pollution more evenly; laws limit possible pollution; polluting industries are preferably exported to developing countries

Evaluate pollution from an ecospheric point of view, not focusing on its effects on human health, but on the effects on life as a whole, including life conditions of every species and ecosystem

Resources

Emphasize resources for human beings, especially for the present generation in affluent societies. Resources belong to those who have the technology to exploit them. Resources will not be depleted because, as they get rarer, higher market prices conserve them, and substitutes will be found through technological progress. Plants, animals, and natural objects are valuable only as resources for humans; if they should have no use to humans, they can be destroyed

Deeplyconcernedwith resources and habitats for all life-forms for their own sake; no natural object is regarded solely as a resource; this view leads to a critical examination of human modes of production and consumption

Population

See overpopulation as a problem of developing countries. The question of the optimum human population is discussed without reference to the optimum population of other life forms. The destruction of wild habitats is accepted as a necessary evil

Recognize that excessive pressures on planetary life stem from the human population explosion. The pressure stemming from industrial societies is a major factor, and population reduction in these societies, as well as in the developing countries, is imperative

Cultural diversity and appropriate technology

Regard industrialization of the kind manifested in the west as an example for developing nations

Concerned with maintaining cultural diversity, and limiting the impact of western society upon presently existing non-industrial societies, defending the fourth world against foreign domination, and applying local, soft technologies where appropriate

Pollution

Source: After Naess (1 986)

1 ~

2 18

LIFE, HUMANS, AND MORALITY Box 8.2 RELIGION, NATURE, AND WORLD WILDLIFE FUND

THE

Inautumn 1986, auniquealliance wasforged between conservation, as represented by the World Wildlife Fund, and five of the world’s chief religions. The venue was Assisi, in central Italy, home of thepatronsaint of animals.Apilgrimage, conference, cultural festival, retreat, and interfaith ceremony in the Basilicz of St Francis formed the basis of a permanentbondbetweenconservation and religion. Forreligions,ecologicalproblemsposemajor challengesinrelatingtheologyandbelieftothe damage to Nature and human suffering caused by environmental degradation. All too ofien, religious teaching hasbeenused or abusedtojustifythe destruction of Natureandnatural resources. In the declarations issued at Assisi, leading religious figuresmappedoutthe way to re-examine and reverse this process. Andin 1987, the Baha’i’ Movement, which had its origins in Islam, joined the World Wildlife Fund’s New Alliance. At the same time, starting with the conference and retreat heldin Assisi,religiousphilosophersarehelping inject some powerful moral perspectives into conservation’s ill-defined ethical foundations. Adevelopment of the Assisieventwas the making of the Rainbow Covenant at Winchester Cathedral,EnglandinOctober 1987. Covenants have long a history. For theancientHittites, covenants were made between rulers and subjects for the benefit of both. For Jews and Christians, ‘The Covenant’ is between God and his people, and was made after the Flood. In the Bible, Noah, at God’s

A balancing act: biorights, human

command, builds an ark to save species onEarth from extinction when God sends a grand Flood to punish humanity. After the Flood has finished, God makes a covenant - t h e first in the Bible. He makes just withNoah,butwithall hiscovenantnot creatures on Earth, in the seas, and in the skies. God promises never wantonly to destroy life again, and sets the rainbow in the sky to bear witness to his promise. TheRainbowCovenant has beeninspired by God’s covenantwithNoah,and was specifically designed by the International Consultancy on Religion, Education and Culture (ICOREC) to be usablebyanyone,religious or not. So, unlike the Hittite and biblical covenants, it is unilateral a contrite pledge by humans to other living beings and their ecosystems. The rainbow’s arc embraces not only those who live, but those who will live; it encompasses all living creatures and the earth, air. and water that sustain them. This is the Rainbow Covenant: Brothers and Sisters in Creation, we cwenant this aky wid you and with all mation yet to be; With awy living mature and all that contains andsustain. you. With allthat is on Earth and within the Earth itself;. Withall thatlives in thewaters and with the water themselves; With all that pies inthe skies and with the sky itseq We establish this covenant, that all our PWS will be US^ to prevent your destntction. We confus that it is our own kind who p:tt yon at risk 4 dpath. We ask fw your t m t and as a symbol of our intention i ~ ’ mark this cmwant with you by the rainbow. This is the sign of the covenant betum ourselm and living thing that is found on the Earth.

Ecosystem management arose in the late 1980s and is advocated by many scientists and other people Do humans still have rights in the world of the deep interested In the environment (e.g. Agee and Johnson ecologist? People are partof Nature. The philosophy 1988). Its ultimate aim is to enhance and to ensure behind a new brand of conservation - ecosystem the diverslty of species,communities,ecosystems, and landscapes. It is a fresh and emergmg model of management - accepts that simple fact.

rights, and ecosystem management

LIFE, H U M A N S , A N D M O R A L I T Y resource management. It covers spectrum a of approaches. Amild,technocentric version simply extends multiple use, sustained yieldpolicies,prosecuting stewardship a approach, in which the ecosystem I S seen merely as ahuman life-support system (e.g. Kessler et al. 1992). In this v~ew, public demands for habitat protection, recreatlon, and wildlife uses are simply seen as constraints to maximizing resource output (Cortner andMoote 1994). A more radical, ecocentric approach is to accept Nature on itsownterms, even where doing so means controlling ~ncompatible human uses (e.g. Keiter and Boyce 1991). This extreme, but eminently sensible, form of ecosystem management reflects a willingness to place environmental values, suchas blodiversity and animal rights, and social and cultural values, such as the upholding of human rights, on an equal footing. It means accepting that environmental concerns are not purelyfactual; they have do withvalues and ethics. Peopleexert a profound influence on ecological patterns and processes, and in turn ecological patterns and processes affect people. The connection between technology and the environment is well studied; not so the connectionbetween social system and the environment. However,policies are tending to move away from the administrator-as-neutral-expert approach to policies that engender public deliberation and the discovery of shared values. Naturally, such extension of ecolog~cal matters to the soc~al and political arena presents difficulties, though these may not be insuperable (e.g. Irland 1994). Ecosystem management accepts that human values must play a leading role in policy decision-making. Conservation strategies must takeaccount of human needs andaspirations;and they must integrate ecosystem, economic, and social needs. The key players in ecosystem management are scientists, policy makers, managers, and the public. The public, many of whom have a keen interest ~n environmental matters, are becoming more involved in ecosystem management as professlonals recognize the legitimacy of claims that various groups make on natural resources. In Jervis Bay, Australia,the marine ecosystem is used by many existingand proposed conflicting interests (national park, tourlsm, urbanization,militarytraining)(Wardand Jacoby

1992). Similarly, in the forests of the south-western United States, ecosystem, economic, and social needs are considered in policy decision-making concerning ecology-based, multiple-use forest management (Kaufmann et a / . 1994) (Figure8.1).

B I O G E O G R A P H Y ,E C O L O G Y ,A N D C O N S E R V A T I O NP R A C T I C E Biogeographical and ecological ideas inform conservation practice. When ruling theories ~n ecology and biogeography change, conservation practice soon follows sult. This generalization, though valid, shouldbetreatedcautiously, as therelationships between ecological science and environmental policy are complicated (Shrader-Frechette and McCoy 1994). Nevertheless, it is fair to say that the major paradigm shifts in twentieth-century ecological thought have engendered quite distinct conservation philosophles. Four paradigms, which were mentioned in Chapter 7, are recognized: endurlng equilibrium, balancedecosystems, disequilibriumcommunities, and the edge of chaos.

Environmental exploitation and enduring equilibrium Environmental problems in the offing

In his Walden;or, Liji in the Woods (1893 edn), Henry Davld Thoreau expressed concern for wild Nature and wild environments. H e urged that, when a conflict arises between Nature and human society, then the first thing to consider are the laws of Nature. George Perkins Marsh wrote ManandNature;or,Physical GeographyasModified by Human Action in 1864. H e examined‘thegreater,morepermanent,andmore comprehensive mutations which man has produced, and I S producing in earth, sea, and sky; sometimes, indeed,with conscious purpose,but for themost part, as unforeseen thoughnatural consequences of actsperformed for narrower and more immediate ends’ (Marsh 1965 edn: 19). He believed that ‘Man is everywhere a disturb~ng agent. Wherever he plants

219

220

LIFE, HUMANS, AND MORALITY

Social needs I

ecosystem and human benefits, costs, and risks (National Environmental ’ Policy Act)

Economic and social needs

I I

+

+

+

Ecological principles

Species

4

I

,

Biological integrity

loss

Implementation

+

Ecological principlesfilter

unrest Ecological, economic,

Economic socialDegraded and environments sustainability instability

Monitoring

and feedback

Figure 8.1 T h e Integration of ecologlcal, economtc, and social needs In a decwon-analysis model. Economlc and social needs are tested agalnst an ‘ecologlcal filter’, whlchIS shown In the shaded box. The aim IS to determine economlc and

social actlons that will produce the most deslrable balance between blologlcal Integrity and ecological, economic, and soclal sustainability Bowlng fully to economlc and soclal needs would lead to specles loss and envlronmental degradation. Bowlng fullyto ecological needs would leadto soclal unrest and economlc Instability A compromlse posltlon allows the malntenance of blological Integrity while catermg for economic and soclal needs. The resulting decision model leads to the implementation of an envlronmental policy.The effects ofthe policy are carefully monitored and evaluated. Ifthe policy should fail to work as desired, then amendments can be made and the process started anew, until a satlsfactory outcome IS achieved. Source: After Kaufmann et al. (1994)

LIFE, H U M A N S , A N D M O R A L I T Y hisfoot, theharmonies of natureareturnedto discords’(Marsh 1965 edn: 36). Marsh was the first modern scholar to see humans as a factor, and not merely as a subject, in Nature. In1922,Robert Lionel Sherlock published an enormously important book entitled Man as a Geological Agent: A n Aa-otmt o f His Artton o n lnanitnute Nattrre. From extensive observatmns withtn the British Isles, Sherlock showed the effects that human activities were havingon the landscape, the effects that forestry, grazlng, agricultural management, and the technologlcal and d u s t r i a l activity of twentlethcentury society were havingonlandscape processes. Specifically, he looked at the effects of road and railway construction, open-cast coal minlng, and slate, stone, gravel, and sand quarrying; and the management of waterways and coasts. He gathered starlstlcs glving the amounts of material extracted or removed, and furnished estimates of the rates of erosion and sedimentatlon. During the late 1920s and the 19.30s. the recognition of the human Impact on the soil and the need for soilconservatlon was a pressingconcern,largely because of the dust-bowl that devastated the Great Plains reglon In the Unlted States. The main problems of environmental degradation were found m a d y in western Europe and North Amerlca.

land use. Thispractlce, heaverred, shouldstop environmental degradation andplace the entire nation on a biologically and economically sustainable footing.Inother words, hewas proposing ecological princlples to gulde conservation practlce and achleve a relationship with Nature that ensured an enduring equilibrium.

Balanced ecosystems

The balanced ecosystem view, and its extension in the Gaia hypothesis, supplanted ‘enduring equilibrium’ as a blueprintfor Nature’s design.I t guided conservation strategy in two different ways. First, ecocentric environmentalistsarguedthatunrestratned interference withNature’sdevelopmentstrategy leads to ecosystem damage,andthatthe world’s endangered ecosystems should be defended against the excesses of human actlons. ‘Spaceship Earth’, a term first used by Adlai Stevenson, a Unlted StatesAmbassador, in a speech given before the United NatlonsEconomic and Social Council in Geneva on 9 July 1965, became a leading environmental idea. Theimage of Spaceship Earth has now lost Its potency. Nonetheless, i t sparked off an envlronmentally frlendly brand of economics. Free enterprlse and Marxist economists placed humans at the stage centre and tended to disregard the natural environment, seelng it as a free and inexhaustible reservoir Broken equilibrium and a bottomless rubbish dump (Cottrell 1978: 7). Whatever theroot cause ofenvlronmental degradation This exploitative attitudefostered traditional resource should be, Clements’s notton ofenduring equilibrium management practice that maximizes productmn suggested that humans had erred. Its message for con- of goods and services through sustained yield from servation was clear: the climax state is Nature’s deslgn balancedecosystems; adoptsutilitarian values that for bioticcommunities so humansshould respect regard human consumptlon as the best use of and preserve it. I t was thought that the equilibrium resources; and holds a continuous supply of goods for betweenhumansandNature had been upset and human markets as the purpose of resource manageshould be restored, that Nature’s deslgn princlples had ment(CortnerandMoote 1994). Such barefaced been forsaken and needed Immediate relnstatement. ‘resourclsm’ IS ecologically unsound. I t fails to recogThls response toenvlronmentaldegradatlon was nize limltstoexplomtionand, in consequence, a evldent in Paul Sears’s Daert.r on the Alurch (1949, the growingnumber of species, and even entire ecofirst edition of whlch appeared In 19.35, In the after- systems, are currently endangered. But, unsound or math of the Amerlcan dust-bowlcalamity. Sears not, I t persists. Even now, ecosystems are vlewed by advocated that each county In theUnitedStates some as long-lived, multi-product factories (Gottfrled should appoint an ecologist to adv~se on matrers of l992), o r , if you prefer, as Nature’s superstores.

222

LIFE, H U M A N S , A N D M O R A L I T Y the In mid-l960s, a new ‘environmental economics’ emerged that was sensitive to the needs of the environment. Kenneth Ewart Boulding (1966, 1970), for instance,complainedthathumans have operated a ‘cowboy economy’,in which success is measured in the amount of material turned over and in which resources are exploited in a profligate and reckless manner. Instead, he insisted, life on a small and crowded planet demanded a‘spaceship economy’, inwhich success is gauged by turnoverandthe watchwords areconservation, maintenance,and recycling.Eugene P. Odumemployedthe spaceship analogy as recently as 1989. He recalled the crisis on board the spacecraft Apollo 13. An explosion in the craft had caused damage to the vessel and threatened the lives of the crew.After several anxious hours, the crew managedto reach the safety of thelunar module and abandoned ship. Odum compared this event to the current environmental crisis on Earth. Our life-support systemsarebeingdestroyed,but, unlike the Apollo 13’s crew, humans cannot abandon ship and find a safe haven. The only option is to stay onboardSpaceshlp Earthand repair thedamage. Unfortunately, he said, humans do not know how to repair the planet, although he hinted that a civilization capable of putting people on the Moon should have witenoughtocomprehendthe mechanlcs of ecospheric dynamics and keep the cogs of the ecological machine running freely. Luckily, he observed, the Earth has its own regeneration systems, its own means of healing wounds. The view that the Earth has itsown powers of healing is supported by the Gaiahypothesis,the medicalanalogy beingmade clear in Lovelock’s (1989) use of the term ‘geophysiology’. (As an aside, I wouldpointoutthatthe impossibility of indefinitely sustaining life in a spacecraft was exploredin Brian Aldiss’sfictional ‘Helliconia’ trilogy, which also used an exotic setting tospeculate on thepossibilities of the Gaia hypothesis.) Second, technocentric environmentalists used the balanced ecosystem view of Naturetopromote the understanding of balanced ecosystem function as a basis for planetary management. Both ecocentrics andtechnocentricsthoughtthey could determme

what was safe for humans to do and what was not. Largely, they were mistaken (see Worster1994: 416). Many ideas of theequilibrium ecologlsts are used today. Equilibrium ecological thought also underpinsthe theory of islandbiogeography, as formulated by Robert H. MacArthurandEdward 0. Wilson in the 1960s, which is still used to aid the designing of nature reserves. Some conservationists have espoused the steady-state Gaia hypothesis as a blueprint for planetarymanagement (Myers 1987), and somevery green environmentalists have used it as a design for living. The equilibria1 ecological theme persists in the notion of sustainability. This is the well-meaning, but possibly misguided, idea that the ecosphere should be developed without compromising the ability of future generations to satisfy their needs. Or, put another way, sustainable development urges that diverse, functioning ecosystems should be preserved withoutdamagingthe economy; and that economies and human welfare need preserving as much as theintegrity of ecosystems (Gerlach and Bengston 1992). It was first presented in 1980 at an internationalforum in theWorld Conservatlon Strategy, by the Internatlonal Unlon for the Conservation of Nature(IUCN),and has become mainstay of much environmentalist philosophy. The IUCN identified three key areas for action: the maintenance of essential ecological processes and lifesupport systems, thepreservation of genetic diversity, andthesustainable use of species and ecosystems (Allen 1980). But the idea of sustainability is questionable. One problem lies in integratinghumanand biophysical factors. Humans interact with Nature through culture, often in symbolic ways that are not comprehended by biological or physical ecosystemmodels (Gerlach and Bengston 1992). In other words, humans can generate wants and capabilities of meeting these wants that lie outside the naturalecological order. This is often difficult for scientists to understand, and leads to their focusing on envlronmental protectionandrestoration,ratherthan facing the greater challengeof understanding social and cultural interactions with thebiophysical world. Thevery Idea

L I F E , HUMANS, AND M O R A L I T Y of sustainability itself is a curious human construct. Laudable though the idea of sustainability might be, there are problems with it, not the least of which is the lack of a definition.It was originally taken to mean meetingthe needs of the present generation without compromising the ability of future generationsto satisfy their needs (World Commission on EnvironmentandDevelopment1987),butat least eight interpretatlons are available (Gale and Cordray 1991).It is therefore a buzzword that lacks bite. Anotherproblem is its inadvertent arrogance shouldthe present generation be so presumptuous and foolhardy as to anticipate the needs of future generatlons?Thesepithyquestionsshouldnot be ducked In ecosystem management initiatives

Evolutionary disequilibrium Some time In the 197Os, the notions of homeostasls, balance, and stability i n Nature collapsed. They were supplanted by the idea of evolutionary disequilibrium. Thedramatlcchange of paradigms was, In part, a reflectlon of the human soclety at that time. I t arose when i t wasGashionable to see neither Nature nor society as a stable entity,steadfastly wlthstanding all attempts to dislodge it from a global equilibrlum state. Instead,a view emerged of all hlstory as a record of disturbancespringingfromboth‘cultural and natural agents, including droughts, earthquakes, pests, viruses, corporate takeovers, loss of markets, new technologies, increaslng crime, new federal laws, and even the invasion of Amerlca by French literary theory’ (Worster 1994: 424, emphasisin original). Disequilibrium ecologists seemed divided among themselves on the advice they should give to society about how to act over the environment. One group, reflecting some of the new disequilibrium thlnking, began to challenge the public perception that ecology and environmentalism were the same thing. Some ecologlsts became disenchanted with trying to conserve a healthy planet. Nature is characterized by highlylndivdualistic assoclatlons, so why attempt to constrain I t ? This anarchlc argument, if taken to theextreme, could have revlved social Darwinism andstood in antitheslstothe conservation ethic of

co-operation and collective action suggested by Odum’s balanced ecosystems. In the event, a mildly anarchic view of Nature was taken by some ecologists (p.215).Anothergroup of ecologists, instark contrast,drew differentconcluslons from thedisequilibriumtrendswithinthendiscipline.Daniel B. Botkin(1990) was one of themostarticulate advocates of a new, chastened set of environmentalist policies. H e favoured an environmentalism that was more friendly towards manipulating and dominating Nature,butthat tolerated modern technology and progress, and human deslres for greater wealth and power. This was not the preservation of a balanced nature, so much as a responsible attitude to technological developments and their effects on an environment that would inescapably change. Botkin’s specific proposals were somewhat vague. He cautioned that we should not engineer Nature at an unnatural rate nor In novel ways. But In a worldwhere disturbance is commonplace and change is the rule, how may the unnaturally rapld andthe novel be identified and defined!

The edge of chaos The implications of chaotic ecosystem dynamics for conservation are far from clear. How are conservationists to use a chaotic design? Answers to this question have to confront a paradox: a chaotic view of Nature I S at once exhilaratingandthreatening. ChaoticNature, so irregularandindividualisticin character, appears almost impossibly difficult to admlre or to respect, to understand or to predict. It seems to be a world In which the security of stable, permanent rules IS gone forever, a dangerous and uncertain world that Inspires no confidence (Prigogine andStengers1984:2 12-1 3). Thisdark aspect of chaos might promote a feeling of alienation from the naturalworld,and cause peopletowithdrawinto doubt and self-absorptlon (cf. Worster 1994: 413). It mlght also setpeople wondering how theyshould behave In a world where chaos reigns. With natural disturbance foundeverywhere,why shouldhumans worry about doing their own bit of disturbing? Why not join individuals of all other specles and do their

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LIFE, H U M A N S , A N D M O R A L I T Y own thlng, free of guilt that in doing so they would cause any specific damage? As Worster (1994: 413) put it, what does the phrase 'envlronmental damage' mean In a world so full of natural upheaval and unpredictability? However, chaos does not have to be portrayed in such gloomy and doom-laden terms. Chaotlc Nature has abrlghtandedifylngaspect,too.In a chaotic world,communltles, ecosystems, and socletiesare sensitivetodisturbance. Small disturbances can, sometimes,growand cause the communities, ecosystems, or societies to change. Consequently, i t is a world In which indivldualactlvlty may have major slgnificance (cf. Prlgogine and Stengers 1984: 3 13). It is apost-modern worldinwhlch Increased Inclividualityand diversityencourages agreat overall harmony (a notlon akin to Taoist beliefs). Moreover, the newfangled theory of chaotic dynamlcs is leading to the discovery of hiddenregularities In natural processes, the application of whlch is prowng most salutary (Stewart 1995). Satellites can now be sent to new destlnations, ferried by gravitational forces through a fractal Solar System, with far less fuel than was once thought possible. A dishwasher, involvlng tworotatingarmsthatspln chaotically, has been invented in Japanthat removes dirt fromdishes using less energy than in a conventlonal dishwasher. Some forests and fisheries are being better managed through an understanding of thelr chaotic behaviour. And, chaos theory is glvlng us a breathtaking view of our Universe, and persuading us to look again at our place wlthin It. It offers us a new design with which to reconsider our world, and a new framework wlthln which to review our fears and our hopes.

Forward from the past: the biodiversity bandwagon By the last decade of twentieth century, ecologlsts were undecided about the implications of their subject for modern technological civilization. Nonetheless,theyfoundthemselves regrouplng around a new conservation ideal - biodiversity. I t mattered notthatNature was chaotlcandunpredictable.It was also gloriously diverse, and, for all its disturbing

and perverse ways, and its abiding ability to evade I t still needed our love, our respect, and our help (cf. Worster 1994: 420). This V I ~ Whas supplied a conservation ethlc, and a source of inspiration (and funding) for biogeographers, for the closing years of the twentieth century, and perhaps beyond.

our understanding,

SUMMARY There aremoral grounds for grantlngrightsto animals, plants, andeven landscapes. In the late 1990s attltudes towards anlmals are more sympathetic than they were In the past. Thls sympathy I S seen in the publicoutcryagalnst cruel methods of culling, in the measures taken to stop the wildlife trade, and in the new role for zoos In conservation. Attitudes towards Nature are vaned and reflected in different brands of environmentalism. The two main opposing brands ofenvironmentalismare ecocentrism and technocentrism. Ecosystem managementaccommodates human needs andaspirationswlthin anecocentric perspective.Biogeographical and ecological theory mforms conservation practice. During the twentieth century the leadingideas in biogeography andecology have changed.In consequence,conservation guidelines have changed. As the twenty-first century draws near, the chaotic world of the theoretical ecolo p s t stands parallel with the blodiverse world of the practlcal ecologist.

E S S A YQ U E S T I O N S Should animals, plants, and landscapes be granted rights?

To what extent does biogeographical and ecological theory inform conservation practice? What is so special about ecosystem management?

LIFE, H U M A N A SN , MDO R A L I T Y

FURTHER READING Anderson, E. N . (1996) Ecologies of the Heart: Emotfon. Belid and the Enzironnrent, New York: Oxford Universlty Press. Botkm, D. B. (1990) Discordant Harmonies: A Neur Ecology for the Tuznty-First Century, New York: Oxford University Press. Commlttee onScientific Issues in the Endangered Species Act, National Research Council (1995) Scienle and the Endangered Spec.ies Al.t, WashingtonDC: National Academy Press. Cooper, N . andCarling,R.C. J. (eds) (1996) Ecologists and EthicalJudgernents, London: Chapman & Hall. Norton, B. G. (1991) Touwd Untty Environmentalists, New York: Oxford Press.

Among University

Ryder,R. D. (ed.) (1993) Anutral Welfare and the Envirotment, London: Duckworth and RSPCA. Shafer, C. L. (1990) Nature Reserves: Island Theory and Consetvatton Practtte, Washlngton and London: Smlthsonlan Institunon Press. Sylvan, R.andBennett,D. (1994) The Greening of Ethm: From Hnmun Charwnurn t o Deep-Green Theory, Cambridge: Whlte Horse Press; Tucson, Universlty of Arlzona Press.

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GLOSSARY

Table

GI

The geological time-scale for the last 570 million years

Era and subera

Period

Epoch

Age range (millions of year ago) Start

Cenozoic era Quaternary sub-era

Pleistogene

Tertiary subera

Neogene Palaeogene

Holocene Pleistocene Pliocene Miocene Oligocene Eocene Palaeocene

0.0 1 1.64 5.2 23.3 35.4 56.5 65

Finish

0 0.01 1.64 5.2 23.3 35.4 56.5

Mesozoic era Cretaceous Jurassic

Triassic

Late Early Late Middle Early Late

Middle Early

97 145.6 157 178 208 235 24 1 245

65 97 145.6 157 178 208 235 24 1

290 362.5 408.5 439 510 570

245 290 362.5 408.5 439 510

Polaeozoic era Permian Carboniferous Devonian Silurian 0rdovician Cambrian

GLOSSARY abiotic Characterized by the absence of life; inanimate. acidophiles Organisms that love acidic conditions.

atonal soils Soils in which erosion anddeposition dominate soil processes, as insoilsformedin river alluvium and sand dunes. bacteria Micro-organisms, usually single-celled, that exist as free-living decomposers or parasites.

adaptation The adjustment or the process of adjustment by which characteristm of an entire organism, or some of its structures and functions,become better suited for life in a particular envlronment.

barrier Any terrainthathinders dispersal of organisms.

aerobicDependingon, presence of oxygen.

basal metabolic rate The rate of energy consumption by an organism at rest.

or characterized by,

the

algae Simple, unicellular or filamentous plants. alkaliphilesOrganismsthat environments.

flourish in alkaline

reactlon of organismstoglven

benthic Pertalning to the bottom of a water body.

allelopathicPertainingtothe influence of plants upon each otherthroughtoxicproducts of metabolism - a sort of phytochemical warfare. allogenic Originating from outslde a community or ecosystem. anaerobicDependingon, absence of oxygen.

behaviourThe circumstances.

or preventsthe

or characterizedby,

the

anthrax An infectious and often fatal disease caused by the bacterium B a c i l h antbraczs; common in cattle and sheep.

biocideA poison or othersubstance used tokill pests (and inadvertently other organisms). bioclimatesTheclimaticconditionsthat living things.

affect

biodiversity The diversity of specles, genetic information, and habltats. biogeochemical cycles The cycling of a mineral or organic chemical constituent through the biosphere; for example, the carbon cycle.

apical buds Buds located at the tips of shoots.

biologicalcontrolThe use of natural ecological lnterrelatlonships to control pest organisms.

aquifer Asubsurface geological formation contamng water.

biomagnification The concentration substances up a food chain.

archipelagoAgroup group.

of islands,typically

a large

aridity The stateor degree of dryness. atmosphereThe gaseousenvelope of theEarth, retained by the Earth’s gravitational field. autogenic Originating from inside a community or ecosystem. autotrophsOrganisms - greenplantsandsome mlcro-organisms - capable of making their own food from inorganic materials. available moisture Precipitation less evaporation.

of certain

biomantle The upper portion of soil that is worked by organlsms. biomass The mass of living materlal In a specified group of animals, or plants, or in a community, or in a unit area; usually expressed as a dry weight. biome A community of animals and plants occupying a climatically uniformarea on a continental scale. biosphere Life andlife-supporting systems - all livingbeings,atmosphere,hydrosphere,and pedosphere. biota All the animals(fauna) and plants (flora) living in an area or region.

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228

GLOSSARY biotic Pertalning to life; anlmate. biotic potential The maximum rate of population growth that would occur In the absence of environmental constraints. borealforest A plantformation-type associated wlthcold-temperateclimates (cool summersand long wlnters); also called talga and coniferous evergreen forest. Spruces, firs, larches, and pines are the domlnant plants. browsers Organlsms that feed mainly on leaves and

young shoots, especially those of shrubs and trees. bryophytesSimple landplantswlthstemsand leaves but no true roots o r vascular tissue: mosses, hornworts. and liverworts.

cohort Indivlduals In a population born durlng the same perlod of t m e , e.g. during a partlcular year.

commensalism An interaction between two species In which one specles (the commensal)benefits and the other specles (the host) is unharmed. communityassemblyThecomingtogether species to form a communlty.

of

competitionAn interactlonbetweentwospecies trying to share the same resource. competitive exclusion principle The rule that two species wlth Identical ecological recluIrements cannot coexist at the same place. congener An organism belonging to the same genus as another or others.

to the family cactiNewWorldplantsbelonging Cactaceae with thick, fleshy. and often prickly stems.

conifers Trees of the Order Coniferales, commonly evergreen and bearing cones.

calcicoles (calciphiles) Plants that love soils rich in calcium.

consumption Community respiration.

calcifuges (calciphobes) Plants that hate in calcium.

soils rlch

capillary pressure The force of water in a capillary tube. carrion Dead and decaying flesh

continentaldriftThe differential movement contlnents caused by plate tectonlc processes. corolla The inner envelope of a flower; fused or separate petals.

I t IS

of

made of

corridors Dispersal routes offerlng little or no remtance to migratlng organisms.

period A slice of geological tlme carrying capacity The maxlmum population that an Cretaceous spanning 145.6 million to 65 million years ago; the environment can support without environmental youngest unit of the Mesozoic era. degradation occurrlng. caviomorph rodents A suborder of the Rodentla

cullingThe act of removing o r killing‘surplus’ animals In a herd or flock.

Cenozoic era A slice of geological time spannlng 65 million years ago to the present; the youngest unlt of the geological eras.

cursorial Adapted for running.

chasmophytes Plants that live in crevlces. chomophytes Plants that live on ledges.

cuticle A protecttve layer of cutin (a wax-like, waterrepellent materlal) covering the epldermls (outermost layer of cells) of plants.

chronosequence A time sequence of vegetatlon or soils constructed by uslng sltes of different age.

cycad Any gymnosperm of the Family Cycadaceae looklng like a palm tree but topped with compound, fern-like leaves.

circulatory system The system ofvessels which blood is pumped by the heart.

decomposer An organism that helps organic matter.

through

to

break down

GLOSSARY demeA group of interbreeding Indivduals; a local population. density-dependent factors Causes of fecundity and mortality that become more effectlve as population density rises. density-independentfactors Causes of fecundity and mortality that act independently of populatlon density; natural disasters are an example. detritivoreAnorganismthatcomminutesdead organic matter. detritus Dislntegrated matter. diatom A microscopic,unrcellular, marine(planktonic) alga with a skeleton composed of hydrous opaline silica. dicotyledonous plant species Plants belonglng to the Dlcotyledoneae, one of the two major divisions of flowering plants. They are characterlzed by a pair of embryonic seed leaves that appear at germination. disharmonious community community A of animals and plants adapted to a climate that has no modern counterpart.

El Niiio Theappearance of warm water in the usually cold water regions off the coasts of Peru, Ecuador, and Chile. endangered species A specles faclng extlnctlon. endemicspeciesA place.

species nativeto

a particular

environmentThe biological (biotic)and physical (ablotic) surroundings that Influence Individuals, populations, and communities. environmentalethicsA philosophy dealingwith theethics of theenvironment,includinganimal rlghts. environmental factor Any of the biotic or abiotic components in the environment, such as heat, moisture, and nutrient levels. environmental gradient A continuous change in an environmental factor from one place to another; for example, a change from dry to wet conditlons. environmentalism A movementthatstrivesto protect the environment against human depredations.

dispersal The spread of organisms into new areas.

Eocene epoch A slice of geologlcal time spanning 56.5 million to 35.4 million years ago.

dispersalrouteThepath organlsms.

epiphyte A plant that grows on another plant or an object, uslng I t for support but not nourishment.

followed by dispersing

distributionThegeographical area occupied by a group of organlsms (species, genus, family, etc.); the same as the geographical range. d r o u g h t A prolonged period with no rain. ecological equivalents Species of different ancestry but with the same characteristics livlng In different biogeographical regions. Also called vicars. ecosphere The global ecosystem - all life plus its life-support systems (air, water, and soil).

estuarine Of, pertaining to, or found In an estuary. evaporation The diffusion of water vapour Into the atmosphere from sources of water exposed to the alr (cf. evapotranspiration). evapotranspiration Evaporation plus the water dischargedintotheatmosphere by plant transpiration. evergreen A plant with year round.

ecosystem Short for ecolog~calsystem - a group of organismstogetherwlththe physical environmentextinctionThedemise taxon. with interact. which they ecotone A transirlonal zone between twoplant communities.

foliage that stays green all of a specles o r any other

extirpation The local extinction of a species or any other taxon.

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230

GLOSSARY extremophiles Organisms that relish ultra-extreme conditions.

and comprising large parts of South America, Africa, India, Antarctica, and Australia.

fauna All the animals living in an area or region.

grazer An organism that and herbs.

as measured by fecundityReproductivepotential thenumber of eggs,sperms, or asexual structures produced.

feeds on growing grasses

gross primary production Production respiration losses are accounted.

before

filter route A path followed by dispersing organisms that,owingtotheenvironmental or geographical conditions, does not permlt all species to pass.

habitat The place where an organism lives.

flora All the plants living in an area or region.

heath A tract of open and uncultlvated land supporting shrubby plants, and especially the heaths (Erzcu spp).

food chain A simple expresston of feeding relatlonshipsinacommunity,startingwithplantsand ending with top carnivores.

habitatselectionThe selection of particular a habitat by a dispersing individual.

heliotropism The growthof a plant towards or away food web A network of feeding relationships within from sunlight. a community. helophytes Marsh plants. frugivorous Pertaining to fruit-eating.

heterogeneous An environment comprising a mosaic of dissimilarspatialelements;non-uniform environment.

GaiahypothesisThe idea thatthe chemical and. physical conditions of the surface of the Earth, the atmosphere,andthe oceans are actlvely controlled by life, for life.

heterotrophs Organismsthatobtainnourishment from the tissues of other organisms.

geneticvariabilityA measure of thenumber genetic diversity wlthin a gene pool.

homeostasisAsteady-statewithinputscounterbalancing outputs.

of

genus (plural genera) A taxonomic group of lower rank thana family that consists of closely related specles or, in extreme cases, only one specles. geodiversity The diversity of the physical environment. geographical range Thearea occupied by a group of organisms (species, genus, family, etc.); the same as the distribution.

geosphere The solid Earth - core, mantle,and crust. geothermal springs Springs of water made hot the Earth’s internal heat.

by

homeotherm An animalwhichregulatesits body temperature by mechanisms withln its own body; an endotherm or ‘warm-blooded’animal (cf. poikilotherm). homogeneousReferringtoa spatial elements; uniform.

mosaic of similar

humification The process of humus formatlon. hummock-hollow cycle A peatland cycle involving the infilling of a wet hollow by bog mosses until a hummock is formed and colonized by heather. The heather eventually degenerates and the process starts again.

germination The time of sprouting in plants.

h u m u s An amorphous, colloidal substance produced by soil micro-organisms transforming plant litter.

G o n d w a n a Ahypothetical, Late Palaeozoic supercontinent lying chiefly in the Southern Hemisphere

hurricaneAviolentandsometimesdevastating tropical cyclone.

GLOSSARY hydrophyte A plantthatgrows submerged in water.

wholly or partly

hydrosphere All the waters of the Earth. hyperparasite A parasite parasite.

that lives on another

hyperthermophile An organismthatthrives ultra-hot conditions. Ice Age An oldterm interglaclal sequence.

in

for theQuaternary glaclal-

iceage A timewhen Ice formsbroad sheetsin middle and high latitudes (often in conjunction with the widespread occurrence of sea ice and permafrost) and mountain glaciers form at all latitudes. interspecific competition Competition among species.

LaurasiaTheancient,NorthernHemisphereland mass that broke away fromGondwanaabout 180 million years ago and subsequently split into North America, Greenland, Europe, and Northern Asia. leaf-areaindexTheratio of total leaf surface to ground surface. For example, a leaf-area index of 2 would mean that if you were to clip all the leaves hanging over 1 m2 of ground, you would have 2 m2 of leaf surface. legume A plant of the family Leguminosae, characteristically bearingpodsthatsplitintwoto reveal seeds attached to one of the halves. lichen A plant consisting of a fungus and an alga. lifetable A tabulation of thecompletemortality schedule of a population. life-form The characteristics of a mature animal or plant.

intestinal endosymbionts Symblotic organisms living in another organism's gut.

littoral Shallow water zone

intraspecificcompetitionCompetitionwithin species.

a

local extinction (extirpation) The loss of a species from a particular area.

intrazonal soils Soils whose formation is dominated by local factors of relief or substrate; an example is soils formed on limestone.

macronutrients Chemical elements needed in large quantltles by living things.

introducedspecies or introduction A species released outslde Its natural geographlcal range.

maquis A shrubbyvegetation trees and bushes.

isolines A lineon value.

mass extinctions Extinction episodes in which large numbers of specles disappear.

a mapjoiningpoints

of equal

Jurassic period A slice of geological time spanning 208 million to 145.6 million years ago; the middle unit of the Mesozolc era. keystone species A specles that plays a central role ina communlty or ecosystem, its removal havlng far-reachlng effects. landbridge A dry land connection between two land masses that were prevlously separated by the sea. land ethic A set of ethlcal princlples declaring the rights of all naturalobjects(animals,plants,landscapes) to continued exlstence and, In some places at least, continued exlstence In a natural state.

of evergreen small

meadow Grassland mown for hay. mechanistsProponents of the vlew thatall biological and ecological phenomena may be explained by the interaction of physlcal entities. megafauna Large mammals, such as mammoths and sloths,contrastedwithsmallmammals, such as rodents and insectivores. megaherbivores Large browsers and grazers, such as elephants and rhinoceroses. mesophyte A plant flourishing under mesic (not too dry and not too wet) conditions.

23 1

23 2

GLOSSARY Mesozoic era A slice of geological time spanning 245 million to 65 million years ago comprising the Triassic, Jurassic, and Cretaceous perlods. metabolic processes The many physical and chemical process involved in the maintenance of life. micronutrients Chemical elements required small quantities by at least some livlng things.

in

orobiomes Biomesassociated climatic zones on mountains. overwinterTo place.

withthealtitudinal

spendthewinter

in aparticular

Palaeocene epoch A slice of geological time spanning 65 million to 56.5 million years ago; the oldest unit of the Palaeogene period.

visible tothe

PalaeogeneperiodA slice of geological t m e between 6 5 million and 23.3 million years ago.

mineralization The release of nutrients fromdead organlc matter.

Palaeozoic era A slice of geological time spanning 570 million to 245 million years ago; comprises the Cambrian, Ordovician, Silurian, Devonian, Carboniferous, and Permian periods.

micro-organism An organismnot unaided naked eye.

Miocene epoch A slice of geological tlme spanning 23.3 million to 5.2 million years ago. mutualism An obligatoryinteraction between two species where both benefit. natural selection Theprocess by which the environmental factors sortand sift geneticvariabilityand drive evolution. Neogene period A slice of geological t m e between 23.3 million and 1.64 million years ago. net primary production Gross primary production

less respiration.

Pangaea The Triassic supercontlnent comprising all the present continents. pathogen Anagent, such as bacterlum or fungus, that causes a disease. pedobiomes Areas of characteristic vegetation produced by distinctive soils.

pedosphere All the soils of the Earth. Permian period A slice of geological time spanning 290 million to 245 million years ago.

neutralism An lnteractlon in whlch neither species is harmed or benefited.

photoperiod The duration of darkness and light.

been cut, or a forest that has been undisturbed for a long tlme.

phytotoxic Poisonous to plants.

photosynthesis The synthesisof carbohydrates from carbon dioxide and water by chlorophyll, using light neutrophiles Plants that like neutral conditlons, not as an energy source. Oxygen is a by-product. too acldic and not too alkaline. phytomass The mass of living plant materlal in a conversion of atmospheric nitrogen-fixing The community, or in a unit area; usually expressed as a nitrogen into ammonium compounds by some dry welght. bacteria. phytoplanktonPlantplankton:theplantcommunutrientA chemical element required by at least nity in marlneand freshwatersystemwhich floats some living things. free in the water and contains many species of algae old-growthforestAvlrgln forest that has never and diatoms. plantformationThevegetatlonalequivalent of a biome, that IS, a community of plants of like physOligocene epochA slice of geological time spanning iognomy (life-form) occupying a climatically uniform 35.4 million to 23.3 million years ago; the youngest area on a continental scale. unit of the Palaeogene period.

GLOSSARY Pleistocene epoch A slice of geological tune spanning 1.64 million to 10,000 years ago; the older unlt of the Plelstogene period (or Quaternary sub-era). of geoPleistogene period The mostrecentslice logical time between 1.64 million years ago and the present;subdividedinto Pleistocene and Holocene epochs.

Pliocene epoch A slice of geological time spanning 5.2 millionto 1.44 million years ago;theyounger unit of the Neogene period. poikilotherm An organism whose body temperature is determined by the ambient temperature and who can controlits body temperature only by taking advantage of sunandshadeto heat up or to cool down; a 'cold-blooded' animal (cf. homeotherm). prairie An extensive area of natural, dry grassland; equlvalent to steppe In Eurasla. profundal Deep water zone. propagule The reproductive body of a plant. protocooperation non-obligatory A interaction between two species where both benefit. psychrophile Organisms that temperatures.

flourish at low

race An anlmal or plant populatlon that differs from other populations of the same specles in one or more hereditary characters.

salinity Saltiness. saltatorial Adapted for hopping and jumping. savannah Tropical grassland. saxicolous Growlng on or living among rocks. scavenger An organism feedingon dead animal flesh or other decaying organic material. seasonality The degree of climatlc contrast between summer and winter. sessile Permanently attached moving.

or fixed; not free-

social Darwinism Darwln's doctrme of the survlval of the fittest applied to society. soil Rock at, or near, the land surface that has been transformed by the blosphere. solar radiation The total electromagnetlc radiation emltted by the Sun. Electromagnetic radiatlon is energypropagatedthrough space orthrougha material as an interaction between electric and magnetic waves; occurs at frequencies rangmg from short wavelength,high frequencycosmlc rays, through medium wavelength, medium frequency vlsible light, to long wavelength,low frequency radio waves. speciation The process of species multlplication. species A reproductively isolated collection of interbreeding populations (demes).

radioisotopeA form of a chemical elementthat undergoes spontaneous radioactlve decay.

steppe An extensive area of natural, dry grassland; equlvalent to prairiein North American terminology.

rain shadow A region wlth relatively low rainfall that is sheltered from raln-bearingwinds by high ground.

or stem through which air and water vapour pass.

respiration A complexserles of chemical reactions in all organlsms by which energy is made available for use. The end products are carbon dioxlde, water, and energy. rhizome A root-like stem, usually horizontal, growing underground with roots emerging from Its lower surface and leaves or shoots from Its upper surface.

stomata (singular stoma) Epidermal pores in a leaf sweepstakes route A path along whlch organisms may disperse, but very few will do so owing to the difficulties involved. symbiotic Pertaining to two or more organisms of differentspecies livlngtogether.Ina narrow sense equivalent to mutualism. taxon (plural taxa) A population or group of populations (taxonomic group) that is distinct enough to

233

234

GLOSSARY be given a distinguishing name and to be ranked in a definite category.

territoryAn area defended by an animalagainst other members of its specles (and occasionally other species).

ungulates Hoofed mammals. vascular plant A plant containing vascular tissue, theconducting system which enables waterand minerals to move through it.

vegetation Plant life collective; the plants of an area. Tertiary period A slice of geological time spanning vernalization The exposure of some plants (or their 65 million to 1.64 million years ago and the youngest seeds) to a period of cold that allows them to flower unit of the Cenozoic era; now designated a sub-era. or to flower earlier than usual. theory of islandbiogeography A modelthat vicariance The splitting apart of a species distribuexplains the species diverslty of Islands chiefly as a tion by a geological or environmental event. function of distance from themainlandand island vicars Species of different ancestry but with the same area. characteristlcs living in different biogeographical thermophiles Organisms that love hot conditions. regions. Also called ecological equivalents. toposequence The sequence of soils and vegetation vitalists Proponents of the Idea that some kind of found along hillslope, a from summitto valley living force resides in organisms. bottom. volatilization To convert to vapour. trace elementA chemical element used by an organism In minute quantities and vital to its physiology. wetlands All the places in the world that are wet for at least part of the year and support a characterlstic tree-lines The altitudinal and latitudinal limits of soils and vegetation; they Include marshes, swamps, tree growth. and bogs. tree-throw The toppling of trees by strong winds.

xerophyte A plant adapted for life in a dry climate.

Triassic period A slice of geological time spanning 245 million to 208 million years ago; the oldest unit of the Mesozoic era.

zonobiome One of nine, climatically defined, major community units of the Earth.

or

zoomass The mass of livinganimalmaterialin a community, or in a unit area; usually expressed as a dry weight.

tsunamis The scientific (and Japanese) name for tidal waves.

zooplankton Animal plankton; aquatic invertebrates living in sunlit waters of the hydrosphere.

trophic Of, feeding.

or pertaining to, nourishment

FURTHER READING

General biogeography books

Borhldi, A. (199 1) Phytogeography and Vegetatton Ecology of Cuba, Budapest: A k a d h l a l Kiad6.

Cox, C. B. and Moore, P. D. (1993) Biogeography A n Ecologtcal and Euoluttonaty Approach, 5thedn,Oxford:Bowman, R , I, (ed,) (19(;6) The Gala'pagos Blackwell. (Proceedings of theSymposium of theGalipagos Already a classic. A very popular textbook. International Sclentlfic Project), Berkeley and Los George,

w,

(1962) Anitna(Geography,

London:Angeles,California:

Helnemann. drift, H. C. Written before the revival of continental well but worth a look. Pears, N. (1985) Basic Btogeography, 2nd edn, Harlow: Longman. Gressitt, J. A good textbookonecologicalbiogeography. Tivy, J. (1992) Biogeography: A Study o f Plants tn the Ecosphere, 3rd edn, Edinburgh: Oliver & Boyd. A good introductory textbook.

University of Californla Press.

(ed.) (1984) and BiogeograPhY in Sri L n k a (Monographiae Biologicae, Vol. 57), The Hague: W. Junk.

L.(1982) (ed.) Biogeography and Ecology of New Guinea, 2 vols (Monographiae Biologicae,Vol. 42), The Hague: W. Junk. Kuschel, G . (ed.) (1975) Biogeography and Ecology in Neu, Zealand (Monographlae Biologicae,Vol. 27), The Hague: W. Junk.

Regional biogeography Here is a small selection of regional biogeographlcal and ecological texts. A GEOBASE search will reveal many more. Battistini, R. and Richard-Vindard, G . (eds) (1972) Bzogeography and Ecology in Madagascar (Monographiae Biologicae, Vol. 21), The Hague: W . J u n k .

Stoddart, D. R. (ed.) (1984) Biogeography and Ecology ofthe Seychelles Zslands (Monographiae Biologicae, Vol. 5 5 ) , The Hague: W . J u n k . Whitmore, T. C. (ed.) (1981) Wallace's Line and Plate Tectontcs (OxfordMonographs inBiology, Vol. l), Oxford: Clarendon Press.

236

F U R T H E RR E A D I N G Whitmore, T. C. (ed.)(1987)BiogeographicalEvolutron of the Malay Archipelago (Oxford Monographs in Biology, Vol. 4), Oxford: Clarendon Press.

Williams, W.D. (ed.) (1974) Biogeography and Ecology zn Tasnrunta (Monographlae Biologicae, Vol. Hague: The25), Junk. W. Woods,C. A. (ed.)(1989) Bzogeography of the We.rt Indies: Past. Present. and Fzrtzrre, Gainesville,Florida: Sandhill Crane Press.

aspects

Ecological Dansereau, (1957) p.

Biogeography: An Eco/ogll~/

Perspective, New York: Ronald Press. Old but have a look.

Forman, R. T. T. (1995) L n d Mosaics: The Ecology o f hzdsrapes and Regions, Cambridge: Cambridge University Press. A weighty tome on landscape ecology with some useful sections for biogeographers.

Kormondy, E. J. (1996) Concepts of Ecology, 4th edn, Englewood Cliffs, NJ: Prentice Hall. An excellent basic ecology text. Physzologtcal Plant Ecology: Larcher, W. (1995) Ecophyszology and Stress Phystology of Functional Groups, 3rd edn, Berlin: Springer. First-ratecoverageofphysiologicalaspectsof plant ecology. Stoutjesdijk. and Barkman, P. (1992) J.J. Microdimate, Vegetatron and Furma, Uppsala, Sweden: Opulus Press. Unusual but highly interesting book. Viles, H. A.(ed.)(1988) Brogeonlorphology, Oxford: Basil Blackwell. Links life to geomorphology.

Whelan, R. J. (1995)The Ecology of Fire, Cambridge: Cambridge University Press. A clear introduction to the subject.

Historical aspects (1995) C. Global Bzogeography Brlggs, J. (Developments In Palaeontology andStratigraphy, 14)~ Amsterdam: Elsevier. An advanced text but worth dipping into. Cox, C. B. and Moore, P. D. (1991) Bzogeogruphy: A n Erological and Evolutionary Approach, 5th edn, Oxford: Blackwell. basic coverageOf historical aspects.

Cronk, Q. C. B. and Fuller, J. L. (1995) Plant Invaders: The Threat t o Natural Ecosystons, London: Chapman & Hall. Many examples. Zoogeography: The Darlington, P. J., Jr (1957) Geographical Dzstributron o f Aninfal.r, New York: John Wiley & Sons.

and Plants, London: Chapman & Hall. A little gem.

Hallam, A. (1995) A n Ozrtline of Phanerozoic. Biogeography, Oxford: Oxford University Press. A palaeontological perspective. Nelson, G. and Rosen, D. E. (eds) Vicariance Biogeugraphy: A Crrtrqzre (Symposium of the Systematics Discusslon Group of the American Museum of Natural History, 2 4 May 1979), New York: Columbla University Press. Only for the brave. Udvardy, M. D. F. (1969) Dynumtc Zoogeography, u'rth SpeL'ialReference to LandAnzmah, New York:Van Nostrand Remhold. Rather old, but still deserves to be read.

FURTHER READING Wallace, A. R. (1876) The Geographrcal Distrihutron of Animals: with a Study of the Relattons of Ltvrng and Extrnct Faunas asEluc1datzng the Past Changes of the Earth's Suvface, 2 vols, London: Macmillan. All biogeographers should read this great work. Woods,C.A.(ed.) Biogeography of the West Indies: Past. Present. and Future, Gamesville, Florida: Sandhill Crane Press. Caribbean case studies.

Single populations Hanskl, I. and Gilpin, M. E. (1997) Metapopulation Biology: Erology,Genetrcs. and Evolution, New York: Academic Press. Advanced but worth reading. Kormondy, E. J. (1996) Concepts of Ecology, 4th edn, Englewood Cliffs, NJ: Prentlce Hall. Relevant sections worth reading. Kruuk, H. (1 995)W i l d Otters: Predatzon and PoprdaOxford: Oxford University Press. A good case study.

trons,

I. P.,Smith,R.H.,and Perry, J.H.,Wolwod, Morse, D. (1997) Chaos in Real Data: Analysis on Nonlineur Dynanmlr from Short Ecologiczl Time Serres, London: Chapman & Hall. If mathematics is not your strong point, forgetit. Taylor, V. J.andDunstone,N.(eds) (1996) The Exploitation o f M a m d Populations, London: Chapman & Hall. Examples of exploitation. Tuljapurkar, S. and Caswell, H.(1997) StructuredPopulatlon Models, New York: Chapman & Hall. Somewhat demanding.

Interacting populations Crawley, M. J.(1981) Herbluory: TheDynamirs of Anmal-Plant Interac-tions (Studies in Ecology, Vol. lo), Oxford: Blackwell Scientific Publications. A superb book, despite the mathematics. An excellent basic text.

Grover, J. P. (1997) Raource Competitron, New York: Chapman & Hall. Up-to-date exposition. Jackson, A. R. W. and Jackson,J. M. (1996) Environmental Science: TheNatural Environment andHuman Impact, Harlow: Longman. Excellent text. Kingsland, S. E. (1985) ModelingNature: Eprsodes rn the History of Population Brology, Chicago and London: Unlverslty of Chicago Press. A first-rate account of the history of population biology. More exciting than it might sound. Kormondy, E. J. (1996) Concepts o f Ecology, 4th edn, Englewood Cliffs, NJ: Prentice Hall. Clear explanations of basic ideas. MacDonald, D. (1992) The Velvet Clauj: ANatural History ofthe Carnivores, London: BBC Books. All about carnivores with beautiful photographs.

Communities Archibold, 0. W. (1994) Ecology o f World Vegetation, New York: Chapman & Hall. Covers ecological aspects of plant communities. Chameides, W . L. and Perdue, E. M. (1997) Bmgeoo'hen~~czl Cycles: A Cot~I~uter-lntwactiveStudy o f Earth System Sc1enr.c andGlobalChange, NewYork: Oxford University Press. If you demand much, this is the one for you. Hochberg, M. E., Colbert, J., and Barbault, R. (eds) (1996) Aspects of the Genesrs and Marntenance o f Biological Dtversity, Oxford: Oxford Unlversity Press. Contains many case studies, though not an easy read. Jeffrles, M. L. (1997) Blodiversltyand London and New York: Routledge.

Conservation,

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F U R T H E RREADING Lawton, J. H. and May, R. M. (eds) (1995) Extinctton Rates, Oxford: Oxford University Press. Lots of recent figures on extinction rates. Morgan, S, (1995) El.ology and Envtronment: The of Lge, Oxford: Oxford University Press. A good starting text.

Community change

of Wetlands, Committee Characterizatlon on National Research Council (1995) Wetlands: cycles and Boundaries* Washington, DC: National Academy Press. Readable case study.

Polis, G . A.andWinemiller, K. 0. (1995) Food Webs: Integration ofparternsandDynamtlJ, New York: Chapman & Hall. Not easy, but worth a look.

Gates, D. M. (1993) Climate Change and Its Btologrcal Consequences, Sunderland, Massachusetts:Sinauer Associates. A clear account.

Quammen, D. (1996) The Song of the Dodo: Island Age of Extrnctions, London: Biogeography in an Hutchinson. Highly readable.

Huggett, R. J. (1993) Modelling the Human Impact on Nature: Systems Analysis of Environmental Problems, Oxford: Oxford University Press. If youlikemodellingbutarenottoomathematically minded, this might be of interest.

Reaka-Kudlka, M. L., Wilson, D. E., and Wilson, E. 0. (1997) Bzodiversity 11: Understanding and Protecting Our Biological Resources, Washington,DC:National Academy Press. Sure to become a classic.

EnvtronmentalChange:The Huggett,R.J.(1997) Evolving Ecosphere, London: Routledge. Provides a broad perspective.

Rosenzweig, M. L. (1 995) Speczes Dzversity zn Space and Time, Cambridge: Cambrldge Universlty Press. Fascinating, but not easy.

Jackson, A. R. W . andJackson,J.M.(1996) Environmental Science: TheNatural Enzsronment and Human Impact, Harlow: Longman. A basic textbook with several relevant sections.

Schultz,J.(1995) The Ecozones of the World:The Ecological Dtvzsions of the Geosphere, Hamburg: Springer. A verygoodsummary of theworld’smain ecosystems.

Matthews, J . A. (1992) The Ecology of RecentlyDeglaciated Terrain: A Geoecologrcal Approarh to Glacier Forelands and Prtmnary SuccesJton, Cambridge: Cambridge Universlty Press. An excellent case study and lots more.

Szaro, R. C. and Johnston, D. W. (1996) Biodiversity in Managed Landscapes: Theory and Practice, New York: Oxford Untversity Press. Try it.

Peters, R. L. and Love~oy, T. E. (eds) (1992) Global Warming and Biolopcal Dtuersity, New Haven, Connecticut and London: Yale University Press. A host of examples in this one.

Diversity of Life, Wilson, E. 0. (1992) The Cambrldge, Massachusetts: Belknap Press of Harvard University Press. A highly readable book.

Schwartz, M. (1997) Conservation z n Highly Fragmented LndslilpeJ’ New Chapman & Topical and highly recommended.

F U R T H E RREADING

Ethical issues and conservation This is a selection from a long list. Anderson, E. N . (1996) Ecologies of the Heart: Emotion, Belid and the Environment, New York: Oxford Unlversity Press. Botkin, D. B. (1990) Discordant Harmonies: A Neuj Ecology for the Twenty-First Century, NewYork: Oxford University Press. Committee onScientific Issues in theEndangered Species Act, National Research Council (1995) Science and the Endangered Specres Act, Washington,DC: National Academy Press. Cooper, N. andCarling, R. C. J. (eds) (1996) Ecologrsts and Ethical Judgements,London: Chapman & Hall. Norton, B. G. (1991) Toward Unity Among Envzronmentalists, New York: Oxford Universlty Press. Ryder, R. D.(ed.) (1993) AnimalWeyareand Environment, London: Duckworth and RSPCA.

the

Shafer, C. L. (1990) Nature Reserves: Island Theovy and Consenlation Practice, Washington,DCand London: Smithsonian Institution Press. Sylvan, R. andBennett,D. (1994) The Greenrng of Ethrcs: From Human Chauvinism t o Deep-Green Theory, Cambridge: White Horse Press; Tucson: University of Arizona Press.

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