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Magurran y McGill (1)

Magurran y McGill (1)

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Published by: Maria Jesus Sutta on Jun 15, 2012
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Challenges and opportunitiesin the measurement and assessmentof biological diversity
Anne E. Magurran and Brian J. McGill
‘When we look at the plants and bushes clothing anentangled bank, we are tempted to attribute their pro- portional numbers and kinds to what we call chance.But how false a view is this! Every one has heard thatwhen an American forest is cut down, a very differentvegetation springs up; but it has been observed thatancient Indian ruins in the Southern United States,which must formerly have been cleared of trees, nowdisplay the same beautiful diversity and proportion of kinds as in the surrounding virgin forests.’
—– Darwin, Origin of Species (1859)
1.1 Introduction
By the 1800s naturalists had begun to document afactthatwassurelyapparenttotheearliesthumans:that species vary markedly in how common theyare. Audubon, for example, noted that bird speciesin North America differ in abundance by up toseven orders of magnitude (McGill 2006; McGilletal.2007).Darwin’s(1859)reflectionsontheobser-vation that some taxa are abundant yet many arerare, and that there are geographical differences inthe abundances and types of species, shaped histhinking about natural selection. In drawing atten-tion to the ‘proportional numbers and kindsofspecies in nature, however, Darwin also identifiedquestions that continue to challenge ecologists acentury and a half later.Humans have a strong intuitive sense of the dis-tinction between high diversity and low diversityassemblages, as encapsulated, for example, by thecontrast between a tropical rain forest and a mono-culture planted for timber. Moreover, the inequalityof species abundances that Darwin reported isso pervasive as to have acquired the status ofan ‘ecological law’ (McGill et al. 2007). However,quantifyingthebiologicaldiversityofacommunity,assessing differences in this diversity over space ortime, and doing so in ways that are useful to thosewho seek explanations for the natural patterns aswell as to the managers and policy makers chargedwith the sustainable use of wild nature, is by nomeans as simple as it first appears. The reason forthis difficulty is that biological diversity is a mul-tifaceted concept that can be defined and docu-mented in different ways. Being clear about exactlywhat we mean by biological diversity (or biodiver-sity) is the first step towards measuring it. Eventhen the user can be confronted by a myriad mea-sures,someofwhichwilldoabetterjobthanothers.The goal of this book is to guide readers throughthe entangled bank of biodiversity measures andprovideanup-to-dateandaccessibleaccountofthisimportant and rapidly expanding field.
1.2 State of the field
2010markstheUnitedNationsInternationalYearofBiodiversity (UN IYB), an initiative that highlightsthegrowingneedtoprovideinformativeandrobustassessments of biological diversity. The objectivesof the IYB are to promote the protection of biodi-versity and encourage organizations, institutions,companies, and individuals to take direct actionto reduce the constant loss of biological diver-sity worldwide. However, 2010 is also the targetdate set by the 2002 World Summit on Sustainable
Development ‘for significant reduction in the cur-rent rate of loss of biological diversity’. It is by nowgenerally accepted that the 2010 target will not bemet, and of course there are many reasons for this,one of which is the ongoing debate about how bestto measure biological diversity.The measurement of biological diversity has res-onances with the parable of the blind men andthe elephant where one felt the trunk, one an ear,one a tusk and another a leg, only to find thatthey disagreed about the nature of the creature theyhad encountered. Quantifying biological diversityis in some ways like trying to measure an elephant(Nanney 2004). An elephant might be described interms of its morphology or behaviour or neural cir-cuitry, or DNA. Gene expression might be trackedthrough time or compared amongst individuals.The social interactions in different herds might also be explored. All of these approaches are valid, butclearly different. The investigator needs to spec-ify which aspect of the elephant is being assessed,and then make the case that the types of measuresadopted are employed correctly and are suitablefor the purpose for which they are bring used.In the same way that there is no single methodthat entirely captures the essence of elephantness,so there is no one metric that perfectly quantifies biological diversity. A large, and growing, num- ber of diversity measures have been developed(Southwood&Henderson2000;RoyalSociety2003;Magurran 2004). Southwood (1978) (p. 421) wrylynoted the ‘explosive speciation’ of diversity indicesand the habit of inventors of new measures to con-demn their predecessors. Biodiversity assessmentembraces not just the classical measures of richness,evenness, and species abundance distributions butalso evaluations of occurrence, range size, and vul-nerability as well as functional traits and phyloge-netic diversity. It is clear that there can be no single‘best buy’ index that will suit all needs. Instead,clusters of measures have been developed to cap-ture a certain aspect of biodiversity, with someof these performing this task better than others.Although it may seem like a disadvantage to haveto choose amongst a range of metrics, the ability toexamine biological diversity in different ways notonly helps ecologists gain a deeper understandingof how ecosystems function, but also sheds light onissues of practical concern such as the link betweendiversity and ecosystem services.The quantification of biological diversity proba- bly began with Darwin, who in 1855 recorded allthe plants in the meadow, Great Pucklands, nextto his family home at Downe. His list ran to 142species. It was over 50 years before the first accountof the relative abundance of species was pub-lished by Raunkaier in 1909. Other early, yet piv-otal, papers by Motomura (1932) and Fisher (Fisheret al. 1943) laid the foundations for the investiga-tion of species abundance distributions. Additionalinsightful and significant contributions (includingthose by Preston (1948), MacArthur (1960), May(1975), Pielou (1975), Taylor et al. (1976), and Sug-ihara (1980) followed. These provide the frame-work within which biodiversity is measured, andstill repay careful reading. Two developments thatoccurred towards the end of the twentieth century,however, shifted the measurement and assessmentof biological diversity to a new gear. The first ofthesewastherecognitionthatbiologicaldiversityisa crucial global resource as well as one that is beinglost at an accelerating rate, the second the advancesin computing power that have fostered substan-tial improvements in statistical and modellingtechniques.
1.3 What is in this book
Although vastly improved analytical and comput-ing resources are of tremendous benefit to ecolo-gists, the rapid development of new methods, theinconsistent and sometimes confusing applicationof old ones, and the lack of agreement in the liter-ature about the best approach means that users can be bewildered. This book provides an up-to-dateaccount of the methods used to measure and assess biological diversity and places particular empha-sis on the practical issues involved in measure-ment. It extends the discussion in Magurran (2004)and includes many new developments as well as are-examination of familiar approaches. Our focus ismeasurement of the variety, abundance, and geo-graphical occurrence of taxa, reflecting the sensein which Darwin used the term ‘diversity’. Theauthors have wide experience of the measurementof biological diversity and have contributed to
recent advances. As befits a vibrant field, our con-tributors do not invariably agree on which metricsand approaches are the best ones, although there is broadconsensusabouttheessentialsofbiodiversityassessment. The various chapters set out the issuesthat investigators must consider and explain theadvantages and disadvantages of different meth-ods, thus helping the reader to reach a sensible con-clusion about the best course of action in their ownparticular study system.The book is primarily aimed at those who needto measure and assess biological diversity. We havein mind upper-level undergraduates engaged inresearch projects, graduate students, and postdoc-toral researchers, as well as environmental man-agers and conservation biologists. Although notdesigned as a theoretical ecology text, we nonethe-less hope that the issues raised in the book willspark the interest of modellers and theoreticians. Ineach chapter the author(s) describe the state of thefield, discuss recommendations and future direc-tions, and end with key points.The book begins with an overview of the basicmeasurement issues involved in biodiversity stud-ies. As Scott Bonar, Jeffrey Fehmi, and NormanMercado-Silva make plain, survey design andmethodology play a crucial role in the success—or otherwise—of biodiversity investigations. Theseare well-known concerns, although they often getmuch less attention than they deserve; Chapter2sets out the points that must be addressed beforeaninvestigationcanevenbegin.IncontrastStephenBuckland, Angelika Studeny, Anne Magurran, andStuart Newson highlight a problem that is poten-tially of great importance but one that has, untilnow, been almost entirely overlooked. Most diver-sity statistics proceed on the assumption that indi-viduals and species have been collected at randomfrom the community of interest and most investiga-tors use these statistics irrespective of whether thisassumption is fulfilled. In practice individuals andspecies often vary considerably in how easy theyare to detect. Buckland et al. show how detectabil-ity can affect the conclusions drawn from diversitystatistics, and provide advice on how to deal withthe issues raised, although it is clear that assessingindividual and species variation in detectability iseasier for some taxa than for others.The next section deals with the approaches thatare sometimes described as measures of ‘speciesdiversity’. Although the term ‘biodiversity’ canmean many things, and is sometimes used in ageneric way (EASAC 2009), most scientists, man-agers, and policy makers identify species richnessas a central component. In many ways speciesrichness—the number of species in a given local-ity or assemblage—is the iconic measure of bio-logical diversity. It is used to identify biodiver-sity hotspots and plays an important role in con-servation planning. Species richness also accordswell with our intuitive sense of biological diversity.However,despiteitswideappealandapparentsim-plicity, accurate estimates of species richness can beremarkably difficult to achieve. This is not just atthe global level, where it is still unclear, to withinat least an order of magnitude, how many speciesinhabit the planet, but also at local levels andeven for taxonomically well-characterized organ-isms. The reason for this difficulty lies in the obser-vations of Darwin and Audobon—because mostspecies are rare, an increase in sampling effort willalmostalwaysleadtoanincreaseinrichness.Fortu-natelytherehasbeenintenseinterestinrecentyearsin providing solutions to this problem; NicholasGotelli and Robert Colwell provide a clear accountof the approaches that can be used to make faircomparisons amongst sites where sampling effortdiffers (which can happen even if the investigatorthinks that it is the same) and examine the newgenerationofnon-parametricestimatorsthatcanbeused to deduce the minimum number of speciespresent in an assemblage.Although species richness is widely used as ameasure of biological diversity, investigators oftenwant to find a means of quantifying Darwin’s ‘pro-portional numbers and kinds’ in a single statis-tic. As noted at the beginning of this chapter allcommunities consist of species that vary in theirabundance. However, communities (or localities)can also differ from one another in terms of theirproportions of species, that is, in how ‘even’ theirspecies abundance distributions are. The degreeof evenness can shed light on the processes thatshape a community’s structure or provide a gaugeof the impacts on it. There are a swathe of measuresthat populate the ground between species richness

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