Biological Inventory Using Target Taxa: a Case Study of the Butterflies of Madagascar Author(S): Claire Kremen Source: Ecological Applications, Vol
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Biological Inventory Using Target Taxa: A Case Study of the Butterflies of Madagascar Author(s): Claire Kremen Source: Ecological Applications, Vol. 4, No. 3 (Aug., 1994), pp. 407-422 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1941946 . Accessed: 07/02/2014 15:59 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecological Applications. http://www.jstor.org This content downloaded from 128.32.85.74 on Fri, 7 Feb 2014 15:59:56 PM All use subject to JSTOR Terms and Conditions Ecological Applications,4(3), 1994, pp. 407-422 ? 1994 by the Ecological Society of America BIOLOGICAL INVENTORY USING TARGET TAXA: A CASE STUDY OF THE BUTTERFLIES OF MADAGASCAR' CLAIRE KREMEN2 Centerfor ConservationBiology, Stanford University, Stanford, California 94305 USA Abstract. Biotic inventoriesprovide criticaldata forconservation planning, but fre- quently,conservation decisions are made withoutsurveys, due to lack of time, funds,or appropriatemethodology. A method,target taxon analysis, is thereforeproposed for stream- liningregional biotic inventories, while simultaneously increasing their taxonomic coverage and spatial resolution.In this method,regional inventories focus on a numberof narrowly definedtarget taxa, chosen to representcollectively an array of highertaxa. Such target taxa should be informationrich; in other words, the patternof species distributionsin these taxa should correlateeither with patternsof environmentalheterogeneity or with distributionalpatterns of species in unrelatedtaxonomic groups. It is suggestedthat clades that experiencedan evolutionaryradiation withinthe regionare likelyto be information richfor conservation planning at or withinthis regional scale. Such clades willbe identifiable as low-ranking,species-rich taxa with highendemism. The informationrichness of these potentialtarget clades can then be evaluated by directgradient methods of analysis that relate communitycompositional change to environmentalfactors, or by correlatingdis- tributionalpatterns of species among separate targetclades. To assess this approach to biological inventory,a species-richgenus and subgenusof endemic butterfliesfrom the island of Madagascar were chosen as targettaxa and were evaluated forinformation richness in comparison to the entirebutterfly fauna of Mada- gascar. Using canonical correspondenceanalysis and otheranalytical techniques, the sub- genus of Malagasy Henotesia species (Satyrinae)proved to be as good or betterthan the entirebutterfly fauna at delineatinga varietyof environmentalgradients at both local and landscape scales. The endemic genus Strabena (Satyrinae)was only able to delineate such patternsunder a restrictedset of conditions. However, this genus, while species rich in Madagascar, was not exceptionallydiverse nor were its species membersabundant within the study area. It is concluded that targettaxon analysis is a potentiallyuseful tool for providinghigh-quality data while expandingcoverage of taxonomic diversityfor conser- vation planning. Key words: area of endemism;biodiversity inventory; biogeography; biological indicators;conser- vationplanning; environmentalgradient; Madagascar; ordination;species richness;species turnover. INTRODUCTION or that are solicited by planners are of limited taxo- To set aside representativeareas around the globe nomicscope: birds,mammals, and some vascularplants thatwill protectthe fullestrange of biologicaldiversity (Murphyand Wilcox 1986, di Castriet al. 1992). Many requires far more complete knowledge of the distri- taxonomicallydiverse groups that could provide more bution and abundance of organismsthan is currently fine-graineddatabases have been ignored(Collins and available (Soule and Kohm 1989, Lubchenco et al. Thomas 1991). The biggeststumbling block preventing 1991). In the past, boundaries of naturereserves were inclusion of a wider variety of taxa in biodiversity frequentlyestablished withoutreference to biotic in- inventoriesis our greatlack of taxonomic knowledge; ventories(Terborgh and Winter1983, Scottet al. 1987, only 1 700 000 species are described (Wilson 1985, Fa 1989). Increasingly,however, conservation organ- Vane-Wrightet al. 1991), while estimates of species izations and governmentalagencies are asking forbi- diversityrange from 10 to 80 x 106 (Endler 1982, ological surveydata to use in makingdecisions about Erwin 1982, 1988, 1991b, May 1988, Stork 1988, Gas- which areas to protect(e.g., the Manaus Conference: ton 1991). The vast majorityof species are inverte- Kuliopulos 1990, Prance 1990; see also Jenkins1985, brates (70%) (Wilson 1988). Noss 1987, Thomas et al. 1990). Given the tremendousnumber of species and our Frequently,the only surveydata that are available tremendousignorance of them, practicalityprohibits an exhaustive inventorywithin a time framerelevant to conservation [see estimates in Wilson (1985) and I Manuscript received 31 August 1992; revised 29 April 1993; accepted 1 June 1993. Wheeler (1990) forestimated lengthof time to com- 2 Present address: The Xerces Society, 10 SouthwestAsh plete a total inventory].By necessity,inventories must Street,Portland, Oregon 97204 USA. rely on focal or indicatortaxa (di Castri et al. 1992). This content downloaded from 128.32.85.74 on Fri, 7 Feb 2014 15:59:56 PM All use subject to JSTOR Terms and Conditions 408 CLAIRE KREMEN Ecological Applications Vol. 4, No. 3 Rather than selectingindicators on an ad hoc basis, phries and Parenti 1986, Cracraftand Prum 1988, inventories for conservation planning should target Myersand Giller 1988). However,given that congruent taxonomic groups that are biogeographicallyinfor- patternsabound, the processes that produced them need mative,relatively well-characterized taxonomically, and not be determinedin order to develop an efficientin- easy to measure or observe (Noss 1990, Pearson and ventorystrategy for conservation. Selecting clades that Cassola 1992). Biogeographicallyinformative taxo- radiated withinthe regionof interestas biogeographic nomic assemblages will delineate areas of endemism, probes will ensure that areas of endemism can be de- centersof diversity,and unique communitiesor hab- tected, whetherthey result from vicariance or other itats-informationof great relevance to conservation processes. planners. Species-richclades thathave radiatedwithin a region The purpose of this paper is to suggesta set of bi- mayalso be good indicatorsof the environmental com- ological criteriathat can be used forselecting biogeo- plexity (habitat and microhabitatvariation) present graphicallyinformative taxa forinventories. I present withina region(definition 2). As species derived from a generallyapplicable method for testingthe infor- a common ancestor,clades will tend to consist of spe- mation richnessof targettaxa and illustratethis ap- cies thatare relativelysimilar ecologically (Brown 1984, proach with surveydata on butterfliesfrom a region Brooksand McLennan 1991). Wherethey exist in sym- of Madagascar. patry,species fromhigh-diversity clades may frequent- ly be segregatedon a fine spatial or temporal scale. Selection criteriafor choosing Examples includeecological replacementseries by con- information-richtarget taxa genersalong elevational, latitudinal,or habitat gradi- A taxon is definedhere as biogeographicallyinfor- ents,and nicheor resourcepartitioning (Heinrich 1976, mative if, in a given geographicarea, analysis of the McClure and Price 1976, Adams and Bernard 1977, patternof species distributionsin thisassemblage cor- Southwood 1978, Grant 1981, 1983). Again,regardless relateswith spatial patternsof: (1) species distributions of mechanism (e.g., competitiveexclusion, character of unrelated taxonomic groups, and/or (2) environ- displacement, species packing, etc.), the diversityof mental heterogeneity.Such taxa will then serve as in- species that are partitionedwith respectto habitat or dicatorsof geographicallyreferenced ecological or bio- microhabitatcan be an indicatorof the level of envi- diversitycharacteristics of a region(see also Noss 1990). ronmentalheterogeneity present in an area. For ex- I hypothesizethat clades (monophyleticassemblages ample, Murdoch et al. (1972) foundthat plant-feeding of species) that have experiencedan evolutionaryra- Homoptera diversitywas positively correlated both diation within a region are likely to be information withplant species diversityand vegetationcomplexity; rich,as definedabove, forregional conservation plan- fora review see Lawton (1978). In seeminglyhomo- ning.Such clades will be recognizedby theirhigh spe- geneous habitats, transitionalzones of high species cies diversityand endemism withinthe region of in- turnoverfor the membersof the clade may demarcate terest.The rationaleto supportthis basis forselecting subtlydistinct community or habitattypes. Even con- informativetarget taxa is as follows. tinuous variation in distributionsof members of the The patternsof species distributionsof two or more clade may identifyotherwise