Molecular Taxonomy and the Conservation of the Red Wolf and Other Endangered Carnivores Author(S): C

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Molecular Taxonomy and the Conservation of the Red Wolf and Other Endangered Carnivores Author(s): C. Alexander Brownlow Source: Conservation Biology, Vol. 10, No. 2 (Apr., 1996), pp. 390-396 Published by: Wiley for Society for Conservation Biology Stable URL: http://www.jstor.org/stable/2386855 . Accessed: 24/06/2013 16:30

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This content downloaded from 128.192.114.19 on Mon, 24 Jun 2013 16:30:12 PM All use subject to JSTOR Terms and Conditions MolecularTaxonomy and the Conservation ofthe RedWolf and Other Endangered Carnivores

C. ALEXANDER BROWNLOW" Yale School of Forestryand EnvironmentalStudies, 205 ProspectStreet, New Haven,CT 06511, U.S.A.

Abstract: Recentpublicationshave reaffirmedthat the red wolf(Canis rufus)isa hybridcof thecoyote and the gray wolf Besides the implicationsthese results uill likelyhave forfuture conservation /fJortsanld allotment of resourcesthrough the Endangered SpeciesAct for recoveryof the red wolf;it is likelythat broader consequences will be felt throughoutthe conservationcom;nu;iity as species come under the scrutinsyof el mor-e powerfulmeans of taxonomic identification.As molecuilartechnology is refinedin its abilityto resolvetaxo- nomic historiesand uncertainties,it is likelythat hybridization event(s) will be recognizedin more species. Thismay be ofparticular importancefor large carnivores,whose smallpopulationisizes mnakethemn suscepti- ble to bybridizationepisodes withclosely related, symnpatric species. Because of negativeperceptions, powerful antipredatoradvocates, conservationand resource constraints,anid an enigmatichybrid policy withinthe EndangeredSpecies Act, how red-wolftaxonomy is decided by the US. Fish and WildlifeService mnay affect the futureof large carnivoresin general.

Taxonomiamolecular y la conservaci6ndel lobo coloradoy otrasespecies de carnivorosen peligro Resumen: Publicaciones recienteshan reafirmado el estado taxon6mico del lobo colorado (Canis rufuss), identificandolocomo tin hibrido entrecoyote y lobo gris.Adem7is de las consecuencias qite estos resultados tendreinsobre los esfuerzosde conservaci6nfuturos y sobre la alocaci6n de recursos,a travis del Acta de Es- pecies en Peligro,para la recuperaci6ndel lobo colorado, es probable que se establescan consecuencias mas amplias a lo largo y ancho de la comnunidadconservacionista a medida que ncdsespecies caigan bajo el es- crutiniode metodospoderosos de identificaci6ntaxon6rmica. Es probable que, a mnedidaque la tecnologia molecular es refinadaen su habilidadpara resolverhistoriasy dudas taxon6micas,se reconozccanevenitos de hibridaci6n en mcisespecies. Esto puede ser particular7nenteimportante para los grandes carni'voros,cuyos tamafnospoblacionales pequefnoslos hacen susceptiblesa episodios de hibridaci6n coniespecies simpditricas estrechamenterelacionadas. Debido a percepcionesnegativas, a los poderosospartidarios anti de pred.adores, a las limitacionesde la conservaci6ny los recursosy Cl unanenigmcitica e hibridapolitica dentrodcel Acta de Especies en Peligro,laforma en que el Servici6nde Pesca y ViclaSilvestre resuielva la Taxonomniadel lobo colorado podria afectarelfuturo de los grandes carnivoroseni genieral.

Introduction fus), addingfuel to the continuingdebate over its taxonomic positionwithin the Canidae. The resultsof these Two recentlypublished articles (Wayne & Jenks1991; studies,all molecular-geneticin form,indicate unambig- Roy et al. 1994a, 1994b) have provided data further uouslythe hybridnature of thissouthern canid, empha- questioningthe species statusof the red wolf(Canis ru- sizingits genetic similaritiesto an ever-expandingcoy- ote (Canis latrans) population. They also have the potentialof placing a strainon a reintroductionprogram thathas been referredto by the U.S. governmentas one *Currentaddress for correspondence:2634 SE Kelly,Portland, OR of the "most significantsuccess stories"of the Endan- 97202. Act Fish and Service Paper submittedJanutary 23, 1995; revised inaniuscriptaccepted gered Species (ESA) (U.S. Wildlife April20, 1995. 1994) because an obscure "hybridpolicy" (O'Brien & 390

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Mayr 1991; Littell1992:21) will now likelybe called TheRed Wolf: Conservation and Controversy upon in an effortto delist the red wolf (Gittleman& Pimm 1991; Rennie 1991). The consequences of this Recognized as Canis niger as early as 1791 and first case maybe greaterthan currently understood because, identifiedscientifically in 1851 (Nowak 1979; Cohn as populationsget smallerdue to naturalor anthropic 1987), the red wolf (Canis rufus) historicallyranged factors,not only does the likelihood of hybridization throughoutthe southeasternUnited States into Texas amongcertain groups of animalsincrease, but the bene- and southernIllinois (Nowak 1979; Peek et al. 1991; fitsof hybridizationalso increase,often adding outside Phillips1991b) and was consideredabundant through- alleles to an alreadydepauperate genetic base. Giventhe out the westernpart of its rangethrough the firsthalf of ESA's historyof last-minutelistings and science's often this century(Peek et al. 1991). Perhaps numberingin last-minute,crisis-oriented conservation efforts (Soule the hundredsof thousands(Parker 1984), the red wolf, 1985), more populationsmay come to the point where likethe closelyrelated gray wolf (Canis lupus), was sub- hybridizationis a viable option for populationsurvival ject to the persecutionand loss of habitatthat accompa- (Cohn 1987; Parker1990; Lehmanet al. 1991). Further- nied the movementof European settlerswest, untilby more, to combat the demographicand genetic effects the 1930s it had been extirpatedeast of the Mississippi that small population size and subsequent inbreeding (Nowak 1979; Cohn 1987; Peek et al. 1991). The small can produce (O'Brien et al. 1985; Fergus1991; Packeret populationthat remained in the 1960s in the coastal ar- al. 1991), the deliberateintroduction of outsidegenetic eas of southerneasternTexas and southwesternLouisi- materialof closelyrelated species or subspecies maybe ana was subjectedto furtherstress in the formsof heavy utilizedby conservationofficials as a last-ditcheffort to parasiteloads and hybridizationwith an expandingcoy- preventspecies extinction(Avise 1989; Avise & Nelson ote (Canis latrans) population,a closely relatedcanid 1989). more adaptable to human landscape alterations.De- As molecular-genetictechnology becomes more so- claredan endangeredspecies in 1967 underthe ESA,the phisticated,allowing researchers to furtherresolve the red wolf numberedfewer than 100 by the time it re- taxonomicidentities of species of uncertainorigins, it is ceived priorityconservation measures under the 1973 likelythat many species and populationswill displayge- ESA (U.S. Fish and WildlifeService 1994). In the mid- netic materialindicating hybridization at some point in 1970s all remainingred wolves were capturedfor cap- the recent or distantpast. Many of these animalswill tivebreeding purposes before the species was lostto hy- have been historicallyconsidered conservationfocal bridization.By 1980 the red wolfwas consideredextinct points and standardbearers (O'Brien et al. 1990; Leh- in the wild (Cohn 1987; Meese 1989). man et al. 1991), and in our subsequentefforts to appro- Only 40 out of 400 wolves trappedwere considered priate efficientlyand effectivelythe limitedfunds pro- "pure" enough forbreeding purposes (Cauley in Cohn vided by the ESA,these historicallypopular species may 1987; Jenks& Wayne 1992), and the remaininganimals then be ignored.Carnivores, with historicallylow num- were destroyedbecause of infiltrationof coyotegenetic bers-made even smallerby yearsof directand indirect material.Of the 40 only 14 individualseventually made persecutionby humans-due to theirtop trophicposi- up the foundingpopulation (Cohn 1987; U.S. Fish and tion, potentiallyhave an increased likelihoodto inter- WildlifeService 1994). By 1984 the captive population breed cross-specificallyduring times of demographic numberedno more than 50 animals (Parker 1984). In stress. These factors,combined with likely calls for July1986 the U.S. Fish and WildlifeService (USFWS) delistmentby those who perceivepredators as a threat, proposed reintroductionof the red wolf into the re- have the potentialto change significantlyhow we ap- centlyobtained Alligator River National Wildlife Refuge proach carnivoreconservation. Using the red wolfas an (ARNWR)in coastal NorthCarolina (U.S. Fish and Wild- example, I discuss the potentialconsequences thatthe lifeService 1986a), and in Novemberof thatsame year combinedeffects of carnivorenatural history, public at- fourpairs were flownand releasedinto the refugeunder titudestoward carnivores, limited resources for conser- an "experimental"status that permits limited "taking" of vationefforts, destructive human activities, and superior the wolves under conditionsapproved by the USFWS taxonomicidentification technology may have upon fu- (Parker & Phillips 1991). In 1990 the Great Smoky ture carnivoreconservation efforts. I focus on carni- MountainsNational Park was consideredas a second re- vores in particularbut will discussother taxa forillustra- lease site (Parker1990). Bythe end of 1993 the red wolf tivepurposes. The definitionof "species"within the ESA populationhad grownto 233-247 individualswith 46- and used by conservationistsand those in decision-mak- 60 animalsin the wild and 187 in captive breedingpro- ing positionsis of obvious consequence to the current gramsaround the country(U.S. Fishand WildlifeService topic. But the role of the definitionhas been well-docu- 1994). Despite local fearsthat the wolfwould harmlive- mented elsewhere (O'Brien & Mayr 1991; Jenks & stock and domesticanimals (U.S. Fish and WildlifeSer- Wayne 1992; Hill 1993), and I will not commentupon it vice 1984, 1986a, b; Kellert 1986; Cohn 1987; Meese here. 1989;Rees 1989a,b; Phillips1991 b), throughthe use of

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This content downloaded from 128.192.114.19 on Mon, 24 Jun 2013 16:30:12 PM All use subject to JSTOR Terms and Conditions 392 MolecularTaxonomy and Endangered Species Brownlowt extensivepublic educationand outreachprograms, the was called forbefore any conclusions on red-wolftaxon- reintroductionhas been consideredexemplary by both omywould be considered.In theirrecent analysis of mi- conservationistsand USFWSand now servesas a model crosatelliteloci in coyotes and grayand red wolves, it for other reintroductionprograms (Phillips 1991a, appears thatRoy et al. (1994a) have providedthis miss- 1991b; U.S. Fishand WildlifeService 1986b, 1994). ing information,further confirming the red wolfs hy- C(onsidered by some to be the only wolf to have bridnature. evolved in NorthAmerica (Nowak 1979, 1992), the red wolfis of size and staturealmost exactly between that of coyotes and graywolves (Nowak 1979, 1992; Wayne 1992), a fact that had persuaded some (e.g., Goldman 1944) of its potentialto hybridizewith these two ani- MolecularTaxonomy and Endangered mals. It was, however,still considered a viable species. Species Conservation Not untilthe early1990s did moleculardata establishab- solutelythe hybridnature of the red wolf (Wayne & As moleculartechniques used in the resolutionof taxo- Jenks1991), puttinginto question its species status.But nomic historiesand eventsbecome more refined,their severalauthors cite anatomical(Nowak 1979, 1992), be- role in the conservationof floraand fauna threatened havioral,and ecologic (Phillips& Henry1992) factorsin withextinction will mostcertainly become of increasing theirsupport of the viable species statusof the red wolf. importance.May (1990) notes the overwhelmingimpor- Nowak (1979) combined skeletaland dental measure- tance of taxonomyand systematicsin conservationen- mentswith data fromthe fossilrecord to conclude that deavors,referring to these disciplinesas "bricks"from not onlyis the red wolfa viable species but it is perhaps which biological science is constructed.Methods such the ancestralstock from which both the coyoteand gray as polymerasechain reaction(PC,R) and otherDNA anal- wolfemerged. Likewise, Phillips and Henry(1992) note yses irrefutablyidentify andl clarify relationships among distinctivewolflike predatory and social behaviorswithin species, oftenwidely separated both taxonomicallyand theNorth Carolina red wolfpopulation, thereby reftiting geographically,including extinct taxa that exist only challengesto a red wolf-coyotehybrid hypothesis. withinmuseum collections (Arnheim et al. 1990; Roy et Recent molecularanalysis, however, may have estab- al. 1994b). Daughteryet al. (1990), studyingallozyme lishedbeyond a doubt the hybridnature of the red wolf differentiationbetween subpopulationsof the tuatara (Wayne & Jenks1991; Roy et al. 1994a, 1994b). Using (Sphenodon punctatus), including live and museum mtDNAanalysis from several extant graywolves, coy- specimens,discovered that the populationactually con- otes, and red wolves, as well as fromseveral museum sistedof two species and a single,possibly extinct, sub- specimens of red wolf, Wayne and Jenks(1991) con- species. Faulty taxonomy applied to S. punctatus histor- cluded that "the red wolf is entirelya hybridform or a icallyby nationalandl international conservationists and distincttaxon that hybridizedwith coyotes and gray decisionmakers resultedl in littleto no internationalpro- wolves over much of its previousgeographical range." tectionand the possible extinctionof several subspe- In particular,they note the unquestionablepresence cies. Only currentmolecular and genetic techniques and influenceof coyote geneticmaterial within the red- could have resolvedthe taxonomicidentities within S. wolf genome. The studysparked controversyas many punctatus, and they have thereforebeen fundamentalin called forthe immediatedelistment of the red wolffrom the establishmentof appropriateconservation efforts. the ESA,stating that the ftundsallotted for its reintroduc- Avise (1989) providlesfurther arguments for the use of tion(roughly $160,000 per yearfor the ARNWRpopula- moleculartechniques in the conservationof endangered tion; Phillips1991b) mightbe betterspent upon other, species. less questionableand more recognizablespecies (Gittle- It is extremelylikely that as more species are scruti- man & Pimm1991; Rennie 1991). Likewise,despite any nized throughmolecular means the identificationof hy- indicationthat the red wolfwas a threatto livestock,the brid eventsin the clistantor recentpast will be discov- studywas adopted by the AmericanSheep Industryin ered. Hybridlizationoccurs throughmany media, both theirpetition to the Secretaryof the Interiorto delistthe naturaland anthropic.C(ombined with introductionsof redwolf from the ESA (Phillips & Henry1992). The USFWS oftendominant domestic and/or game species (C ourte- denied the petition,remarking that it did not provide nay 1978), however,the continuingincrease in habitat substantialenough evidence to warrantdelisting (Henry loss and the resultantfragmenting of populationshave 1992). On the other hand, manyresearchers rallied in the potentialto increase significantlythe likelihoodof defense of the red wolf, noting the potentialbias in hybridization.This may be of particularimportance for Wayneand Jenks'sstudy and its analysisof mtDNAonly closelyrelated, sympatric carnivores with smallpopula- (Dowling et al. 1992a, 1992b; Nowak 1992; Phillips& tion sizes thatare the restultof ecologic and anthropic Henry1992), what Avise (1989) has controversiallyre- factors.Lehman et al. (1991) have discoveredthe intro- ferredto as "molecularchauvinism." Further research gression of coyote I)NA into gray-wolfpopulations in

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This content downloaded from 128.192.114.19 on Mon, 24 Jun 2013 16:30:12 PM All use subject to JSTOR Terms and Conditions Bi-ovroloii Mllolecularhlaonom)' antdE Edantigered cd$)ecies 393 the northernUnited States that have been reduced in activities that fuLrthermarginalize and fragment carni- numberto the point thatAllee effects(the inabilityto vore populations. findan appropriatemate) have now become apparent. The endangered Florida panther (Felis concolor coryi) Similarly,as red-wolfpopulations decreased to ex- was found to have a remarkablyhigh amount of abnor- tremelylow numbersin the southernUnited States, hy- mal sperm throughout its male population (O'Brien et bridizationwith coyoteswas facilitatedwhen appropri- al. 1990; Fergus 1991). Considered a subspecies of the ate mates could not be located (Parker 1990; Jenks& North American cougar, loss of habitat in the wake of in- Wayne 1992). Domestic pets also have a profoundef- creasing development has fragmented the Florida pan- fect, exemplifiedby two recentlypublished stuclies. ther population to a remnant of no more than 50 individ- Gottelliet al. (1994), researchingthe causal factorsbe- uals in the wild areas south of Lake Okeechobee. In a hind the endangermentof the Ethiopianwolf (Canis si- population of such small size the sperm abnormalities mensis,considered by manyto be the world's mosten- discovered by O'Brien et al. are not surprising. In addi- dangeredcanid) have discovereda dangerousinflux of tion to these abnormalities, however, the researchers genetic materialfrom the domestic dog (Cans Jatm/il- also discovered genetic material within the population iaris). Combinedwith continued habitat loss ancdreduc- that was not local in origin. It has since been discovered tion in population size, the hybridizationof the Ethio- that panthers from South America were released in the pian wolf may take it beyond conservation'srescue. Everglades by the I.S. Park Service in the 1950s and Similarly,declining populations of the Europeanwildcat 1960s in an effortto boost population size and decrease (Felis silvestris)in Britainare being inundatedby ge- the likelihood of genetic defects due to small population netic materialfrom local house cats (Hubbard et al. size (O'Brien & Mayr 1991; Fergus 1991). South Ameri- 1992), potentiallycompromising any ftitureconserva- can genetic material is now found in roughly 50% of the tion efforts.Instances of hybridizationdue to the com- population (O'Brien et al. in Fergus 1991; U.S. Fish and bined effectsof habitatdegradation and the intentional Wildlife Service 1991). But with potentially detrimental introductionof a more competitivespecies by, gener- genetic factors now found throughout the male popula- ally,short-sighted enthusiasts has led to the possible en- tion due to small population size and inbreecling,O'Brien dangermentof several troutspecies in the southwest and Mayr (1991) suggest that this influx of new genetic (Dowling & Childs 1992) and northwest(Leary et al. material may be of benefit to the population in combat- 1993) UnitedStates. ting potentially detrimentaleffects through the addition To combat the extremedemographic (Allee) and ge- of new alleles. netic effectsproduced by small population size, re- Demographic factors resulting from small population searchersmay be faced with the dilemmaof eitheral- size and the subsequent effortsat conservation through lowingthe introductionof genetic material from outside hybridization are perhaps nowhere better illustrated the (sub)species,thereby compromising the species-spe- than in the case of the Dusky Seaside Sparrow (Arnmo- cific"purity" of the endangeredpopulation, or allowing dracnuts nmaratimnitsnigrescens) of Florida's Atlantic the species to run its course to possible extinction. (Coast. Considered by many to be a melanistic subspecies Small population size often results in inbreedingbe- of the more common Seaside Sparrow (Amnmnodramus tween closely relatedindividuals, which then in many mnaratibnns)(O'Brien & Mayr 1991; Bean 1993), the instancesmanifests itself in reduced fitnessthroughout Dusky Seaside Sparrow underwent severe population the population as a whole. Unchecked,this loss of fit- decline during the 1950s and 1960s because of severe ness has the potentialto cause population extinction. loss of habitat. By 1980 only five individuals remained, Largefelids in particularappear to be at riskdlue to loss all males. In a last-ditcheffort to save the species, these of fitness.Packer et al. (1991), studyinga naturallyiso- individuals were brought into captivity and a breeding lated and increasinglyinbred population of lions in the program was established with females of the morpho- Ngorongorocrater in Tanzania,have discoveredsignifi- logically similar Scott's Seaside Sparrow (Ammnodramus cant levels of abnormalsperm throughout the male pop- miaratimrnspeninsulae) from Florida's gulf coast (Avise ulation.Similar findings have been recordedfor the Afri- & Nelson 1989; O'Brien & Mayr 1991; Bean 1993). The can cheetah(Acinonyxjubatus), which appears to have intentional hybridizationsubsequently denied the Dusky undergone an extreme population bottleneckseveral Seaside Sparrow protection under the ESA, and conser- thousandyears ago (O'Brienet al. 1985, 1987). Although vation effortswere eventually halted. The last male died the likelihoodof hybridizationin these two cases is ab- on 16 June 1987 and the subspecies were pronounced sent (the Ngorongorolion population is not a subspe- extinct. But post-mortem molecular data indicated that cies of Africanlion; the cheetah is the only species A. m. nigrescens was virtuallyidentical genetically to the withinits genus and thereforecannot naturallybreed more common Seaside Sparrow of the Atlantic coast and outsideof the species), theyclearly illustrate the natural clearly distinct from the gulf coast subspecies with demographicand genetic riskslarge carnivoresface in which it was bred, factors unknown at the time (Avise the wild. These risksare only compounded by human 1989; Avise & Nelson 1989).

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Perceptionof Carnivores Pimm (1991) and Rennie (1991) were quick to point thisout followingWayne and Jenks's(1991) conclusion Historically,carnivores have been perceivedas evil, de- of coyote-redwolf hybridization.Indeed, the articleby plorable creaturesin need of subordinationor eradica- Gittlemanand Pimmwas publishedin the samieedition tion(Lopez 1978; Kellert1980, 1986). Wolves,in partic- as the studyby Wayne and Jenks. ular,have been the subjectof oftenmisguided hatred in thiscountry (and aroundthe world),exemplified by our rigorous and successful effortsat eradication in the TheHybrid Policy of the Endangered Species Act lower 48 statesduring the nineteenthand earlytwenti- eth centuries(Lopez 1978; Kellert1986). But as public Althoughthe EndangeredSpecies Act has no formalpol- attitudestoward predators have evolvedto encompassa icy on hybrids,historical legislation dealing with hybrid more naturalisticand humanistic character (Kellert animalsappears to have been species-specificin nature 1980, 1986), manyof these species have become verita- (Appenzeller1989; Henry1992; Bean 1993). In the case ble icons in thiscountry's conservation movement. The of the DuskySeaside Sparrow,the intentionalhybridiza- bald eagle, grizzlybear, and graywolf all currentlyserve tion withScott's Seaside Sparrowresulted in its exemp- as powerfulsymbols in our effortsto conservethis coun- tion fromany federalconservation funding (Steinhart try's endangeredfauna (Kellert 1986, 1994). Further- 1986; O'Brien & Mayr 1991; Bean 1993). Accordingto more,they are now consideredby many conservationists the USFWS, "once the Dusky had been contaminated as "umbrella"species, whose large habitatrequirements with whatever genes, the biological entitythat was encompass the ranges and ecological requirementsof there is no more; . . the subspecies was extinct biologi- many more, smaller species; conservationof the one cally, in our view" (in Steinhart1986). In a not alto- promotesconservation of the whole. Nevertheless,be- getherdifferent case, however,the USFWS ruled in fa- cause of theirpredatory nature and expansivehabitat re- vor of the red wolf afterit had been petitionedfor quirements,conservation of these animalsis oftencon- delistmentby the American Sheep Industry(Henry sideredto run opposite manyinterests, particularly the 1992; Phillips& Henry 1992). Whetheror not this is a economicallyand politicallypowerful development and case of taxonomic prejudice remainsto be seen. The livestock industries.As molecular techniques are ap- two cases have been compared withinboth biological plied to moreof these predatory animals and the identifi- (O'Brien & Mayr 1991) and legal (Hill 1993) frame- cation of hybridizationevents is furtherresolved, these works. The conclusions of these respective studies, interestsare furtherarmed with data that may legally however,are verydifferent. Whereas O'Brien and Mayr benefittheir causes under an enigmatic"hybrid policy" (1991) call forthe redefinition,clarification, and subse- of the ESA (O'Brien and Mayr1991; Littell1992:21). In- quent consistentimplementation of the ESA's hybrid dustrysupport of the hybridpolicy, combined with the policy,Hill (1993) suggeststhe redefinitionof "species" negative perceptions that the public, in general, at- as applied by the ESA. taches to the term"hybird" (often mistaking it to mean "mongrel"or "interbreed";U.S. Fishand WildlifeService 1991), maybe of greatconsequence to futureconserva- Conclusion tion efforts.This has been illustratedin the case of the red wolf,which, upon publicationof itslikely hybrid na- As moleculartechniques with greater capabilities of tax- ture (Wayne & Jenks 1991), was petitionedfor delist- onomicresolution are producedand employedin our ef- ment fromthe ESA by the AmericanSheep Industry fortsto define conservationpriorities, it is likelythat (Phillips& Henry1992). It is likelythat recent publica- manymore species will be identifiedas havingexperi- tions by Roy et al. (1994a, 1994b), furtherquestioning enced a hybridizationevent(s) in theirrecent or distant the red wolf'staxonomic identity, will resultin renewed past. This maybe ofparticular importance to largecarni- effortsto delistthe animal.Similarly, removal of the gray vores whose numbershave been constrainedby both wolf fromthe ESA was soon sought by farmbureaus ecological and anthropicfactors. As a consequence of fromWyoming, Montana, and Idaho followingthe publi- the Allee effectsthat accompany reduced population cationby Lehmanet al. (1991) of wolf-coyotehybridiza- size, the potentialof hybridizingevents increaseswith tion data in the northernU.S. (O'Brien & Mayr 1991). increasedeffort to ensure gene flow and avoid extinc- O'Brienet al. (in Fergus1991) expressfear that develop- tion. These ftindamentaldemographic factors are often ment interestswill seek delistmentof the Floridapan- overlookedin our effortsto defineconservation foci on ther now that its hybridizationwith South American a genetic level (Lande 1988). Futurestudies of the ge- cougarshas been identified. netic makeup of largecarnivores may put furtherstrain Calls for delistmentcome fromwithin the scientific on conservationefforts if these animals, like the red wolf communityas well, as scientistsremind us ofthe limited and the Florida panther,are found to contain genetic fundsavailable for species conservation.Gittleman and materialof a nonspecificorigin. In the handsof interests

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This content downloaded from 128.192.114.19 on Mon, 24 Jun 2013 16:30:12 PM All use subject to JSTOR Terms and Conditions Brownlow Molecutlarhaxonom and Endangered Species 395 thatoppose carnivoreconservation, such data can be an brid (Wayne & Jenks 1991; Roy et al. 1994a, 1994b), awesome legal weapon. Furthermore,it adds additional and its taxonomic status will most likely once again be constraintsto the conservationmeasures necessaryto confronted and decided upon by the U.S. Fish and Wild- save those species and subspeciesundergoing severe ge- life Service. It will likely constitute a landmark decision netic and/ordemographic effects because of humanac- and, for many more species waiting in the wings, may tivities(U.S. Fish and WildlifeService 1991). As the hy- make the differencebetween survival and extinction. brid policy is now interpreted,once the Dusky Seaside Sparrowconsisted only of males all conservationmeasures were effectivelyfor naughtand the bird was al- Acknowledgments lowed to go extinct.Similarly, without genetic exchange from an outside source, the sperm abnormalities I thank Stephen Kellert, Tim Clark, and Dwight Barryof throughoutthe male Florida pantherpopulation may the Yale School of Forestry and Environmental Studies eventuallyrender this species extinct(O'Brien & Mayr for their help during the early stages of this manuscript. 1991; U.S. Fishand WildlifeService 1991). I also thank Russell Lande and two anonymous review- To deny conservationresources and effortsto these ers for their insight and helpful comments and sugges- species or populationsbased upon recentor distanth-y- tions. bridevents produces not onlyecological arguments(the loss of these species maylead to the loss of those organ- ismsfalling underneath their respective "umbrellas") but LiteratureCited also adds furtherto the ethical argumentof our ap- Appenizcller, T. 1989. A sparrow's fall. ScienltificAmericani 264:32-33. proach to endangeredspecies conservationin general. Arnheim, N., T. Wlhite,and W E. Rainey. 1990. Application of PCR: or- In the case of mostlarge carnivoresin the U.S., popula- ganismal and population biology. Bioscienice 40:174-182. tiondecline and fragmentationhas resultedfrom human Avise, J. C. 1989. A role for molecular genetics in the recognition and conservation of endangered species. Trends in Ecology and Evolol- activities.We are becomingincreasingly responsible for tio(n4:279-281. the fact that many species and populationsare in the Avise, J. C., and W. S. Nclson. 1989. Molecular genetic relationships of otherwiseunlikely and precarious position of mating the extinct DuLskySeaside Sparrow. Science 243:646-648. with closelyrelated yet more abundantspecies in order Bean, M. J. 1993. Where late the sweet birds sang. Natural Histoiy 102: to furthergenetic contributionand secure population 83-85. Cohn, J. P. 1987. Red wolf in the wilderness. Bioscience 37:313-316. survival,thereby reducing species-specific "purity." We Courtenay, Jr., W. R. 1978. The introdtuction of exotic organisms. are also responsiblefor the need to introduceoutside Pages 237-252 in H. P. Brokaw, editor. Wildlife and America: Con- genetic materialto allow population (species) persis- tribtutionsto an understanding of American wildlife and its conser- tence. Some argue that these are plausible groundsto vation. Couincil on Environmental Quiality,Washington, D.C. discontinueconservation efforts (e.g., Jenks& Wayne Daugherty, A. R., A. Cree, J. M. Hay, and M. B. Thompson. 1990. Ne- glected taxonomy and contintuingextinctions of tuatara (Spheno- 1992; but the authorsdo expressconcern over thissitu- don). Natture347:117-179. ation). Otherstate that, all else failing,flexibility should Dowling, T. E., and M. R. Childs. 1992. Impact of hybridization on a be permitedin such situationsto a certainextent (U.S. threatened trouLtof the souLthwesternUnited States. Conservation Fishand WildlifeService 1991). Biology 6:355-364. The questionsraised by theseissues are numerousand Dowling, T. E., B. D. Demarais, W. L. Minckley, M. E. Dotuglas, and P. C. Marsh. 1992ci. UJseof genetic characters in conservation biol- controversial.By depriving,through our activities,many ogy. 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