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In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane G.; Collins, Loa C., comps. 2013. Merging science and management in a rapidly changing world: Biodiversity and management of the Madrean Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station.

Biodiversity in the Madrean Archipelago of Sonora, Mexico

Thomas R. Van Devender, Sergio Avila-Villegas, and Melanie Emerson Sky Island Alliance, Tucson, Arizona

Dale Turner The Nature Conservancy, Tucson, Arizona

Aaron D. Flesch University of Montana, Missoula, Montana

Nicholas S. Deyo Sky Island Alliance, Tucson, Arizona

Introduction Floweryrhetoricoftengivesbirthtonewtermsthatconveyimagesandconcepts, leadtoinspirationandinitiative.Onthe1892-1894expeditiontoresurveytheUnitedStates-Mexicoboundary,LieutenantDavidDuboseGaillarddescribedtheArizona-Sonoraborderlandsas“bare, jagged mountains rising out of the plains like islands from the sea” (Mearns1907;HuntandAnderson2002).LaterGalliardwastheleadengineeronthePanamaCanalconstructionproject. In1951,WeldonHeald,aresidentoftheChiricahuaMountains,coinedtheterm‘SkyIslands’fortherangesinsoutheasternArizona(Heald1951).FrederickH.Gehlbach’s1981book,Mountain Islands and Desert Seas: A Natural History of the US-Mexican Borderlands, providedanoverviewof thenaturalhistoryof theSky Islands inthesouthwesternUnitedStates.The‘desertseas’inthevalleysaredesertgrasslandandChihuahuandesertscrubinthenorth,Sonorandesertscrubtothewest,andfoothillsthornscrubinthesouth.VariantsofthetermincludeSkyIslandregion(McLaughlin1995;Gottfriedandothers2005;Skroch2008),sky-islandranges(FelgerandWilson1995,Fishbeinandothers1995),andskyislandbioregion(Fishbeinandothers1995;Skroch2008).Todaywemostlyuse‘SkyIslandRegion’withcapitalletters.ThebestnameinSpanishisIslas Serranas,ratherthantheliteraltranslationIslas del Cielo,becausecielo meansboth‘sky’and‘heaven.’Theanalogytooceanicislands(Warshall1995)islimitedbecauseSkyIslandsdifferfromtrueinsularareasinhighspeciesdiversity,lowlocalandregionalendemism,andlowpercent-agesofnon-nativespecies(McLaughlin1995).Whileanyisolatedareaisapotentialareaforspeciationinsmallpopulations,therearerelativelyfewspeciesrestrictedtotheSkyIslandMountains. Theterm‘Madrean’ comesfromtheSierraMadre.TheMexicanPlateau is a vast area of grasslands and desertscrub between theRockyMountainsinNewMexicoandca.1,300kmtothesouththeTrans-MexicanVolcanicBeltinsouth-centralMexico.ThePlateauis

opentoincursionsoffrigidArcticairfromthenorth,andtheSierraMadresOriental andOccidental create adouble rain shadowandtheChihuahuanDesert.MadreanisageneraltermusedtodescribethingsrelatedtotheSierraMadres.InabiogeographicalanalysisoftheherpetofaunaofSaguaroNationalMonument,UniversityofArizonaherpetologistandecologistCharlesH.Lowewasprobablythefirsttousetheterm‘MadreanArchipelago’todescribetheSkyIslandrangesbetweentheSierraMadreOccidentalinSonoraandChihuahuaandtheMogollonRimofcentralArizona(Lowe,1992).Warshall(1995)andMcLaughlin(1995)expandedanddefinedtheareaandconcept.

Biodiversity In2007,ConservationInternationalnamedtheMadreanPine-oakWoodlandsasaglobalbiodiversityhotspot.ThiswasaverylargeareathatincludedtheSierraMadreOrientalineasternMexico,theSierraMadreOccidental(SMO)inwesternMexico,andtheSkyIs-landrangesnorthofthemintoTexas,NewMexico,andArizona.TheSierraMadreOccidentalextendsinwesternMexicofromZacatecasandJalisconorthtoChihuahuaandSonora.ThesouthwesternUnitedStatesisfamousforitsdiversityofanimalsandplants,andinmanygroupsdiversityincreasessouthwardinSonoraintotheSierraMadreOccidentalandtheNewWorldtropics. SpeciesrichnessisenhancedintheSkyIslandRegionformanyreasons.ThewesternhalfofNorthAmerica ismountainous,withtopographicallydiversehabitatsandvegetationzonedalongeleva-tional-climatic gradients. Biotic communities are concentrated insmallgeographicareascomparedtotheeasternUnitedStateswherevegetationmostlychangesalonganorth-southlatitudinalgradient. TheMadreanArchipelagoisaconvergencezoneforfivebioticprovinces(fig.1).TheRockyMountainsandColoradoPlateaustothenorthhavetemperateclimates.Inthemid-continent,thegrasslandsoftheGreatPlainsextendfromCanadasouthtoNewMexico,Texas,andtheMexicanPlateau.Inthesouth,amosaicwithdesertgrasslandinthevalleysandChihuahuandesertscrubonadjacentlimestoneslopesextendswestwardintosoutheasternArizonaandnortheasternSonora.OnthewesternedgeoftheSkyIslandRegion,SonorandesertscrubsurroundsSkyIslandranges.Inthesouth,moretropicaloakwood-landsandpine-oakforestsareintheSierraMadreOccidentalandSkyIslandmountaintops.Inthelowlands,foothillsthornscrub(FTS)andtropicaldeciduousforestarepresent.ThetransitionbetweentheNewWorldtropicsandthenortherntemperatezoneisatabout29°Nin

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Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

east-centralSonora.ThenorthernlimitsofFTSinSonoraareatabout30°11’NeastofSinoquipeintheRíoSonoraValleyand30°26’NatPresaAngosturaontheRíoBavispeatthesouthernendoftheSierraElTigre.FTSdoesnotreachArizona,butthedistributionsofquiteafewFTSspeciescrosstheborderintosouthernArizonaindesertgrasslandoroakwoodland.

Sky Island Bioblitzes Todaytheterm‘bioblitz’ispopular,meaninganintensiveeffortinashortperiodtodocumentthediversityofanimalsandplantsinanarea.ThefirstbioblitzinthesouthwesternUnitedStateswasthe1848-1855surveyofthenewboundarybetweentheUnitedStatesandMexicoaftertheTreatyofGuadalupeHidalgoof1848endedtheMexican-AmericanWar.TheborderbetweenElPaso,Texas,andtheColoradoRiverinArizonawassurveyedin1855-1856,followingtheGadsdenPurchasein1853.Besidessurveyingandmarkingtheborderwithmonuments, theseexpeditionsmadeextensiveanimalandplantcollections,oftenbyarmyphysicians.BotanistsJohnM.Bigelow,CharlesC.Parry,ArthurC.V.Schott,EdmundK.Smith,GeorgeThurber,andCharlesWright(fig.2)andzoologistJohnH.ClarkcollectedbetweentheSantaRitaCopperMines(=SilverCity),NewMexicoandSantaCruz,SonorainthemodernArizona-Sonoraborderlands.OnthesecondUnitedStates-MexicoBoundarySurveyin1892-1894,EdgarA.Mearns(fig.3)collected30,000animalandplantspecimens.ThemammalswerepublishedinthebookMammals of the Mexican Boundary of the United States (Mearns1907). In1890,NorwegianCarlS.LumholtzledananthropologicalexpeditionthroughnortheasternSonora.Hepublishedapopulartraveloguein1902.Sweden’sCarlV.Hartmanwasabotanistinchargeofgeneralbiologicalcollections(fig.4).

Río Mayo-Yécora Baseline

Inthe1930s,HowardScottGentrywasarecentgraduatefromtheUniversityofCaliforniaatBerkeley.Insearchofemployment,hewenttotheNewWorldtropicsintheÁlamosareainsouthernSonora,inhopesofsellingbiologicalspecimenstomuseumcollections.His

plantcollectionswerepublishedinthebookRío Mayo Plants(Gentry1942).Inthe1970s,80s,and90s,PaulS.MartinledexpeditionsfromtheUniversityofArizona,whichretracedGentry’stravels,explorednewareas,andresultedinarevisionoftheRíoMayoflora(Martinandothers1998).ThislargeareaincludestheSierraMadreOccidentalineasternSonoraandwesternChihuahua. StudiesofthefloraandfaunaoftheYécoraareaintheSMOineastern Sonora provide baseline information needed to comparebiotasintheMadreanArchipelagowiththemainlandSierraMadreOccidental,puttingthebiodiversityofSkyIslandrangesinregionalcontext.MexicanFederalHighway16(MEX16),oneofthefewhighwaysthatcrossestheSMO,providesawest-to-eastelevationaltransectfromtheRíoYaqui(180m)easttotheChihuahuaborder,42kmeastofYécora(fig.5).

Figure 2—(A) The U.S.-Mexico Boundary and Gadsden Purchase Surveys of 1848-1857 went through Guadalupe Canyon (photo by T. R. Van Devender). (B) Arthur C.V. Schott and (C) Charles Wright were expedition botanists.

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Figure 3—(A) The second U.S.-Mexico Boundary Survey in 1892-1894 was led by Edgar A. Mearns. (B) The Arizona-Sonora Boundary is between the Sierra San José and the Huachuca Mountains (photo by T. R. Van Devender).

Figure 1—Map of the biotic convergence in the Madrean Archipelago (map by Nick Deyo).

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Van Devender, Avila-Villegas, Emerson, and others Biodiversity in the Madrean Archipelago of Sonora, Mexico

deYécoraregionon36fieldtripsfrom1995to2008.Thisflorawith1,776taxain3,300km2isverydiverse,ca.30%morediversethantheHuachucaMountainsinsoutheasternArizona(Reina-G.andVanDevender2005).Thereare2,125observationsof301speciesofbirdsfromtheMunicipiodeYécoraintheMadreanArchipelagoBiodiversityAssessment(MABA)database(http://www.Madrean.org).

Borderlands Exploration

HigherSkyIslandmountainrangessuchastheSierrasdelosAjos,Elenita, Marquita, and Tigre have extensive pine-oak forest andmixed-coniferforestsonthehighestpeaks(fig.6;Marshall1957).ThebiotasoftheseareasaretemperatemixturesofSierraMadreanandRockyMountainspecies. Inthe1930sand1940s,theUniversityofMichiganhadaveryactivebiologicalexplorationprograminMexico.In1935,BarryCampbellcollectedamphibiansandreptilesintheSierraelTigre,Sonora.From1938-1941,StephenS.Whiteledthreebotanicalexpeditionstoex-ploretheRíoBavisperegionofnortheasternSonora(fig.7).TheRíoBavispeflorawith1200species(currently995taxaafterrevision)in549generain114familiesfromawiderangeofhabitatsintheregionwasthefirstmodernfloraintheSonoranborderlands(White1948). Ornithologist and intrepid explorer JoeT.Marshall studied thebirdsanddominantplantsofpine-oakwoodlandinmountainrangesin southeasternArizona,northeasternSonora, and in thenorthernSierraMadreOccidentalinChihuahuainthesummersof1951-1953

Figure 4—The Lumholtz Expedition. (A) Carl S. Lumholtz. (B) Botanist Carl V. Hartman. (C) Sketch of the church in Bacadéhuachi in Lumholtz (1902).

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Figure 5—Sunset in the Sierra Madre Occidental west of Yécora, Sonora (photo by Erik F. Enderson).

Between1953and2005,herpetologist-ecologistCharlesH.LowecollectedamphibiansandreptilesintheYécoraareafortheUniver-sityofArizonaHerpetologicalCollectionon33tripsinvolving26people.From2004to2008,ErikF.EndersonandRobertL.BezydocumentedtheYécoraareaherpetofaunathroughphotographyon40fielddays.VanDevenderandAnaL.Reina-GuerreroattheArizona-SonoraDesertMuseum(ASDM)studiedthefloraoftheMunicipio

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Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

(Marshall,1957).InSonora,heworkedintheSierrasAconchi,Azul,“Cananea”(=Elenita-Mariquita),Oposura(=LaMadera),andPinitos. Beginningin1963,RaymondM.Turner,J.RodneyHastingsandlater Tony L. Burgessmade numerous trips to Sonora and Baja,California,todocumentthedistributionsof414speciesofplantsintheSonoranDesert.Theirobservations,collections,andecologicalsummarieswere included in thebookSonoran Desert Plants. An Ecological Atlas (Turnerandothers1995).

Beginningin1964,StephenM.RussellandGaleMonsonwenttoSonora toobservebirdson150 tripsand692fielddays.Theirobservationsof525specieswerepublishedinthebookThe Birds of Sonora(RussellandMonson1998).Beginningin2000,AaronD.FleschexpandedontheirfieldobservationsinnorthernSonora(Flesch2008). Inthe1980sand1990s,theUnitedStatesFishandWildlifeService,ÁreaNaturalProtegida(ANP)Ajos-Bavispe(aComisiónNacionaldeÁreasNaturalesProtegidas[CONANP)reserve),UniversityofArizona,andTheNatureConservancyPlantCollectionsorganizedexpeditionstotheSierradelosAjosjusteastofCananea:FrankW.Reichenbacherin1982,PaulS.Martinin1983and1991,PeterWarrenandEstherSaucedoin1989,andRichardS.Felger,MarkFishbein,George M. Ferguson, Dale S. Turner, James Malusa, FlorentinoGarza-S.,andMartínHaroinOctober1992andJuly1993(Fishbeinandothers1995).Turner,Garza-S.,andHaroreturnedonazoologicalexpeditionin1998alongwithCecilSchwalbe,TaylorEdwards,CarlOlson,YarPetryszyn,DavePrival,andothers(Albertiandothers1998). Withafewexceptions,documentationofanimalsandplantsintheArizona-Sonoraborderlandswasmostlyneglectedafter the1848-1857and1892-1894boundarysurveys.Evencommon,widespreadspecieshadnotbeencollectedorobserved.Beginningin2001,VanDevenderandReina-Guerrero(2005)begancollectingandobservingplantsinLa Frontera,theareainnorthernSonorawithin100kmoftheArizonaborder,aspartofseveralinventoryprojects. LeonardoVarela-Espinosa andAlbertoBúrquez-Montijo at theUniversidadNacionalAutónomadeMéxicoinHermosillostudiedthefloraoftheSierraSanJavieronseventripsfromJuly1996toMay1997(Varela-Espinosa2005).TheSierraSanJavieristhesouthernmostSkyIslandwithtropicaldeciduousforestbelowoakwoodland.Since1994,HerbarioUniversidaddeSonora(USON)hasgrownintoamajorregionalcollection.In2003-2004,CuratorJ.JesúsSánchez-Escalante,ManuelEspericueta-Betancourt,andReynaA.Castillo-GámezfromUSON,andVanDevenderandReina-GuerreroatASDMstudiedthefloraoftheSierradeMazatán(Sánchez-Escalanteand

Figure 7—Stephen S. White and José Vera-Santos in the Sierra El Tigre on 13 October 1941.

Figure 6—(A) View from the Sierra de los Ajos (photo by Dale S. Turner). (B) Mixed-conifer forest in Sierra El Tigre (photo by George M. Ferguson).

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others2005).Thisrangeisthesouthwestern-mostSkyIslandlocated70kmeastofHermosilloincentralSonora.BetweenMarch2009andSeptember2011,Sánchez-Escalante,DeniseZ.Ávila-Jiménez,DavidA.Delgado-Zamora,andLilianaArmenta-CotafromUSON,andVanDevenderandReina-GuerreroatSkyIslandAlliancestudiedthefloraofRanchoElAribabiintheSierraAzulonsevenfieldtrips.ElAribabiConservationRanchintheSierraAzulwasdesignatedanÁreaNaturalProtegidaPrivadabyCONANPinMarch2011. InJuly-August2005,biologistsfromtheUniversityofArizonaandtheSonoranDesertResearchStationinventoriedthefaunaandfloraalongtheRíoÁros,atributaryoftheRíoYaquiineast-centralSonora (O’Brien and others 2006). Participants included CharlesO’Brien,AaronD.Flesch,EricWallace,MichaelD.Bogan,SamiaCarrillo-Percástegui,SkyJacobs,andCharlesvanRiperIII. InApril-May2008,SkyIslandAllianceorganizedanexpeditiontotheSierraElTigretoobserveplantsandanimals(Avila-VillegasandJacobs2008).ParticipantswereSergioAvila-Villegas,PaulCondon,SkyJacobs,MattSkroch,RobertVilla,RodolfoVilla,andlandowners.

Madrean Archipelago Conferences Asrecentlyastwodecadesago,fewpeopleknewoftheMadreanArchipelagoasadistinctregionandtheevidenceforitsimportancewasscatteredamongmanydisciplinesintwocountries.InSeptem-ber1994,aconferenceentitledBiodiversity and Management of the Madrean archipelago: The Sky Islands of Southwestern United States and Northwestern Mexicowasorganizedby theRockyMountainForest andRangeExperimentStation,U.S.ForestService, alongwith Sky IslandAlliance (DeBano and others 1995).Therewere69presentationsand20posters.ThesecondMadreanArchipelagoConferenceinMay2004wasentitledConnecting Mountain Islands and Desert Seas: Biodiversity and Management of the Madrean Archipelago.Therewere93presentationsand14posters.ThelackofinformationfromtheSkyIslandRegioninMexicowasnotedinbothconferences. InMay2012,thethirdMadreanArchipelagoconferenceassembledthecurrentstateofourknowledgeabouttheuniquenaturalandcul-turalresourcesoftheMadreanregionandcontinuedthediscussionofmanagementpracticesusefulformaintainingthoseresourceswith24sponsoringorganizations.Itbroughttogetherresearchers,partnersin resource stewardship, land managers, educators and students,government officials, consultants, and the interested public frombothsidesofthebordertoexaminetheMadreanArchipelagoofthesouthwesternUnitedStatesandnorthwesternMexico.Therewere151presentationsandposters,includingsubstantialadvancesintheknowledgeofanimalandplantdistributionsinMexico.

Madrean Archipelago Bodiversity Assessment OneoftheconclusionsofbothoftheMadreanconferenceswasthattherewasanurgentneedforinformationfromtheMexicanportionoftheSkyIslandRegion.SkyIslandAlliancewasaprincipalorganizerforbothoftheseconferences.Inspring2009,theNorthernMexicoConservation Program at SIA initiated theMadreanArchipelagoBiodiversityAssessment(MABA)project,inpartfundedbyagrantfromtheVeoliaEnvironmentFoundation.Theconceptofdocument-ingallplantandanimalspeciesintheMexicanSkyIslandRegionforuseinconservation,landmanagement,research,andeducationwasproposedbyformerSIAExecutiveDirectorMattSkroch,DaleTurner,andAaronD.Flesch.

TheonlineMABAdatabase(http://www.madrean.org)wascre-atedtomakeobservationsandimagesavailabletothepublic.TheFLORAhalfofthedatabaseisdirectlylinkedtotheSouthwesternEnvironmentalInformationNetwork(SEINet)database,whichhasdataforover2millionherbariumspecimens.TheFAUNAhalfofthedatabasewasanewdatabase.TheMABAdatabasediffersfromSEInetinthatmanyrecordsareobservationsratherthanspecimens,andthatobservationsareoftenillustratedwithcolorimages.Histori-calrecordsarefrommuseumcollections,literature,fieldnotesandotherdatabases.NewobservationsaremadeonfieldtripstoremotehighdiversitySkyIslandranges.Studyareaswereselectedbasedontheneedforbiologicalobservations,conservationinitiatives,andop-portunitiestocollaboratewithlandowners,landmanagers,andlocalresearchers. WhileobservationsaremadeonallMABAtrips,expeditionstohighpriorityareaswith25-45participantsareespeciallyproductive.These are unprecedented groups of animal and plant specialists,landowners,agencybiologists,universityprofessorsandstudents,journalists,photographers,andvolunteers(fig.8).In2009-2011,therewerefiveMABAExpeditions.InSeptember2009,17participantswenttotheSierraSanLuisonCuencaLosOjosFoundationproperty.InMarch2010,25participantswenttotheSierraElTigreonprivaterancheswithintheANPAjos-Bavispe.InAugust2010,39participants,includingprofessorsandstudentsfromthenearbyUniversidaddelaSierra(UNISIERRA),went to theSierraLaMaderaonprivateranches,partiallywithintheANPAjos-Bavispe.InApril-May2011,40participantswenttotheCiénegadeSaracachiandtheSierraSanAntonio.TheCiénegaisanimportantwetlandnaturalareaonRanchoAguaFríathatwasproposedasanÁreaNaturalProtegidaEstatalbytheComisióndeEcologíayDesarrolloSustentabledelEstadodeSonora(CEDES)in2010.TheSierraSanAntonioisprivatelyowned.InAugust2011,45participantswent to theSierraBacadéhuachi,whichismostlyprivatelyowned.RincóndeGuadalupe,ownedbytheCatholicDioceseofHermosillo,hasexcellentpotentialforprotectionasanaturalarea. TheMABAdatabaseistheprimarysourceofbiologicalrecordsforthestateofSonora.FLORAcontains112,623Sonoranobservations,including20,606fromMABA,20,331fromthebookSonoran Desert Plants. An Ecological Atlas (Turnerandothers1995),and13,285fromtheComisiónparaelUsoyConocimientodelaBiodiversidad(CONABIO).FAUNAcontains128,843Sonoranrecords,including60,856fromMABA,39,022fromthebookThe Birds of Sonora (Rus-sellandMonson,1998),33,993fromtheeBirddatabase,and27,404fromCONABIO.Species numbers in the MABA database provide preliminary diversity estimates for the Madrean Archipelago in So-nora. There are ca. 2880 plant taxa in northeastern Sonora. Insects are very diverse, but only 1380 taxa are documented in the Sky Islands Region. Vertebrates are better represented: fish (39 species), amphib-ians and reptiles (104 species), birds (358 species), and mammals (76 species). Surprisingly, diversity appears to decrease southward, but it probably reflects the north-northwest to south-southeast orientation of the ranges, and decreasing area and fewer biological inventories to the south.

Conclusions TheMABAprojecthaspulledtogetherinformationfrombiologi-calexpeditionsintheMadreanArchipelagofromthelast160years.The knowledge of plant and animal distributions has increaseddramaticallysince1950,reflectingfieldworkbyAmericanbotanists,herpetologists,ornithologists,andmammalogists,thedevelopmentofbiologyprogramsattheUniversidadesdeSonoraanddelaSierra,

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Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

theCONABIOnationalbiodiversitydatabase,andtheSonoranstateÁreaNaturalesProtegidasprogrambyCEDES.The2010bookDi-versidad Biológica de Sonora,editedbyFranciscoMolina-FreanerandVanDevender, summarized the stateofknowledgeofplants,animals,andvegetationinSonora.Theseaccomplishmentsareonlythebeginning—theopportunitiesforadditionalnaturalhistorystudiesintheMadreanArchipelagoareenormous!

Acknowledgments For their support and participation onMABAExpeditions,wethankMario Cirett and Rosa Elena Jiménez-Maldonado ofANPAjos-Bavispe;GertrudisYanes-Arvayo,HugoSilva-Kurumiya,MaríadelaPazMontañez-A.,andtheirstudentsofUNISIERRA;J.JesúsSánchez-Escalante of UNISON; Martín Villa-Andrade, GonzáloLuna-Salazar,MartínReyes-Juárez, andRogelioMolina-Freaner;andEnriqueYescasofSonoraEs magazine.Fortheirfieldobserva-tionsandgreatcompany,wethankAnaLiliaReina-Guerrero,JohnPalting,GeorgeFerguson,ChipHedgcock,SkyJacobs,CarlTomoff,SteveHale,RobertVilla,andTrevorHare.Wethankcocineros del campo OmarGutiérrez-Castro andRobertoTorres-García for tor-tillas y frijoles. MarcTrinksandCarolinePatrick-BirdwellhelpedwithMABAtriplogistics.WethankMelanieEmersonandtheSIABoardofDirectorsfortheirsupportoftheMABAproject.BioblitzFathersPaulMartin,SteveRussell,WendellMinckley,CharlesLowe,RaymondTurner,HowardGentry,PaulMartin,StephenWhite,Joe

Figure 8—MABA group at Rincón de Guadalupe in the Sierra Bacadéhuachi in August 2011 (photo by Chris Marzonie).

Marshall,EdgarMearns,JohnLemmon,JohnBartlett,andWilliamH.Emorywalkedthebiodiversitypathlongbeforeus.

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