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  Pleasecitethisarticleinpressas:Garbino,G.S.T.,etal.,Taxonomyofthepygmymarmoset( Cebuella Gray,1866):Geographicvariation,speciesdelimitation,andnomenclaturalnotes.Mammal.Biol.(2018),https://doi.org/10.1016/j.mambio.2018.09.003 ARTICLE IN PRESS G Model MAMBIO-41047;No.of    Pages8MammalianBiologyxxx(2018)xxx–xxx Contents lists available at ScienceDirect Mammalian   Biology  journal homepage: www.elsevier.com/locate/mambio Original   investigation Taxonomy   of    the   pygmy   marmoset   ( Cebuella   Gray,   1866):   Geographicvariation,   species   delimitation,   and   nomenclatural   notes Guilherme   S.T.   Garbino a , ∗ ,Daniel   M.   Casali a , b ,Fabio   O.   Nascimento c , José   Eduardo   Serrano-Villavicencio c , d a Pós-Graduac¸ãoemZoologia,InstitutodeCiênciasBiológicas,UniversidadeFederaldeMinasGerais,Av.AntônioCarlos,6627,Pampulha31270-901,BeloHorizonte,   MG,   Brazil b LaboratóriodeEvoluc¸ãodeMamíferos,DepartamentodeZoologia,InstitutodeCiênciasBiológicas,UniversidadeFederaldeMinasGerais,Av.AntônioCarlos,   6627,Pampulha31270-901,BeloHorizonte,MG,Brazil c Mastozoologia,MuseudeZoologia,UniversidadedeSãoPaulo,AvenidaNazaré,481,Ipiranga,04263-000,SãoPaulo,SP,Brazil d CentrodeInvestigaciónBiodiversidadSostenible(BioS),CalleFranciscodeZela,1556,Lince,Lima,Peru a   r   t   i   c   l   e   i   nf   o  Articlehistory: Received4April2018Accepted12September2018AvailableonlinexxxHandledbyDaisukeKoyabu Keywords: Callitrichidae CebuellapygmaeaCebuellaniveiventris GMYCTypelocality ab   s   t   ra   ct Weexamined   arecent   taxonomic   appraisal   of    Cebuella   by   Boubliet   al.With   an   increasedsamplespanningmostof    the   geographicrangeof    the   genus,   we   investigated   ventral   pelage   variation   in Cebuella   totestthe   propositionof    Boubli   et   al.   thatthere   were   three   geographically-restricted   phenotypes.   Additionally,we   conducted   amodel-based   species   delimitation   test   using   published   cytochrome   b   data   and   verifiedthe   typelocalities   of    theavailablenomina.   Contrary   toBoubliet   al.,   ouranalysis   showed   thatventralpelage   of    Cebuella   varies   consistently   along   geography,   indicating   the   existence   of    two   species-grouptaxa,   not   three   assuggested   by   the   formerauthors.Themodel-basedspecies   delimitation   alsoindicatedthat   Cebuella iscomposed   of    more   than   one   species.   Therange   of    the   two   species   proposed   here   is   differentthan   assumed   by   previous   authors,   with   one   taxon   occurring   northof    middle/lower   Rio   SolimõesandRíoNapoand   the   other   from   south   of    these   rivers.After   an   extensiveliterature   review,   weshowthatthe   type   localityof    Cebuellapygmaea   can   be   reliablyrestricted   tothe   northern   margin   of    RioSolimões.Consequently,   the   oldest   available   names   forthe   two   species   are   Cebuellapygmaea   for   thepopulationsnorthof    middle/lower   Rio   Solimões   and   RíoNapo,   and Cebuellaniveiventris   for   the   populations   south   of these   rivers.©2018   Deutsche   Gesellschaft   f ¨ur   S¨augetierkunde.   Publishedby   Elsevier   GmbH.All   rights   reserved. Introduction Pygmymarmosets, Cebuella Gray,1866,arethesmallestanthro-poidprimates,witha   bodymassof    85–140g(Soinietal.,1988)and askulllengthrangingfrom33.7to38.9mm(Hershkovitz,1977).Theyhaveamanecoveringtheears,armsthataremuchlargerthanthehindmembers(i.e.a   largeintermembralindex),andnoprotoconeinthefirstupperpremolar(GarbinoandMartins-Junior,2017).Thegenus Cebuella isa   memberofthemarmosetclade,whichalsocontainsthegenera Callithrix and Mico (Schneideretal.,   2012). Cebuella shareswithbothgeneradentaladaptationstotreegougingsuchasenlargedlowerincisors,a   V-shapedmandibleanda   highfrequencyofverticalclinging(Kinzeyetal.,1975;Rosenberger, ∗ Correspondingauthor. E-mailaddress: gstgarbino@hotmail.com(G.S.T.Garbino). 1984).Dependingontheauthor,thepygmymarmosethasbeenplacedinthesamegenusastheothermarmosets(Barrosoetal.,1997;Elliot,1913;Mittermeieretal.,1988;Rosenberger,1981; Townsend,2001),initsown   subgenus(Gray,1866;Groves,2001, 2005),orinaseparategenus(Garbino,2015;Hershkovitz,1977; Hill,1957;Rylandsetal.,2009;RylandsandMittermeier,2013). PygmymarmosetshaveanAmazoniandistribution,occurringbetweentheRíoCaquetáin   southernColombiaandtheleftbankof theRioMadeira,in   Brazil,beingprobablylimitedtotheeastbytheentirePurusandMadeirainterfluvium,andto   thewestbytheRíoMayoandtheRíoHuallagainwesternPeruvianAmazonia(Rylandsetal.,2009;VanRoosmalenandVanRoosmalen,1997). Thetaxonomichistoryof  Cebuella beginswithSpix(1823),   whogavethefirstscientificnameforthepygmymarmoset, Iacchuspyg-maeus ,basedona   singleanimalcollectednearTabatinga,Brazil.Inhis   description,theventralpelagecolorationwasnotmentioned.Wagner(1833)doubtedthevalidityof    thetaxonandaffirmedthatSpix’sspecimenprobablywasajuvenileofsomealreadyknown https://doi.org/10.1016/j.mambio.2018.09.0031616-5047/©2018DeutscheGesellschaftf ¨urS¨augetierkunde.Publishedby   ElsevierGmbH.Allrightsreserved.  Pleasecitethisarticlein   pressas:Garbino,G.S.T.,etal.,   Taxonomyof    thepygmymarmoset( Cebuella Gray,1866):Geographicvariation,speciesdelimitation,andnomenclaturalnotes.Mammal.Biol.(2018),https://doi.org/10.1016/j.mambio.2018.09.003 ARTICLE IN PRESS G Model MAMBIO-41047;No.of    Pages82 G.S.T.Garbinoetal./MammalianBiologyxxx(2018)xxx–xxx species.Lesson(1840,p.192)describedtheunderpartsofthespeciesasreddish( roussàtre )and,followingWagner,didnotruleoutthepossibilitythatSpix’s Iacchuspygmaeus wasa    juvenile.BasedonspecimenscollectedbyFrancisdeLaportedeCastelnauinPeru(thefirstonesknowntoscienceafterSpix’sdescription)andonaColombianspecimen,IsidoreGeoffroySaint-Hilaire(1851,p.61)confirmedthatSpix’s  pygmaeus representedanewspeciesandnotjustajuvenileofanotherform.Thesameauthor,studyingthespecimenswhichhepreviouslyreported(Geoffroy,1851),stated thattheventralpelageof    thespeciesvariedfromwhitetoreddish( Ledessousducorpsvariedu   blancauroussâtre )(Geoffroy,1855).Afteranalyzingthedentitionof    Spix’stypeandtakingnoticeof Castelnau’sspecimens,Wagner(1855)wasalsoconvincedthat  pyg-maeus wasavalidspecies.Inhisdescriptionof    Spix’stype,Wagner(1855,p.126)mentionedthattheventrumwasunicolored,withayellowish-claycolor( DieUnterseiteisteinfärbiglehmgelbich ).Elliot(1913,p.233)reproducedWagner’sdescriptionofthetypeandadded,mostprobablybasedonspecimenshe   analyzed,thefollow-ingdescriptionoftheunderparts:“throatandupperpartof    breastyellowishbrown;abdomengray”.Lönnberg(1940)comparedtheventralpelageof    specimensfromLagoIpixunawithspecimensfromJoãoPessoa(todayknownas   Eirunepé),bothsitessouthoftheRioSolimões.Thisauthorcon-sideredthatthosespecimensfromJoãoPessoa(RioJuruá)werethetypical  pygmaea andthosefromtheLagoIpixunawith ¨purewhitepelage¨onthechestandabdomen,adistinctsubspecies, Cebuella pygmaeaniveiventris .daCruz-Lima(1945)f ollowedLönnbergin recognizingapolytypic Cebuellapygmaea ,   asdidCabrera(1958),Vieira(1955),Hill(1957),   NapierandNapier(1967),andNapier (1976).Hershkovitz(1977),basedon121specimens,recognizedamonotypic Cebuellapygmaea andtreatedtheonlyothernameattributedtothegenus, C.    p.niveiventris Lönnberg,1940,   asajuniorsynonymof  Cebuellapygmaea .Theauthorarguedthatthewhiteandocherventerphenotypeswereindividualvariantsof    thesamespecies.However,36of    the121specimenswerefromanimpre-cise   localitylabeledas“nearTabatinga”,makingitimpossibletoascertainonwhichmarginofRioSolimõestheywerecaptured,andconsequentlytoevaluateif    thedistinctpelagepatternsreflecteddistincttaxonomicentities(Boublietal.,2018).Another24spec- imenswereofunknownlocalities.Recognitionof    asinglespecieswithnosubspeciesofpygmymarmosetwasfollowedafterHer-shkovitz(Coimbra-Filho,1990;Mittermeieretal.,1988;Townsend, 2001).VanRoosmalenandVanRoosmalen(1997),   basedonspecimenscollectedandobservedontheMadeira/Purusinterfluvium,arguedthat niveiventris wasa   validnameandassumedthatthistaxonwasdistributedsouthof    RioSolimões,while  pygmaea occurrednorthof thatriver.Tagliarioetal.(2000)firstnoteda   highlevelof    moleculardivergencewithin Cebuella ,but   thegeneticsampleswereobtainedfromcaptivespecimensofunknownprovenance,andnopheno-typiccomparisonsweremade.Groves(2001)followedVanRoosmalenandVanRoosmalen(1997)inclassifying Cebuellapygmaea withtwo   subspecies.Rylandsetal.(2009)alsorecognizedtwosubspeciesin   Cebuella pygmaea butnotedthatthegeographicvariationanddistributionofbothsubspeciesshouldbefurtherinvestigated.VossandFleck(2011)analyzedspecimensfromPeruandnotedthatskinswithamorewhitishventrumwerepresentsouthofRioSolimões/Amazonas,butneverthelessconcludedthattherewasnotaclearphenotypicbreakamongpopulationsandfollowedHershkovitz(1977)in   treating Cebuellapygmaea asmonotypic.Boublietal.(2018)carriedouta   genome-scalephylogeneticanalysisandshowedthatthereissignificantdivergencebetweenpopulationsof  Cebuella fromthenorthandsouthofRioSolimõesthatwaspartiallycongruentwithdifferencesinpelagecolor.Theyconcludedthatthereareatleasttwo   speciesandthreediagnosablemorphotypesofpygmymarmoset: Cebuella aff.  pygmaea fromtheupperRioJapurá, Cebuella cf.  pygmaea fromtheupperRioJuruá,westintoPeruandEcuador,and Cebuella cf. niveiventris fromthesouthern(right)marginofRioSolimões.Thenameavailableforeachtaxon,however,wasunclearbecausetheauthorsindicatedthatitwasnotpossibleto   ascertainwhetherSpix’stypelocalityof  Iacchuspygmaeus wasto   thenorthorsouthof    theRio   Solimões.Astheventralpelagecolorationof    Cebuella seemstobeanimportanttaxonomiccharacterwhichhasnotbeenthoroughlyinvestigatedinpreviousstudies,wepresentthefirstexplicitanal-ysisof    itsvariationalonga   widegeographicrange,includingspecimensfromEcuador,Colombia,Peru,andBrazil.Tofurthertestthetaxonomichypothesisof    Cebuella proposedbyBoublietal.(2018),   wecarriedouta   speciesdelimitationanalysisconsider-ingthecytochrome b treeobtainedbythepreviouslymentionedauthors.Finally,wediscussthetypelocalityof    Iacchuspygmaeus Spix,whichiscrucialindeterminingwhichnamesareavailableforthetaxarecognizedhere. Materialandmethods Morphologicalvariation Inorderto   explorethegeographicvariationintheventralpelageof  Cebuella ,   weanalyzedmuseumskinsin   thefollowingcollections–FieldMuseumofNaturalHistory(FMNH),InstitutoAlexandervonHumboldt(IAvH),InstitutodePesquisasdaAmazônia(INPA),MuseodeHistoriaNaturaldelaUniversidadNacionalMayordeSanMarcos(MUSM),MuseuParaenseEmílioGoeldi(MPEG),MuseudeZoologiadaUniversidadedeSãoPaulo(MZUSP),SmithsonianNationalMuseumofNaturalHistory(USNM),andUniversidadeFederaldeRondônia(UNIR).We   alsoexaminedphotosof    Cebuella fromtheSwedishMuseumof    NaturalHistory.We   assessedthevari-ationofventralpelagecolorationbyclassifyingthepatternsintodiscretecharacterstates,calculatingtheirfrequency,andmappingthem(seeGarbinoetal.,   2016f ora   similarapproach).We   analyzedatotalof    44taxidermiedskins(Table1). Model-basedspeciesdelimitationtest  Recentmethodologiesforspeciesdelimitationemploycom-putationalanalysestoidentifyputativespecies.Althoughthesemethodshavebeenpopularizedforothertaxa,theyarerarelyusedinNeotropicalprimates(Melloetal.,2018).Totesthowmany speciesof    Cebuella thereare,we   employedaGeneralizedMixedYuleCoalescent(GMYC)test.GMYCevaluatesthethresholdtimethatmaximizesthelikelihoodofatransitionbetweenaspecia-tionmodel(Yule)andacoalescencemodel,thusindicatingwhichregionsofthetreecouldbe   describedasinterspecificdivergencesamonglineagesandregionsofintraspecificdivergences(FujisawaandBarraclough,2013).Thesinglethresholdanalysisconsidersthatonlyonepointof    transitionamongintraandinterspecificportionsofthetreewillbeinvestigated,whilethemultiplethresholdoneallowsthepossibilitythatmultipleof    suchtransitionsarepossi-blealongthebranches.Thismethodisa   discovery-type,unilocusspeciesdelimitationapproachandwas   appliedinorderto   evaluatethemolecularinformationpresentinBoublietal.(2018)thatindi- catetheexistenceofmorethanonespeciesof    Cebuella .For   that,we   conductedtheanalysesconsideringtheultrametrictreeofthemitochondrialcytochrome b –CYTB–genepublishedinBoublietal.(2018),   including14samplesof    Cebuella and9othercallitrichinesasoutgroups,followingthesuggestionsof    Talaveraetal.(2013)f orsmalldatasets.Weperformedsingleandmultiplethresholdanal-ysesandcheckedthesignificanceof    themostlikelydelimitation  Pleasecitethisarticleinpressas:Garbino,G.S.T.,etal.,Taxonomyofthepygmymarmoset( Cebuella Gray,1866):Geographicvariation,speciesdelimitation,andnomenclaturalnotes.Mammal.Biol.(2018),https://doi.org/10.1016/j.mambio.2018.09.003 ARTICLE IN PRESS G Model MAMBIO-41047;No.of    Pages8 G.S.T.Garbinoetal./MammalianBiologyxxx(2018)xxx–xxx 3 Fig.1. Individualvariationinventralpelageinthetwo   speciesof    Cebuella : Cebuella pygmaea (allindividualswerescoredwithcharacterstate0).   Fromlefttoright:Brazil,Maguarizinho(MZUSP11468),Brazil,Serrinha(MZUSP11466),Colombia,Leticia(FMNH71003). Cebuellaniveiventris (characterstates1–3).Fromlefttoright:Brazil,Eirunepé(NRM612171),Brazil,Eirunepé(NRM612173),Ecuador,Oriente(FMNH54290),Brazil,Eirunepé(NRM612127),Brazil,Eirunepé(NRM612176),Brazil,Eirunepe(NRM612170),Brazil,Uarini(MPEG37114),Peru,Maniti(FMNH87135). schemewithrespectto   thenullhypothesisofasinglespecies.TheanalysiswasperformedintheExelixisLab’swebserver(GMYC–http://species.h-its.org/gmyc/). Results Variationinventralpelage Werecognized4characterstates,asfollows(Fig.1):(0)entirelyocherventer,withnodistinctchromogeneticfieldsbetweengularregionandchest/abdomen.(1)   grayishordirtywhitechestandabdomen.Clearcontrastbetweenochergularregionandtheimmediatelyposteriorchro-mogeneticfield.(2)grayishordirtywhitechestandabdomen,butwitha   whitesagittalmedianline.Clearcontrastbetweenochergularregionandthe   immediatelyposteriorchromogeneticfield.(3)whiteventer,withwhitehairsreachingtheunderpartsof forearmsandhindlegs.Clearcontrastbetweenochergularregionandtheimmediatelyposteriorchromogeneticfield.Weobservedtwogeographicallyrestrictedpatternsoftheven-tralpelagecoloration(Fig.2).Animalswithancompletelyocher venter(characterstate0)arerestrictedto   thenorth(right)mar-ginofthemiddle/lowerRio   Solimõesandnorth(right)marginof theRío   Napo(localities1,2,   5,8,   11,   12in   Fig.2).Southof    theserivers,allspecimenshaveatwo-zonedventrum,withanochergularregioncontrastingwiththechestandabdomen,whichcanrangefromentirelywhiteto   grayish(characterstates1,   2,   and3).Amongthethreecharacterstatesfoundsouthof    Napoandmid-dle/lowerSolimõesrivers,thepurewhiteventer(state3)wasbyfar   themostcommonstate,as72%ofoursample(24of    33adultspecimens)fromthatregionpresentedit.   Animalswitha   grayishventer(state1)or   withgrayishventerwithathinwhitemedianline(state2)comprised15%and12%of    thesample,respectively.Ourresultsdisagreewiththeproposalof Boublietal.(2018,p.173),thatthe Cebuella populationsalongtheupperRioJuruáandEcuadorandPeru(referredbytheformerauthorsas Cebuella cf.  pygmaea )havea   distinctphenotypefromthepopulationsalongthesouth-ernmarginoftheRioSolimões/Amazonas(referredas Cebuella cf. niveiventris ),asthepredominantstate3iswidespreadalongthisentireregion.We   examinedthespecimenCCM19ofBenjaminCon-stantreferredbyBoublietal.(2018)ashaving“someorangepelage onitsunderparts”butwedidnotfindanyorangeorocherhairsontheventrum.Theabdominalregionof    thisspecimenispartlynaked,thehair   probablylostduringthepreparationof    theskinandtheorangishskinisevident.Itis   possiblethatBoublietal.regardedtheskincolorasthepelagecolorinthisregion. GMYCspeciesdelimitation Thesinglethresholdanalysissuggested13entities(=species)intheentirephylogenyandfourin Cebuella .   TheLikelihoodRatioTest(LRT)indicatedthatthisdelimitationresultissignificantlybetterthanthenullhypothesisof    a   singlespecies(LR=   10.76359,LRTresults=   0.004599552).Themultiplethresholdanalysissug-gestedfivespeciesof    Cebuella (14includingtheoutgroups),and,asinthesinglethresholdanalysis,allowedus   to   rejectthenullhypothesis(LR=   10.93688,LRTresults=0.004217802).Theputa-tivespeciesandassociatedindividualsareshownin   Fig.3.   Bothanalysescorrectlyplacedtheoutgroupspeciesasseparatelineages(seeSupplementaryfile). Thetypelocalityof  Iacchuspygmaeus Spix,1823 Spix(1823)described Iacchuspygmaeus in   LatinandFrench.IntheLatindescription,hegavethelocalityas(p.32): Habitatinpagomunito,limitrophoTabatingaadripasfluviiSolimöens .Thispart   of    thetextcanbetranslatedas“livedin   a   village,limitrophetoTabatinga,onthebanksof    riverSolimões”.InthedescriptioninFrench(p.   33),SpixwrotethathereceivedthepygmymarmosetalivefromtheforestsofTabatinga: Iel’airec¸uvivantdesforêtsde   Tabatinga,maisilvécutpeudesemainesetmouritdemelancolie . JohannAndreasWagner,whodescribedsomeof    thematerialcollectedbySpixandMartiusontheBavarianexpedition,alsostatedthatthepygmymarmosetwas   collectedintheforestsof Tabatinga,ontheupperRioAmazonas(Wagner,1855,p.127): SpixhatdasvonihmbeschribeneExemplarinderWäldernvonTabatingaamobernAmazonenstromerhalten. FromSpix’saccounts,itis   evidentthattheholotypeof    Iacchus pygmaeus wasobtainedaliveandwas   keptbrieflyasa   pet.Wagner(1855)confirmedthisinformationbasedonapersonalcommuni-cationofCarlFriedrichvonMartius,whotoldhimthat   thepygmymarmosetlivedwithmembersof    theBavarianexpeditionforaperiod.Thereis   no   evidence,however,thatitwascollectedsouthof   Pleasecitethisarticlein   pressas:Garbino,G.S.T.,etal.,   Taxonomyof    thepygmymarmoset( Cebuella Gray,1866):Geographicvariation,speciesdelimitation,andnomenclaturalnotes.Mammal.Biol.(2018),https://doi.org/10.1016/j.mambio.2018.09.003 ARTICLE IN PRESS G Model MAMBIO-41047;No.of    Pages84 G.S.T.Garbinoetal./MammalianBiologyxxx(2018)xxx–xxx  Table   1 Specimensof  Cebuella analyzedinthisstudy,andtheirlocalitiesarrangedbylatitudeinincreasingorder.The“ID#”fieldrefersto   thelocalitiesnumberedinFigs.2and4.TaxonLocalityDepartamento/EstadoCountryID#CoordinatesSpecimen(s) Cebuellapygmaea Serrinha,RioJapuráAmazonasBrazil11 ◦ 33 ′ S69 ◦ 26 ′ WMZUSP11466 Cebuellapygmaea  Maguarizinho,RioJapurá AmazonasBrazil21 ◦ 37 ′ S69 ◦ 16 ′ WMZUSP11467,MZUSP11468 Cebuellaniveiventris RíoBobonaza,MontalvoPastazaEcuador32 ◦ 04 ′ S76 ◦ 58 ′ WFMNH41491 Cebuella   niveiventris Pastaza,RíoCopatazaOrienteEcuador42 ◦ 07 ′ S,77 ◦ 27 ′ WFMNH54290 Cebuella    pygmaea ZonaReservadaYaguasLoretoPeru   52 ◦ 55 ′ S71 ◦ 31 ′ WMN   2781,MN2782 Cebuella   niveiventris Uarini,leftmarginRioSolimões,Barroso,ReservadoDesenvolvimentoSustentávelMamirauáAmazonasBrazil63 ◦ 00 ′ 16 ′′ S65 ◦ 10 ′ 52 ′′ WMPEG37114 Cebuella   niveiventris Iquitos,RioManiti,SantaCeciliaLoretoPeru   73 ◦ 26 ′ S,72 ◦ 46 ′ WFMNH87135,FMNH87136,FMNH87137 Cebuella    pygmaea RíoAmacayacu,PuertoMogue,nearmouthof QuebradaCabimasAmazonasColombia83 ◦ 41 ′ S,70 ◦ 24 ′ WIAvH1855 Cebuellaniveiventris Maynas,QuistocochaLoretoPeru   93 ◦ 49 ′ 46 ′′ S73 ◦ 19 ′ 19 ′′ WFMNH122750(9286),FMNH122751(9287),FMNH122752(9288) Cebuellaniveiventris LagodoIpixunaAmazonasBrazil103 ◦ 52 ′ S63 ◦ 52 ′ WNRMA59,NRM4504(Photo) Cebuella    pygmaea IgarapéTacana,AltoSolimõesAmazonasBrazil114 ◦ 06 ′ S69 ◦ 33 ′ WMN   11910 Cebuellapygmaea LeticiaAmazonasColombia124 ◦ 15 ′ S69 ◦ 56 ′ WIAvH1198,FMNH71003(6573) Cebuellaniveiventris Yaquerana,RíoGalvez,San JuanLoretoPeru   135 ◦ 12 ′ S72 ◦ 53 ′ WMUSM11100 Cebuella   niveiventris LagoMatupirizinho,RioMadeira,NovoJerusalémAmazonasBrazil145 ◦ 33 ′ 28 ′′ S61 ◦ 07 ′ 20 ′′ WINPACCM251 Cebuella   niveiventris  AltoYavari,mouthof    RioYaqueranaAmazonasPeru   155 ◦ 43 ′ S72 ◦ 58 ′ FMNH88997,FMNH88998,FMNH88999 Cebuella   niveiventris  Democracia,leftbankof RioMadeiraAmazonasBrazil165 ◦ 48 ′ S61 ◦ 26 ′ WINPA4119 Cebuella   niveiventris Eirunepé,leftbankof    Rio JuruáAmazonasBrazil176 ◦ 39 ′ 36 ′′ S69 ◦ 52 ′ 26 ′′ WNRM612127,612170,612171,612172,612173(Photo) Cebuella   niveiventris SarayacuLoretoPeru   186 ◦ 44 ′ S75 ◦ 07 ′ WUSNM16610 Cebuella   niveiventris PauiniAmazonasBrazil197 ◦ 42 ′ S66 ◦ 58 ′ WMZUSP11370,MZUSP11371,MZUSP   11376 Cebuellaniveiventris FeijóAcreBrazil208 ◦ 09 ′ S70 ◦ 21 ′ WMPEG21854 Cebuella   niveiventris Ocidente,rioJuruá,PortoWalterAcreBrazil218 ◦ 33 ′ 58 ′′ S72 ◦ 47 ′ WMPEG22953,MPEG22954,MPEG22961 Cebuella   niveiventris PortoVelhoRondôniaBrazil228 ◦ 46 ′ 46.4 ′′ S63 ◦ 58 ′ 14 ′′ WUFRO-M175 Cebuella   niveiventris BocadoChandless,RioPurusAcreBrazil239 ◦ 8 ′ S69 ◦ 50 ′ W   MZUSP11240 Cebuellaniveiventris RioMacauãAcreBrazil249 ◦ 52 ′ S69 ◦ 23 ′ WMZUSP35733,MZUSP35734,MZUSP   35735 RioSolimões,andthespecies’descriptionin   Latinclearlystatesthatthe   specimenwasobtainedfromavillageadjacenttoTabatinga,meaningthatthereferredsettlementnecessarilywason   thenorthbankofRioSolimões.Theadjective“limitrophe”,usedbySpixtorefertothetypelocality,isespeciallyrelevantif    weconsiderthatthewidthofRioSolimõesin   theregionofTabatingamay   reach3km.   AnotherimportantfactisthatTabatingawasthewestern-mostpointexploredbytheBavarianexpedition–neitherRio    JavarinorBenjaminConstantwereexploredbySpixorothermembersof theexpedition(Papavero,1971).Therefore,thetypelocalityof    Iac-chuspygmaeus shouldbe   consideredtolieintheneighboringareasofTabatinga,onthenorthbankof    RioSolimões.Thereferenceof    Tabatingalyingon“thesouthernsideofRioSolimõesnearthejunctionwithRioJavar´y”(Lönnberg,1940,p.22) shouldberegardedasanerrorandnotasa   typelocalityrestriction,asaffirmedbyBoublietal.(2018,p.179). Taxonomyof  Cebuella Gray,1866 Basedonthephenotypicdifferencesin   ventralpelagecolorationobservedinthisstudy,congruentwitha   considerablydeepdiver-gencefoundin   themolecularphylogeny,werecognizetwo   speciesin Cebuella (Fig.3).Theoldestavailablenamesforthespecies are    pygmaea Spix,1823and niveiventris Lönnberg,1940(Fig.4). Althoughwhendescribing Cebuellapygmaeaniveiventris LönnbergcomparedthetwosyntypeswithspecimensfromJoãoPessoa(=Eirunepé)thathe   consideredtobe“typical  pygmaea ”,boththesyn-typesandtheseriesof    sixspecimensfromEirunepérepresent,inouropinion,variantsofthesametaxonomicentity.AlthoughGMYCsuggestedmorethantwoputativespecies,weoptedfora   moreconservativeapproach,usingintegrationof    evi-dencebycongruence(Padialetal.,2010)andacceptingasspecies onlydelimitationscommonto   allsourcesevaluatedhere(morphol-ogyandmoleculardelimitation).TheresultspresentedbyGMYCrejectedthenullmodelof    asinglespecies.FailuretorejectthenullGMYCmodelwoulddemandtherecognitionofasinglespecies,despitethemorphologicalvariation.Also,theserecognizedtaxapresentanallopatricdistribution,congruentwithour   taxonomicproposal.Hence,werecognizedonlytwospecies,forwhichwehavealsoobservedgeographicallyconsistentmorphologicalvari-ationbetweenthepopulations(Fig.2).Thedistributionof    thetwospeciesof  Cebuella proposedherediffersfromtheonepresentin  Pleasecitethisarticleinpressas:Garbino,G.S.T.,etal.,Taxonomyofthepygmymarmoset( Cebuella Gray,1866):Geographicvariation,speciesdelimitation,andnomenclaturalnotes.Mammal.Biol.(2018),https://doi.org/10.1016/j.mambio.2018.09.003 ARTICLE IN PRESS G Model MAMBIO-41047;No.of    Pages8 G.S.T.Garbinoetal./MammalianBiologyxxx(2018)xxx–xxx 5 Fig.2. Geographicvariationintheventralpelagecolorationof    Cebuella .   Forinformationon   thespecimensandlocalitiesseeTable1. Boublietal.(2018)anditisdetailedbelow,ontheaccountof    eachtaxa.Cebuellapygmaea (Spix,1823)Type-locality “inpagomunito,limitrophoTabatingaadripasfluviiSolimöens”hereinterpretedas“inavillagelimitropheto   Tabatinga,onthenorthernbankof    RiverSolimões”. Diagnosisanddescription Thedorsumhasanbizonalpattern,withchromogeneticfieldofthemanecontrastingwiththesaddleandrumpregions.Manecoveringtheears;hairsmultibandedwithanagoutipatternthroughouttheentiredorsalpartof    thebody.Thepelageof    thedorsumposteriortothemaneismottledwithblack,grayandbuff spots.Thispatternextendsto   theforearmsandlegs,but   thehandsandfeetaremorehomogeneouslybuff-colored.Thetailisringed,witha   patternofalternatingbuff,blackandgrayrings.Thefeetandhandsarelightochraceous.Theinsideofthelimbs,thechest,andtheabdomenvaryindividuallyfromochraceoustoclayorange.Oursampledidnothaveenoughrepresentativesof    distinctage   classestoverifyifthereisvariationrelatedtosexor   age. Variationinventralpelagecoloration We   foundsubtleindividualvariationinthecolorationof    theventralpelageof  Cebuellapygmaea withmostspecimenspresentinga   homogeneouslyochraceousventer,whichmay   belighter,moreorangish(Fig.1).AnexceptionisspecimenMN   11910,fromIgarapéTacana(Brazil),thatpresenteda   clearercolorinthethroatregion,contrastingwiththeadjacentochraceouschest,however,thisis   a juvenileandthevariationsobservedmay   beontogenetic. DistributionCebuellapygmaea occursbetweentheRíoNapoin   thewestandtheRioJapuráin   theeast.Its   southernlimitisboundedbythemid-
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