sitivitypaessler

Orobothriurus(Scorpiones:Bothriuridae)phylogeny,Andeanbiogeography,andtherelativeimportanceofgenitalicandsomaticcharactersCAMILOI.
MATTONI,JOSEA.
OCHOA,ANDRESA.
OJANGURENAFFILASTRO&LORENZOPRENDINISubmitted:23August2011Accepted:30November2011doi:10.
1111/j.
1463-6409.
2011.
00508.
xMattoni,C.
I.
,Ochoa,J.
A.
,OjangurenAflastro,A.
A.
&Prendini,L.
(2012).
Orobothriurus(Scorpiones:Bothriuridae)phylogeny,Andeanbiogeography,andtherelativeimportanceofgenitalicandsomaticcharacters.
—ZoologicaScripta,41,160–176.
ThegenusOrobothriurusMaury,1976(BothriuridaeSimon,1880)displaysanAndeanpat-ternofdistribution,mostofitsspeciesoccurringathighaltitudes(over2000–2500mtoamaximumaltituderecordof4910m)fromcentralPerutoArgentina.
TherecentdiscoveryofseveralnewspeciesandtheuncertainphylogeneticpositionofOrobothriuruslourencoiOjangurenAflastro,2003,requiredareanalysisofOrobothriurusphylogeny.
Thirtybo-thriuridtaxa,includingallspeciesofOrobothriurusandPachakutejOchoa,2004,werescoredfor65morphologicalcharactersandanalysedwithparsimonyunderequalandimpliedweighting.
Theresultingtopologyjustiestheestablishmentofanewgenus,RumikiruOjangurenAflastroetal.
,inpress,forO.
lourencoiandacloselyrelated,newspecies,RumikiruatacamaOjangurenAflastroetal.
,inpress.
ItalsooffersnewinsightsaboutthephylogenyandbiogeographyofOrobothriurusandrelatedgenera.
Charactersfromthemalegenitalia(i.
e.
hemispermatophore),comprisingapproximately26%ofthemorphologicalmatrix,werefoundtobelesshomoplasticthanthosefromsomaticmorphology,contradictingsuggestionsthatgenitaliaareuninformativeorpotentiallymis-leadinginphylogeneticstudies.
Correspondingauthor:CamiloI.
Mattoni,LaboratoriodeBiologaReproductivayEvolucion,CatedradeDiversidadAnimalI,FacultaddeCienciasExactas,FsicasyNaturales,UniversidadNacionaldeCordoba.
Av.
VelezSarseld299,CP:X5000JJC,Cordoba,Argentina.
E-mail:cmattoni@efn.
uncor.
eduJoseA.
Ochoa,DepartamentodeZoologia,InstitutodeBiociencias,UniversidadedeSaoPaulo,RuadoMatao,travessa14,101,SaoPaulo,SP05508-900,Brazil.
E-mail:jaochoac2000@yahoo.
comAndresA.
OjangurenAflastro,MuseoArgentinodeCienciasNaturales''BernardinoRivadavia'',AvenidaAngelGallardo470,CP:1405DJR,CABA,BuenosAires,Argentina.
E-mail:ojanguren@macn.
gov.
arLorenzoPrendini,ScorpionSystematicsResearchGroup,DivisionofInvertebrateZoology,Ameri-canMuseumofNaturalHistory,CentralParkWestat79thStreet,NewYork,NY10024-5192,USA.
E-mail:lorenzo@amnh.
orgIntroductionThescorpiongenusOrobothriurusMaury,1976wasestab-lishedtoaccommodateseveralsmalltomedium-sizedbo-thriuridscorpions,endemictotheAndesandsurroundingsinPeru,ChileandArgentina(Fig.
1),andpreviouslyplacedinBothriurusPeters,1861.
Atthetimeofitscrea-tion,Maury(1976)recognizedeightspeciesinOrobothriu-rus:O.
alticola(Pocock,1899),typespeciesofthegenus;O.
curvidigitus(Kraepelin,1911);O.
peruvianus(Mello-Leitao,1948);O.
dumayi(Cekalovic,1974);O.
crassimanusMaury,1976;O.
incaMaury,1976;O.
parvusMaury,1976.
Maury(1976)recognizedtwospeciesgroupsinOrobothriurus,subsequentlynamedthealticolagroupandtheincagroup(Acosta&Ochoa2001),basedonmarkeddifferencesinsomaticandgenitalmorphology.
NofurthermentionwasmadeaboutthesystematicsofOrobothriurusfor25yearsuntilitsmonophylywasques-tionedinlargeranalysesofscorpionoidandbothriuridphylogeny(Prendini2000,2003a).
Subsequentworksonthegenusledtovarioustaxonomicchangesandadditions.
OrobothriurusdumayiwasreturnedtoBothriurus(Acosta&Ochoa2001)andsixnewspeciesweredescribed(Acosta1602012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176ZoologicaScriptaFig.
1MapofwesternSouthAmerica,plottingknownrecordsofallspeciesofthebothriuridscorpiongeneraOrobothriurusMaury,1976(circles),PachakutejOchoa,2004(stars)andRumikiruOjangurenAflastroetal.
,inpress(squares),topography(contourinterval1000m)andvolcanoes(triangles).
SinglerecordforOrobothriuruswawitaAcosta&Ochoa,2000fromBolivia(indicatedbyquestionmark)probablyduetomislabeling(Ochoaetal.
2011).
C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176161&Ochoa2000,2001;Ochoa&Acosta2002,2003;Ojan-gurenAflastro2003a):O.
wawitaAcosta&Ochoa,2000;O.
famatinaAcosta,2001;O.
iskayAcosta&Ochoa,2001;O.
atiquipaOchoa&Acosta,2002;O.
ampayOchoa&Acosta,2003;andO.
lourencoiOjangurenAflastro,2003.
Oneofthesespecies,O.
lourencoi,fromthecoastalAta-camaDesertinChile,possessesseveralcharactersuniqueamongbothriuridscorpions,includinganenlargedbasaltoothonthemediandenticlerowofthepedipalpchelamovablengerandanapophysissituatedsubmediallyontheinternalsurfaceofthepedipalpchelaoftheadultmale(OjangurenAflastro2003a).
Ochoa(2004)revisedOrobothriurusandpresentedacla-disticanalysisbasedon57morphologicalcharactersscoredforthe11speciesknownatthetime.
Ochoa's(2004)resultssupportedPrendini's(2000,2003a)predictionthatOrobo-thriuruswasparaphyletic,theincagroupbeingmonophy-leticwithBothriurustotheexclusionofthealticolagroup.
Ochoa(2004)establishedthegenusPachakutejOchoa,2004toaccommodatethespeciesoftheincagroupandtrans-ferredfourspeciesfromOrobothriurus:P.
peruvianus(Mello-Leitao,1948);P.
crassimanus(Maury,1976);P.
inca(Maury,1976);P.
iskay(Acosta&Ochoa,2001).
Inaddition,twonewspeciesweredescribed:P.
juchuichaOchoa,2004;P.
oscariOchoa,2004.
FollowingseveralexpeditionstotheAndesofPeru,ChileandArgentina,sevenadditionalspe-ciesofOrobothriurusweredescribed(OjangurenAflastroetal.
2009;Ochoaetal.
2011)andonewassynonymized(OjangurenAflastroetal.
2009),bringingthetotalspeciescountto16(Table1).
AsredenedbyOchoa(2004),thespeciesofOrobothriu-russhareseveralsynapomorphies:anarrow,elongatedhemispermatophore,withapoorlydevelopedloberegion,adividedfrontalcrest,anenlargedbasallobewithatortu-ousstemandasubdistalspoon-likedilationendinginater-minalprocess;anacuteapophysisontheinternalsurfaceofthepedipalpchelamanusoftheadultmale,withtheibtrichobothriumsituatedatitsbase;thecloseproximityandsubparallelorientationoftheventrosubmediancarinaetotheventrolateralcarinaeonmetasomalsegmentV.
Ochoa(2004)omittedO.
lourencoifromhisanalysis,sothisenigmaticspeciesremainedinOrobothriurusbydefault,pendinganempiricalevaluationofitsphyloge-neticplacement.
OrobothriuruslourencoiexhibitsseveralcharactersthatsuggestacloserrelationshiptoPachakutej(e.
g.
hemispermatophorewithapapillosefoldonthebasallobeandasclerotizedapophysisontheinternalfoldoftheinternallobe),butresemblesthespeciesofOrobothriurusinotherrespects(e.
g.
thecarinationofmetasomalsegmentV;OjangurenAflastro2003a:p.
119,g.
9).
Thediscoveryofanewspecies,closelyrelatedtoO.
lourencoi,intheAtacamaDesertofChile(OjangurenAflastroetal.
inpress),raisedfurtherquestionsaboutthephylogeneticplacementofthetwospecies.
AreanalysisofOrobothriurusphylogenywasthereforeperformed,usingarevised,expandedversionofOchoa's(2004)datamatrix,basedon17speciesofOrobothriurus(OjangurenAflastroetal.
2009;Ochoaetal.
2011),includingO.
lourencoi,omittedfromOchoa's(2004)analysis;thecloselyrelated,newspecies;allspeciesofPachakutej;andaselectionofotherbothriuridtaxa.
Resultsofthisanalysisjustifytheestablishmentofanewgenus,RumikiruOjangurenAflas-troetal.
,inpress,forO.
lourencoiandthenewspecies,RumikiruatacamaOjangurenAflastroetal.
,inpress,andoffernewinsightsaboutthephylogenyandbiogeographyofOrobothriurusandrelatedgenera.
Approximately26%ofthemorphologicalcharactersusedintheanalysisarederivedfrommalegenitalia(i.
e.
hemispermatophore).
Bothriuridscorpionsexhibitsomeofthegreatestdiversityinmalegenitaliaamongscorpions(Maury1980;OjangurenAflastro2005;Peretti2010).
Ithasbeenarguedthatgenitalcharactersofarthropodsevolverelativelyrapidly,becauseofstrongsexualselection,andhavelowphylogeneticinertia(Arnqvist&Rowe2002;Table1DescribedspeciesofthebothriuridscorpiongeneraOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress,withcountriesandprovinces(Argentina),regions(Chile)ordepartments(Peru)fromwhichtheyhavebeenrecordedOrobothriurusalticola(Pocock,1899)Argentina:Mendoza,SanJuanOrobothriurusampayOchoa&Acosta,2003Peru:ApurmacOrobothriurusatiquipaOchoa&Acosta,2002Peru:ArequipaOrobothriuruscalchaquiOchoaetal.
,2011Argentina:TucumanOrobothriuruscompagnucciiOchoaetal.
,2011Argentina:LaRiojaOrobothriuruscurvidigitus(Kraepelin,1911)Peru:ArequipaOrobothriurusfamatinaAcosta,2001Argentina:LaRiojaOrobothriurusgrismadoiOjangurenAflastroetal.
,2009Argentina:MendozaOrobothriurushuascaranOchoaetal.
,2011Peru:AncashOrobothriuruspaessleri(Kraepelin,1911)Peru:ArequipaOrobothriurusparvusMaury,1976Peru:Junn,LimaOrobothriurusquewerukanaOchoaetal.
,2011Chile:Tarapaca,Peru:TacnaOrobothriurusramireziOchoaetal.
,2011Chile:CoquimboOrobothriurustamarugalOchoaetal.
,2011Chile:TarapacaOrobothriuruswawitaAcosta&Ochoa,2000Peru:Ayacucho,CuscoPachakutejcrassimanus(Maury,1976)Peru:CajamarcaPachakutejinca(Maury,1976)Peru:CajamarcaPachakutejiskay(Acosta&Ochoa,2001)Peru:Apurmac,CuscoPachakutejjuchuichaOchoa,2004Peru:CuscoPachakutejperuvianus(Mello-Leitao,1948)Peru:JunnPachakutejoscariOchoa,2004Peru:CuscoRumikiruatacamaOjangurenAflastroetal.
,inpressChile:AtacamaRumikirulourencoi(OjangurenAflastro,2003)Chile:Antofagasta,AtacamaOrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1622012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176Eberhard2004).
Thisimpliesthatgenitalcharacterscouldbehighlyhomoplasticandthereforemisleading,oratleastuninformative,inphylogeneticstudies.
Recently,Song&Bucheli(2010)providedtherstempiricalassessmentofthisassumption,showingthatgenitalicandsomaticchar-actershavesimilarlevelsofhomoplasyininsects.
Similarcomparisonshavenotbeenconductedinarachnids,whichhavesclerotizedgenitalialikeinsects.
Wethereforepres-entacomparisonofhomoplasyamonggenitalicandsomaticcharactersinbothriuridscorpions.
MaterialandmethodsTaxaAll16describedspeciesofOrobothriurusandoneunde-scribedspecieswereincludedasingrouptaxainthepresentanalysis(Table2).
Theoutgrouptaxonsamplecomprised13speciesinvebothriuridgenera.
Allsixdescribedspe-ciesofPachakutejwereincludedtotesttheirrelationshipwithRumikiru.
FourspeciesofBothriuruswereincluded,basedonpreviousndingsofacloserelationshipwithspe-ciesofPachakutej(Prendini2000,2003a;Ochoa2004):Bo-thriurusasperPocock,1893;Bothriurusbonariensis(C.
L.
Koch,1842);BothriuruscordubensisAcosta,1995;BothriuruscoriaceusPocock,1893.
Threeexemplarspeciesofother,moredistantlyrelatedbothriuridgenerawerealsoincluded:Centromachetespocockii(Kraepelin,1894);Cercoph-oniussulcatusKraepelin,1908;UrophoniustregualemuensisCekalovic,1981.
ThetreewasrootedonC.
sulcatus,basedonpreviousevidencefortherelationshipsamongbothriu-ridgenera(Prendini2000,2003a).
MaterialexaminedSpecimenswerecollectedbyturningstonesduringthedayorbyultraviolet(UV)lightdetectionatnight(Honetsch-lager1965;Stahnke1972;Sissometal.
1990).
PortableUVlamps,comprisingtwomercury-vapourtubesattachedtoachromiumparabolicreectorandpoweredbyarechargeable7Amph,12Vbattery,orMaglite3DcellTable2Distributionof65charactersamong30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress051015202530354045505560Cercophoniussulcatus0011000010100000020110000011100110000000001110110–1010–––0100––00Urophoniustregualemuensis2101100010100000020110001111110110011100000110110–1010–––0100––00Centromachetespocockii0011000011010010030010020000010110000000000010000–1000–––0100––00Bothriurusasper10110001010111000000001200110000011111102000000100000100–0000––00Bothriurusbonariensis00110001010111000000001210110000011111102000000010020102–0000––00Bothriuruscordubensis1011000111011100000000101011000111111110200000000–0110–––1100––00Bothriuruscoriaceus0000000111011100000000101011000111111100200000000–0110–––1100––00Pachakutejcrassimanus00110001110100000000000000000010011111101000101110020100–1010––11Pachakutejinca00110001100100000000000000000010011111200100101110020100–1010––11Pachakutejiskay00110001110100000000000000000010011111101000101110020100–1010––11Pachakutejjuchuicha00111001110100000000000000000010011111200100101110020100–1010––11Pachakutejoscari00110001110100000000000000000010011111101000101110010100–1010––11Pachakutejperuvianus00110001110100000000000000000010111111001000101110020100–1010––11Rumikiruatacama0–110101110100010410012011010010011111010011100010220100–1010––01Rumikirulourencoi0–110101110100010410012011010010011111010011100010220100–1010––01Orobothriurusalticola111010001010000001000011211110111001110100111$1101120111100011100Orobothriurusampay00110000101000001–00001111110011100000010011111100120111000011000Orobothriurusatiquipa11101000101–00000100001121111010110–0011111201120111100011100Orobothriuruscalchaqui11101000101000000100001121111011100011010011101100120111000012000Orobothriuruscompagnuccii111010001010000001000011&1111011101111010011101100120111100011100Orobothriuruscurvidigitus11101010101000110100001221111010011111010011101101120111100011100Orobothriurusfamatina11001000101000000100001111111011100011010011101100120111000012000Orobothriurusgrismadoi10101000101000000100001121111011101111010–11111101120111100011100Orobothriurushuascaran11111000101000000100001221101011100011010011101201120111100011100Orobothriuruspaessleri11101010101100010100001221111010011111010010111100120111100011100Orobothriurusparvus11011000101000000100001111111011100011010011101200120111000010000Orobothriurusquewerukana11101010101100110100001121111010011111010011101100120111100011100Orobothriurusramirezi11101000101000000100001121111011100111010–11101101120111100011100Orobothriurustamarugal11002010101100110100001111111010111111010–10101100120111100011100Orobothriuruswawita11012000101000001–0000–1111110101011112–0111101200120111000011000Characterstatesarescored0–4,(unknown)or-(inapplicable);polymorphicstates:$=[01];&=[12].
RefertoAppendix1forcharacterdescriptionsandAppendixS1formaterialexamined.
C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176163torcheswith32UVLEDattachmentswereusedforeld-work.
Materialsexamined(AppendixS1)aredepositedinthefollowingcollections:AmericanMuseumofNaturalHistory,NewYork,USA;CatedradeDiversidadAnimalI,UniversidadNacionaldeCordoba,Argentina;FieldMuseumofNaturalHistory,Chicago,USA;InstitutoMiguelLillo,Tucuman,Argentina;LaboratoriodeBio-logaReproductivayEvolucion,UniversidadNacionaldeCordoba,Argentina;MuseoArgentinodeCienciasNatu-rales'BernardinoRivadavia,'BuenosAires,Argentina;MuseoNacionaldeHistoriaNatural,Santiago,Chile;MuseodeHistoriaNatural,UniversidadNacionaldeSanAntonioAbaddelCusco,Peru;MuseodeHistoriaNatural,UniversidadNacionalMayordeSanMarcos,Lima,Peru;MuseodeZoologa,UniversidaddeConcep-cion,Chile;MuseudeZoologia,UniversidadedeSaoPaulo,Brazil;MuseuNacional,RiodeJaneiro,Brazil.
CharactersThedatamatrixcomprises65characters,17fromgenitalic(hemispermatophore)morphologyand48fromsomaticmorphology(Table1;Appendix1).
Fifty-fourcharacterswerecompiledfromOchoa's(2004)originaldatamatrix,and11arenew.
MorphologicalterminologyfollowsVachon(1974)fortrichobothrialnomenclature;Vachon(1952)andPrendini(2000)forpedipalpcarinae;amodi-edversionofPrendini(2000,2003b)usedbyOchoaetal.
(2010)fortergite,sterniteandmetasomalcarinae;Mattoni&Acosta(2005)formetasomalmacrosetae;Ochoa(2004)andMattoni&Acosta(2005)forhemisper-matophore;Stahnke(1970)forothercharacters.
Charac-ters24and45werepolymorphicintwotaxa.
Thirteenmultistatecharactersweretreatedasunordered(nonadditive),exceptinsomeanalyses,inwhichthree(characters40,47and51)wereordered(additive).
Allcharacterswereparsi-mony-informative.
CladisticanalysisAnalyseswereconductedusingTNT1.
1(Goloboffetal.
2008).
Heuristicsearchstrategieswereperformedusingtraditionalsearchwith100randomadditionsequences(Wagnertrees)followedbytreebisection-reconnectionbranchswapping(TBR),keepingupto10treesperrepli-cation(commandsequence:'hold1000;mult=tbrreplic100hold10;').
TheresultingtreeswereusedasstartingpointsforaroundofTBRbranchswapping(command:'bbreak=TBR').
Alltreesfoundduringsearcheswerecol-lapsedunder'rule1'(minimumpossiblelengthiszero;Swofford&Beagle1993;Coddington&Scharff1994).
Thedatamatrixwasanalysedunderequalweights(EW)andimpliedweights(IW)(Goloboff1993).
Thesensitivityofresults(sensuWheeler1995)wasassessedbytreatingallcharactersunorderedandvaryingvaluesoftheconcavityconstantintheIWanalyses(k=1–10,12and15;com-mand:'Piwe=N',where'N'isthekvalue).
Twenty-sixanalyseswereconductedintotal,13(oneEWand12IWanalyses)withcharacters40,47and51ordered,and13inwhichthesecharacterswereunordered.
Resultsofthesen-sitivityanalysisweresummarizedbymeansofa50%majorityruletree(command:'majority=50'),whereasdif-ferentoptimaltreesobtainedfromeachanalysisweresum-marizedbymeansofstrictconsensustrees(command:'nelsen').
Adjustedhomoplasy(ordistortion)inIWanaly-ses,measuredwithaconvexincreasingfunction(Goloboff1993,1997),wascalculatedinTNT(command:'t').
Characteroptimizationwasconductedusingacceleratedtransformation(ACCTRAN)(Farris1970;Swofford&Maddison1987,1992)inWINCLADA1.
00.
08(Nixon1999–2002).
UnambiguousoptimizationswereproducedusingTNT(command:'apo-').
Therelativedegreeofsupportforeachnode(Bremersupport,BS;Bremer1994)wascalculatedusingTNT,bysearchingforsuboptimaltreesonesteplonger,keeping1000treesperreplication,untiltheBSwasobtainedforeachbranch(command:'bsupport;').
RelativeBremersup-port(RBS),whichtakesintoaccounttherelativeamountsofcontradictoryandfavourableevidence(Goloboff&Far-ris2001),wascalculatedsimilarly,usingonlytreeswithinabsolutesupport(command:'bsupport];').
RBSforagroupvariesbetween0(completelyunsupported)and100(entirelyuncontradicted).
Supportvalueswerealsoesti-matedusinggroupfrequenciesunderjackkning(Jackknifesupport,JS;reviewedbyGoloboffetal.
2003),withaprob-abilityofalteration,P=0.
36,byperforming1000pseu-doreplicatesof10randomadditionsequences,eachfollowedbyTBRswapping,keepingupto10trees(com-mandsequence:'mult:noratchetrepl10tbrhold10;res-amplejakrepl1000;').
TheaverageJSofeachphylogenetichypothesiswascalculatedwithTNTandusedasacriterionforchoosingamongalternativetrees(Ka¨llersjo¨etal.
1999).
HomoplasyandsynapomorphyTheconsistencyindex(CI;Kluge&Farris1969)andretentionindex(RI;Farris1989)ofthetreesandcharac-terswerecalculatedwithTNTusingthemacroscript'statsall.
run'developedbyPetersonL.
Lopes(Universid-adedeSaoPaulo).
TheCIandRIarethesimplestcharacterstatisticsusedtodescribelevelsofhomoplasyandsynapo-morphy,respectively.
TheaverageCIandRIwerealsocalculatedforsomecharactersystems,e.
g.
,pigmentation(characters0–5),externalmorphologyexcludingtricho-bothria(characters6–21,29–47),trichobothria(characters22–28)andhemispermatophores(malegenitalia,characters48–64).
OrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1642012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176ResultsAsinglemostparsimonioustree(MPT)of167stepswasobtainedinmostanalysesunderIW(k=3–9,10,12and15,Figs2and4).
ThesensitivityoftheresultsofIWtodifferentvaluesoftheconcavityconstantispresentedinTable3andFig.
5.
Thetopologywasstablein10ofthe12weightingregimesexplored.
Analyseswithweakerconcavityfunctions(i.
e.
thoseinwhichhomoplasticchar-acterswerepenalizedtoalesserextent)demonstratedhigheraveragesupport.
AnalyseswithEWandthreeorderedmultistatecharac-tersobtained18MPTs(167steps,CI=0.
479,RI=0.
813).
Althoughsomebranchescollapsedinthestrictconsensustree(Fig.
3),themajorcladeswereTable3Summaryofstatisticaldifferencesamongmostparsimonioustrees(MPTs)obtainedbyanalysesof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress,withequalweights(EW)andimpliedweights(IW)with12valuesfortheconcavityconstant(k)LengthMPTsCIRIAHAv.
JSEW1671641890.
4790.
4880.
8130.
816–28.
634.
0IWk=1171168770.
4680.
4760.
8040.
80727.
7826.
0332.
935.
8IWk=2167–169167–1682080.
473–0.
4790.
476–0.
4790.
808–0.
8130.
807–0.
80920.
7819.
5833.
036.
2IWk=3167164110.
4790.
4880.
8130.
81616.
6115.
7332.
836.
3IWk=4167164110.
4790.
4880.
8130.
81613.
8713.
1732.
736.
1IWk=5167164110.
4790.
4880.
8130.
81611.
9211.
3432.
736.
1IWk=6167164110.
4790.
4880.
8130.
81610.
469.
9731.
136.
1IWk=7167164110.
4790.
4880.
8130.
8169.
328.
8932.
535.
9IWk=8167164110.
4790.
4880.
8130.
8168.
418.
0332.
536.
2IWk=9167164110.
4790.
4880.
8130.
8167.
667.
3234.
338.
0IWk=10167164110.
4790.
4880.
8130.
8167.
046.
7334.
338.
0IWk=12167164110.
4790.
4880.
8130.
8166.
055.
7934.
541.
9IWk=15167164110.
4790.
4880.
8130.
8165.
004.
8034.
741.
8Valuespresentedforanalyseswiththreecharacters(40,47and51)ordered(additive)orunordered(non-additive).
Length,numberofstepsunderequalweights;MPTs,numberofmostparsimonioustrees;CI,consistencyindex;RI,retentionindex;AH,adjustedhomoplasyunderconcaveweightingfunctionofIWanalyses;Av.
JS,averagesupportusingJackkniferesampling(highestvaluesinbold).
7864529976717967838268500.
010.
020.
020.
100.
040.
050.
170.
020.
220.
110.
040.
540.
100.
230.
200.
060.
040.
010.
210.
160.
100.
060.
10OrobothriuruswawitaOrobothriurusgrismadoiOrobothriurustamarugalOrobothriurusramireziOrobothriurusquewerukanaOrobothriuruscompagnucciiOrobothriuruscalchaquiOrobothriurushuascaranOrobothriurusalticolaOrobothriurusatiquipaOrobothriuruspaessleriOrobothriuruscurvidigitusOrobothriurusampayOrobothriurusfamatinaOrobothriurusparvusRumikirulourencoiPachakutejperuvianusPachakutejjuchuichaPachakutejincaPachakutejoscariPachakutejiskayPachakutejcrassimanusBothriuruscordubensisBothriurusasperBothriuruscoriaceusBothriurusbonariensisUrophoniustregualemuensisCentromachetespocockiiCercophoniussulcatusRumikiruatacamaFig.
2Mostparsimonioustree(adjustedhomoplasy=5.
001,length=167)obtainedincladisticanalysisof30bothriu-ridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress,underimpliedweights(k=15,threecharactersordered).
Jackknifepercentagesindicatedabovebranches.
Bremersupportvalues,inunitsoft,indicatedbelowbranches.
Zerolengthbranchescollapsedunder'rule1'.
C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176165maintainedandwerecongruentwiththoseinthetopolo-giesobtainedbyanalyseswithIW.
Similartopologieswereobtainedwhenallmultistatecharacterswereunordered.
AsingleMPT(164steps,CI=0.
488,RI=0.
816)wasobtainedinmostIWanalyses(k=3–9,10,12and15)andnineMPTswereobtainedwithEW(164steps,CI=0.
488,RI=0.
816;Table3).
Rumikiruwasmonophyletic,stronglysupportedby11unambiguoussynapomorphies,BS=9,RBS=48,andJS=99–100(Figs2–4;Appendix2),andplacedsistertoPachakutejinallanalyses,arelationshipsupportedbyveunambiguoussynapomorphies,JS=62–84,BS=4,andRBS=33.
Pachakutejwasalsomonophyleticinallanalysesandsupportedbyveunambiguoussynapomorphies,JS=77–84,BS=4,andRBS=36.
ThemonophyleticgroupcomprisingPachakutejandRumikiruwas,inturn,sistertoBothriurusandsupportedbyveunambiguoussynapomorphies,JS=75–86,BS=4andRBS=50.
Orobothriurus,asredenedbyOchoaetal.
(2011),wasmonophyleticinallanalysesandsupportedbysixunam-biguoussynapomorphiesandahighBS=5(Figs2–5;Appendix2).
ThelowvaluesofJS=51–67andRBS=29indicatesomecontradictoryevidenceforthemonophylyofOrobothriurus,however.
AmonophyleticgroupoffourOrobothriurusspecies(O.
curvidigitus,O.
paessleri,O.
quewer-ukanaandO.
tamarugal)waswellsupportedinallanalysesbysixsynapomorphies,JS=69–78,BS=3,andRBS=60.
Withtheexceptionofthisgroup,otherrelationshipsamongthespeciesofOrobothriuruswereweaklysupported,andnotconsistentlyrecoveredinallanalyses(Fig.
5).
ConsistencyandretentionindicesforeachcharacterintheIWanalysiswithk=15areindicatedinFig.
5.
TheaverageCIforeachcharactersystemwasasfollows:0.
443(pigmentation),0.
597(somaticmorphologyexcludingtrichobothria),0.
422(trichobothria),and0.
752(hemisper-matophore).
TheaverageRIwasasfollows:0.
747(pig-mentation),0.
715(somaticmorphologyexcludingtrichobothria),0.
808(trichobothria),and0.
851(hemisper-matophore).
TheaverageCIandRIforallsomaticchar-acterswere0.
552and0.
733,respectively.
DiscussionNewgenusjustiedThephylogeneticrelationshipsretrievedinthereanalysisofOrobothriurusphylogenymostlycorroboratedOchoa's(2004)phylogenetichypothesis,whichwasbasedonasmallersampleoftaxaandcharacters.
Thetopologiesofthetwostudiesarecongruentinmostrespects,inspiteofthedifferenttaxonsamples.
BothstudiessupportedthemonophylyofPachakutej,itsplacementinamonophyleticgroupwithBothriurus,andthemonophylyofOrobothriu-rus.
OnlyafewoftherelationshipsamongspeciesofOrob-othriuruswereincongruent.
OrobothriuruswouldberenderedparaphyleticbythegroupingofO.
lourencoi(andthecloselyrelated,newspe-ciesfromtheAtacama)withPachakutej,hadOrobothriurus2/501/501/503/605/292/751/1009/482/1004/334/361/1001/1004/502/332/1001/502/2268635199786075826853OrobothriurusparvusOrobothriurusgrismadoiOrobothriurustamarugalOrobothriurusramireziOrobothriurusquewerukanaOrobothriuruscompagnucciiOrobothriuruscalchaquiOrobothriurushuascaranOrobothriurusalticolaOrobothriurusatiquipaOrobothriuruspaessleriOrobothriuruscurvidigitusOrobothriurusampayOrobothriurusfamatinaOrobothriuruswawitaRumikirulourencoiRumikiruatacamaPachakutejperuvianusPachakutejjuchuichaPachakutejincaPachakutejoscariPachakutejiskayPachakutejcrassimanusBothriuruscordubensisBothriurusasperBothriuruscoriaceusBothriurusbonariensisUrophoniustregualemuensisCentromachetespocockiiCercophoniussulcatusFig.
3Strictconsensusof18mostparsimonioustreesobtainedincladisticanalysisof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,Pachakutej,Ochoa2004andRumikiruOjangurenAflastroetal.
,inpress,underequalweightswiththreecharactersordered(length=167,CI=0.
479,retentionindex=0.
813).
Jackknifepercentagesindicatedabovebranches.
BremersupportrelativeBremersupportvaluesindicatedbelowbranches.
OrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1662012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176notbeenredenedtoexcludethem(Ochoaetal.
2011).
However,thesetwospeciescouldnothavebeenaccommo-datedwithinPachakutejeither(OjangurenAflastroetal.
,inpress).
ThecombinationofseveralsynapomorphieswithPachakutej,andseveraluniquesynapomorphies,justiedtheneedforanewgenus,Rumikiru,toaccommodateO.
louren-coiandthenewspecies,formaldescriptionsofwhichwereprovidedbyOjangurenAflastroetal.
(inpress).
RumikiruismostcloselyrelatedtoPachakutej,basedmainlyoncharactersofthehemispermatophore,i.
e.
,thesharedpresenceofonesclerotizedapophysisontheinternalfoldoftheinternallobeandapapillosefoldinthebasallobe.
However,speciesofPachakutejaremarkedlymorepig-mentedandpossessauniquesynapomorphy,aspatulateter-minalprocessonthebasallobeofthehemispermatophore(Ochoa2004)thatisabsentinRumikiru.
Incontrast,thepedipalpcarinae,especiallyofthefemurandpatella,aremorepronouncedinRumikiruthaninPachakutej.
Thetwogeneraalsodifferinthetrichobothrialpatternofthepedi-palpchelamanus.
TrichobothriumV2issituatedintheCercophoniussulcatusCentromachetespocockiiUrophoniustregualemuensisBothriurusbonariensisBothriuruscoriaceusBothriurusasperBothriuruscordubensisPachakutejcrassimanusPachakutejiskayPachakutejoscariPachakutejincaPachakutejjuchuichaPachakutejperuvianusRumikirulourencoiRumikiruatacamaOrobothriurusparvusOrobothriuruswawitaOrobothriurusfamatinaOrobothriurusampayOrobothriuruscurvidigitusOrobothriuruspaessleriOrobothriurusatiquipaOrobothriurusalticolaOrobothriurushuascaranOrobothriuruscalchaquiOrobothriuruscompagnucciiOrobothriurusquewerukanaOrobothriurusramireziOrobothriurustamarugalOrobothriurusgrismadoi42158153051037036029127024023220117314135129141201102552512481380302051047124001015119041321610411382351341310161422045137036016123245123214047231047227023224249032124142201045143049011133132031015114161341351306215614912426123047220281411101601551541512421391301231221171502431421391251222211181174151514113823814016314714612702406415915124813015815305213213115702328044040238126122113112157150035134133132031071470460430260250111100915805315205112802001901703713612512411231481012137511914151617181920212223246Fig.
4Singlemostparsimonioustreeobtainedincladisticanalysisof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,Pachakutej,Ochoa2004andRumikiruOjangurenAflastroetal.
,inpress,underimpliedweights(k=15,length=167,threecharactersordered).
Charactersoptimizedwithacceleratedtransformation.
Homoplasiousandnonhomoplasioustransformationsrespectivelyindicatedbywhiteandblacksquaresalongbranches,withcharacternumbersaboveandcharacterstatesbeloweach.
Zerolengthbranchescollapsedunder'rule1'.
Nodenumbers,usedinsynapomorphylist(Appendix2),indicatedatnodes.
C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176167sameaxisasV1andV3inRumikirubutnotinthesameaxis,formingananglepaessleriOrobothriurusramireziOrobothriurusquewerukanaOrobothriuruscompagnucciiOrobothriuruscalchaquiOrobothriurushuascaranOrobothriurusalticolaOrobothriurusatiquipaOrobothriuruscurvidigitusOrobothriurusampayOrobothriurusfamatinaOrobothriuruswawitaOrobothriurusparvusRumikirulourencoiRumikiruatacamaPachakutejperuvianusPachakutejjuchuichaPachakutejincaPachakutejoscariPachakutejiskayPachakutejcrassimanusBothriuruscordubensisBothriurusasperBothriuruscoriaceusBothriurusbonariensisCentromachetespocockiiCercophoniussulcatusUrophoniustregualemuensisFig.
5Majorityrule(50%)consensusoftreesobtainedfrom26cladisticanalysesof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,Pachakutej,Ochoa2004andRumikiruOjangurenAflastroetal.
,inpress,underequalandimpliedweights,withthreecharactersorderedorunordered.
Nodeswithpaessleri,O.
quewerukanaandO.
tamarugal),formedawell-supportedgroupinallanalyses.
Thesespeciesalsooccurinfairlyclosegeograph-icalproximityinsouthernPeruandnorthernChile(Fig.
1).
OrobothriurusparvusappearstoberelatedtoO.
wawita,andO.
calchaquitoO.
famatina.
However,thesegroupsandseveralotherswithinthegenuswereweaklysupported(Figs2–4)andsome,e.
g.
,theputativerelation-shipbetweenO.
atiquipafromthecoastofsouthernPeruandO.
grismadoifromanisolatedextinctvolcano,over2000minMendoza,Argentina,cannotbeeasilyexplainedbasedontheknowndistributions.
TheplacementofO.
atiquipaamongseveralspeciesfromArgentinamaybeanartefactofmissingentriesinvecharactersofthefemale,whichisunknowninthisspecies.
Thegeneralweaknessoftheseinternalnodessuggeststheneedforadditionalcharacters,e.
g.
,DNAsequences.
AndeanbiogeographyOrobothriurusdisplaysanAndeanpatternofdistribution,mostspeciesoccurringathighaltitudes(over2000–2500mtoamaximumaltituderecordof4910m)fromcentralPerutoArgentina(Fig.
1).
Onlytwospecies,O.
atiquipaandO.
paessleri,occuratloweraltitudes(between700and950m),inlomasformations,endemicplantcommunitiesalongthePacicdesertcoastline,usu-allysituatedonhills,whereoceanfogprovidessufcientmoistureforthedevelopmentofvegetation(Pefaur1981;Dillon2005),insouthernPeru(Ochoa2004,2005).
Onespecies,O.
tamarugal,inhabitstheisolatedforestsofProso-pistamarugoPhil.
(Fabaceae)innorthernChile.
ThegenusmayhaveoriginatedinsouthernPeru,assuggestedbythebasalplacementinthephylogenyofthesouthernPeruvianspeciesO.
ampay,O.
parvusandO.
wawita.
AllOrobothriurusspeciesappeartohavelimitedvagilityandveryrestricteddistributionalranges(Ochoaetal.
2011).
ThepresenceofdifferentspeciesonbothsidesoftheAndessuggeststhattheoriginofthegenuspredatesthemainupliftofthismountainrange.
Andeanorogenystronglyinuencedthedistributionoforganisms(Vuilleu-mier1971;Antonellietal.
2009;Graham2009)andpro-motedsignicantchangesintheclimateofcentralandsouthernSouthAmerica(Vuilleumier1971;Streckeretal.
2007;Inseletal.
2010),perhapscontributingtospeciationprocessesinOrobothriurus.
HighAndeanOrobothriurusspeciescancopewithharshenvironments,wheretheyareexposedtoverylowtemperatureswhenactiveatnight,andcansurvivethewinterathighaltitudeswhereheavysnowfallsarecom-mon.
Therefore,theabsenceofOrobothriurusfromtheAltiplanoandPunaplateaus(Fig.
1),whichhavebeenwellsurveyedforscorpionsbymembersofourresearchgroup,andwhereotherscorpiontaxa(e.
g.
thebothriuridgenus,BrachistosternusPocock,1893)havebeenrecorded,cannotbeexplainedbytheharshconditions.
Onepossiblefactorcouldbetherecent(10millionyearstothepresent)andsustainedvulcanismthataffectedthecentralandsouthernAndeswherevolcanoesareabundant(Trumbulletal.
2006;Fig.
1),therebyalteringthechemicalandorphysi-calcompositionofthesoil,whichisfundamentalforthesurvivaloffossorialandlapidicolousscorpions(Prendini2001).
Thepotentiallylethaleffectsofvulcanism,perhapscombinedwithextensiveoodinginthenorthernandcen-tralAltiplano(Fornarietal.
2001),andthelowvagilityofOrobothriurusspecies,mightexplaintheirabsence.
Indeed,noscorpionshavebeenrecordedfromthecentralareaoftheAltiplano,wherebothvulcanismandoodingoccurred(Acosta&Ochoa2002;OjangurenAflastro2003b).
Otherhigh-altitudescorpionswithgreatervagilityandwiderdistributionalrangesdooccurintheAltiplano,how-ever(Ochoa&Acosta2002;OjangurenAflastro2002,2003b,2004).
FourspeciesofBrachistosternushavebeenrecordedthere,asmallnumbercomparedwithadjacentareasofChileandPeru(OjangurenAflastro2003b;C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176169Ochoa2005;OjangurenAflastro&Mattoni2006;Ojan-gurenAflastro&Scioscia2007;OjangurenAflastroetal.
2007).
BrachistosternusintermediusLo¨nnberg,1902occupiesthesouthernandeasternpartsoftheAltiplano(OjangurenAflastro2004).
BrachistosternustiticacaOchoa&Acosta,2002occursatthenorthernlimitoftheAltiplano,aroundLakeTiticaca(Ochoa&Acosta2002).
Brachistosternusgalia-noaeOjangurenAflastro,2002isrestrictedtotheslopesofSajamaVolcanoinBolivia,atveryhighaltitudesabove4300m.
BrachistosternuspiacentiniiOjangurenAflastro,2003alsooccursontheslopesofSajamaVolcano,andalongthewesternmarginoftheAndesinChile,inaregionthatdoesnotbelongtotheAltiplano(OjangurenAflastro&Ramrez2009).
AllfourBrachistosternusspeciesarerelatedtootherhigh-altitudespeciesoccurringoutsidetheAltiplano(OjangurenAflastro&Ramrez2009),suggest-ingthatspeciationonorneartheAltiplanofollowedrecentcolonizationeventsfromtheAndes.
ThecentralAndes(PeruvianandBolivianAltiplano)experiencedasurfaceupliftof2300–3400mduringthepast10.
7millionyears(sincetheLateMiocene),termedthe'Quechua'phase(Sebrieretal.
1988;Gregory-Wodzicki2000;Gonzalez&Pffner2012).
Thisphasewasaccompaniedbyintensiedvulcanism(Trumbulletal.
2006).
ThemaindiversicationofOrobothriurusprobablyoccurredmuchearlier(ca.
25millionyearsago),whentheAndeswereabout3000mhigh,andvolcaniceventslessfrequent(Gregory-Wodzicki2000;Trumbulletal.
2006).
Thespeciesfromthelomasformationscouldhaveevolvedmuchmorerecently,perhapsduringthePliocene(5–1.
8millionyears).
ManyplantsandanimalsendemictolomasformationshavecloserelativesintheAndesandarethoughttohaveevolvedfromdispersalandsubsequentisolationassociatedwithrecentexpansionsandcontrac-tionsoftheAndeanvegetation(Herrera1930;Dillon2005;Palmaetal.
2005).
Molecular-basedgeographicalstudiesareneededtoconrmwhethertheseOrobothriurusspeciesevolvedinasimilarmanner.
Homoplasyingenitalicvs.
somaticcharactersIthasbeenarguedthatthegenitaliccharactersofarthro-podsevolverelativelyrapidly,becauseofstrongsexualselection,andhavelowphylogeneticinertia(Arnqvist&Rowe2002;Eberhard2004),implyingthatgenitaliccharacterscouldbeveryhomoplasticandthereforemis-leading,oratleastuninformative,inphylogeneticstudies.
Song&Bucheli(2010)demonstratedempirically,using41publisheddatasets,thatgenitalicandsomaticcharactershavesimilarlevelsofhomoplasyininsects,butcompari-sonsamongarachnidsarelacking.
Wecomparedthelevelsofhomoplasyamonggenitalicandsomaticcharactersinbothriuridscorpions,whichexhibitsomeofthegreatestdiversityinmalegenitalia(i.
e.
hemispermatophore)amongscorpions(Maury1980;Ojan-gurenAflastro2005;Peretti2010).
CharactersfromthemalegenitaliashowedhigherCIandRIindicesthansomaticcharacters,suggestingthatgenitaliccharactersofarachnidsareatleastasinformativeassomaticcharacters,ifnotmoreso.
AsconcludedbySong&Bucheli(2010),genitaliccharacterscontainastrongphylogeneticsignal,despiteahighlevelofmorphologicalvariation.
WeagreefurtherwithSong&Bucheli(2010)thatthecompositenatureofgenitaliarendersgenitaliccharactersystemsphylogeneticallyinformativeatmultiplelevelsinthetaxonomichierarchy.
Differentlevelsofinformationareconstrainedbythefunctionalmorphologyofthestruc-turesinquestion.
Inscorpions,whichinseminatefemalesbymeansofaspermatophoreattachedtothesubstrate,structuresthatperformasimilar,mechanicallyconstrainedfunction,e.
g.
,thelaminaandthebasalportion,areprobablyunderstrongstabilizingselection(Peretti2010).
Thesepartsofthespermatophore(anditscomponenthemispermatophores)wouldthereforebeexpectedtoexhi-bitlimitedvariationamongcloselyrelatedtaxa,insteadprovidingphylogeneticinformationfordeepernodes,andthatisindeedthecase.
Incontrast,structuresthatmakecontactwiththefemalegonoporeandgenitalopercula,e.
g.
,theloberegionandsurroundingareasofthesperma-tophore,areprobablyunderstrongsexualselection(Eber-hard1985;Peretti2010),andwouldthereforebeexpectedtoprovidephylogeneticinformationamongcloselyrelatedtaxaand,indeed,itisthesestructuresinwhichthegreat-estmorphologicaldiversityisobserved(Peretti2003).
Itisalsointerestingtonotethattheleasthomoplasticsomaticcharacterspertainmostlytothecheliceraeandpedipalps(characters9–21,Fig.
6),usedbymalescorpionstograspandsecurefemalesduringmating(Polis&Sissom1990;Carreraetal.
2009).
Bothappendagesmaythereforealsobeunderstrongstabilizingselectionandconsequentlyexhibitlesshomoplasythanotherpartsofthesomaticmorphology.
AcknowledgementsWearegratefultoIvanBenoit(CorporacionNacionalForestaldelGobiernodeChile,CONAF)forassistancewithobtainingpermitstocollectscorpionsinChileanNationalParks,andtotheCONAFstaffatLlanosdeChalle,PandeAzucarandPampadelTamarugalNationalParksforlogisticalassistance;totheSecretaradeMedioAmbiente,GobiernodeMendoza,andtheAdministraciondeParquesNacionales,forissuingpermitstocollectscor-pionsinArgentina;andtotheDireccionGeneralForestalydeFaunaSilvestre,MinisteriodeAgriculturadePeru(ex-NRENA)forpermitstocollectscorpionsinPeru.
WeOrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1702012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176thankJohnAchicahuala,JuanCarlosChaparro,LuisCom-pagnucci,JuanFlores,RobertoGutierrez,PaulaKorob,JuanJoseMartinez,OscarMujica,DanielMuniz,WilliamsParedes,LuisPiacentini,JaimePizarro,EliasPonce,AaronQuiroz,ClaraRimondino,GorkyValencia,J.
L.
Velasquez,M.
Vivanco,UlrichZanabriaandHoracioZeballosforassistingusintheeld.
Wethankthefollowingcuratorsandcollectionsmanagersforprovidingaccesstoorloaningmaterialfromthecollectionsintheircare:LuisAcosta,CatedradeDiversidadAnimalI,UniversidadNacionaldeCordoba,Argentina;PetraSierwald,FieldMuseumofNaturalHistory,Chicago,U.
S.
A.
;LuisGrossoandMarc-elaPeralta,InstitutoMiguelLillo,Tucuman,Argentina;ArielCamousseight,MuseoNacionaldeHistoriaNatural,Santiago,Chile;OlinthoAguilar,MuseodeHistoriaNatu-ral,UniversidadNacionaldeSanAntonioAbaddelCusco,Peru;DianaSilva,MuseodeHistoriaNatural,UniversidadNacionalMayordeSanMarcos,Lima,Peru;JorgeArtigas,MuseodeZoologa,UniversidaddeConcepcion,Chile;RicardoPinto-da-Rocha,MuseudeZoologia,UniversidadedeSaoPaulo,Brazil;AdrianoKury,MuseuNacional,RiodeJaneiro,Brazil.
WethanktheWilliHennigSocietyformakingTNTprogramfreelyavailableandPetersonL.
Lopes(UniversidadedeSaoPaulo)forprovidingsomeofthescriptsusedintheanalyses.
WethankSteveThurston,AmericanMuseumofNaturalHistory,NewYork,forassistancewithpreparingthegures,andtwoanonymousreviewersforconstructivecommentsonanearlierversionofthismanuscript.
ThisresearchwassupportedbygrantsfromtheConsejoNacionaldeInvestigacionesCientcasyTecnicas,Argentina(CONICET)toC.
I.
M.
andA.
A.
O.
A.
;aGenomicsPostdoctoralResearchFellowshipfromtheAMNHtoC.
I.
M.
;aKalbeischPostdoctoralResearchFellowshipfromtheAMNH,andafellowshipfromtheFundacaodeAmparoa`PesquisadoEstadodeSaoPaulo,Brazil(FAPESP201000018-9)toJ.
A.
O.
FieldworkinArgentina(2006,2007)wasnanciallysupportedbyaCONICETgrantPIP6502toA.
A.
O.
A.
FieldworkinChilewasnanciallysupportedbythefollowingsources:C.
I.
M.
,J.
A.
O.
andL.
P.
(2003)fromU.
S.
NationalScienceFoundationgrantEAR0228699toL.
P.
;C.
I.
M.
,J.
A.
O.
andA.
A.
O.
A.
(2005)fromtheAMNH;A.
A.
O.
A.
(2006)fromCONICETgrantPIP6502.
FieldworkinPeru(2008)wasfundedinpartbyaKalbeischPostdoctoralResearchFellowshipfromtheAMNHtoJ.
A.
O.
ReferencesAcosta,L.
E.
&Ochoa,J.
A.
(2000).
NuevaespeciedeOrobothriurus,Maury,delPeru.
(Scorpiones,Bothriuridae).
RevueArachnologique,13,135–144.
Acosta,L.
E.
&Ochoa,J.
A.
(2001).
TwonewspeciesofOrobothriurusMaury1976,fromArgentinaandPeru,withcommentsonthesystematicsofthegenus(Scorpiones,Bothriuridae).
InV.
Fet&P.
A.
Selden(Eds)Scorpions2001.
InMemoriamGaryA.
Polis(pp.
203–214).
BurnhamBeeches,Bucks:BritishArachnologicalSociety.
Acosta,L.
E.
&Ochoa,J.
A.
(2002).
Listadelosescorpionesbolivianos(Chelicerata:Scorpiones),connotassobresudistribucion.
RevistadelaSociedadEntomologicaArgentina,61,15–23.
Antonelli,A.
,Nylander,J.
A.
A.
,Persson,C.
&Sanmartn,I.
(2009).
TracingtheimpactoftheAndeanupliftonNeotropical01234567891011121314151617181920212223242526272829303132333435363738394041424344454647484950515253545556575859606162636400.
20.
40.
60.
81CIRICharacterPigm.
ExternalmorphologyExternalmorphologyHemispermatophoreTrichob.
IndexvalueFig.
6Consistencyindices(CI,blacksquares)andretentionindices(RI,greytriangles)ofcharactersusedincladisticanalysisof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress,calculatedonmostparsimonioustreeobtainedunderimpliedweights(k=15).
Abbreviations:Pigm.
,pigmentation;Trichob.
,trichobothria.
C.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176171plantevolution.
ProceedingsoftheNationalAcademyofSciences,106,9749–9754.
Arnqvist,G.
&Rowe,L.
(2002).
Correlatedevolutionofmaleandfemalemorphologiesinwaterstriders.
Evolution,56,936–947.
Bremer,K.
(1994).
Branchsupportandtreestability.
Cladistics,10,295–304.
Carrera,P.
,Mattoni,C.
I.
&Peretti,A.
V.
(2009).
Cheliceraeasmalegraspingorgansinscorpions:sexualdimorphismandassociatedbehaviour.
Zoology,112,332–350.
Coddington,J.
&Scharff,N.
(1994).
Problemswithzero-lengthbranches.
Cladistics,10,415–423.
Dillon,M.
O.
(2005).
SolanaceaeoftheLomasformationsofCoastalPeruandChile.
InV.
Hollowell,T.
Keating,W.
Lewis&T.
Croat(Eds)AFestschriftforWilliamG.
D'Arcy:TheLegacyofaTaxonomist(pp.
131–155).
MonographsinSystematicBotanyfromtheMissouriBotanicalGarden,104.
Eberhard,W.
G.
(1985).
SexualSelectionandAnimalGenitalia.
Cambridge:HarvardUniversityPress.
Eberhard,W.
G.
(2004).
Male-femaleconictandgenitalia:failuretoconrmpredictionsininsectsandspiders.
BiologicalReviewsoftheCambridgePhilosophicalSociety,79,121–186.
Farris,J.
S.
(1970).
MethodsforcomputingWagnertrees.
SystematicZoology,19,83–92.
Farris,J.
S.
(1989).
Theretentionindexandtherescaledconsistencyindex.
Cladistics,5,417–419.
Fornari,M.
,Risacher,F.
&Feraud,G.
(2001).
DatingofpaleolakesinthecentralAltiplanoofBolivia.
Palaeogeography,Palaeoclimatology,Palaeoecology,172,269–282.
Gajardo,R.
(1995).
LavegetacionnaturaldeChile.
Clasicacionydistribuciongeograca.
Santiago:Universitaria.
Goloboff,P.
A.
(1993).
Estimatingcharacterweightsduringtreesearch.
Cladistics,9,83–91.
Goloboff,P.
A.
(1997).
Self-weightedoptimization:treesearchesandcharacterstatereconstructionsunderimpliedtransformationcosts.
Cladistics,13,225–245.
Goloboff,P.
A.
&Farris,J.
(2001).
Methodsforquickconsensusestimation.
Cladistics,17,S26–S34.
Goloboff,P.
A.
,Farris,J.
S.
,Ka¨llersjo¨,M.
,Oxelman,B.
,Ramrez,M.
J.
&Szumik,C.
(2003).
Improvementstoresamplingmeasuresofgroupsupport.
Cladistics,19,324–332.
Goloboff,P.
A.
,Farris,J.
S.
&Nixon,K.
C.
(2008).
TNT,afreeprogramforphylogeneticanalysis.
Cladistics,24,774–786.
Availableviahttp://www.
zmuc.
dk/public/phylogeny/TNT.
Gonzalez,L.
&Pffner,O.
A.
(2012).
MorphologicevolutionofthecentralAndesofPeru.
InternationalJournalofEarthSciences,101,307–321.
Graham,A.
(2009).
TheAndes:ageologicaloverviewfromabiologicalperspective.
AnnalsoftheMissouriBotanicalGarden,96,371–385.
Gregory-Wodzicki,K.
M.
(2000).
UplifthistoryoftheCentralandNorthernAndes:areview.
GeologicalSocietyofAmericaBulletin,112,1092–1105.
Herrera,F.
L.
(1930).
Lavegetaciondelacostaperuana.
RevistaUniversitaria,Cusco,59,71–79.
Honetschlager,L.
D.
(1965).
Anewmethodforhuntingscorpions.
TurtoxNews,43,69–70.
Insel,N.
,Poulsen,C.
J.
&Ehlers,T.
A.
(2010).
InuenceoftheAndesMountainsonSouthAmericanmoisturetransport,convection,andprecipitation.
ClimateDynamics,35,1477–1492.
Ka¨llersjo¨,M.
,Albert,V.
A.
&Farris,J.
S.
(1999).
Homoplasyincreasesphylogeneticstructure.
Cladistics,15,91–94.
Kluge,A.
G.
&Farris,J.
S.
(1969).
QuantitativephyleticsandtheevolutionofAnurans.
SystematicZoology,18,1–32.
Mattoni,C.
I.
&Acosta,L.
E.
(2005).
AnewspeciesofBothriurusfromBrazil(Scorpiones,Bothriuridae).
JournalofArachnology,33,735–744.
Maury,E.
A.
(1976).
EscorpionesyescorpionismoenelPeruV.
Orobothriurus,unnuevogenerodeescorpionesaltoandinos(Bothriuridae).
RevistaPeruanadeEntomologa,18,14–25.
Maury,E.
A.
(1980).
UsefulnessofthehemispermatophoreinthesystematicsofthescorpionfamilyBothriuridae.
Proceedingsofthe8thInternationalArachnologyCongress(Wien):335–339.
Wien:H.
EgermannVerlag.
Nixon,K.
C.
(1999–2002).
Winclada,version1.
00.
08.
Computersoftwareanddocumentation.
Availableviahttp://www.
cladistics.
com.
Ochoa,J.
A.
(2004).
FilogeniadelgeneroOrobothriurusydescripciondeunnuevogenerodeBothriuridae(Scorpiones).
RevistaIbericadeAracnologa,9,43–73.
Ochoa,J.
A.
(2005).
Patronesdedistribuciondeescorpionesdelaregionandinaenelsurperuano.
RevistaPeruanadeBiologa,12,49–68.
Ochoa,J.
A.
&Acosta,L.
E.
(2002).
Orobothriurusatiquipa,anewbothriuridspecies(Scorpiones)fromLomasinsouthernPeru.
JournalofArachnology,30,98–103.
Ochoa,J.
A.
&Acosta,L.
E.
(2003).
UnanuevaespeciedeOrobothriurusdelSantuarioNacionaldeAmpay,Apurimac,Peru.
RevistaPeruanadeEntomologa,43,1–6.
Ochoa,J.
A.
&OjangurenAflastro,A.
A.
(2007).
SystematicsanddistributionofBrachistosternus(Brachistosternus)ehrenbergii(Gervais,1841),withtherstrecordofstridulationinthisgenusBrachistosternus(Scorpiones:Bothriuridae).
StudiesonNeotropicalFaunaandEnvironment,42,61–69.
Ochoa,J.
A.
,Botero-Trujillo,R.
&Prendini,L.
(2010).
OnthetroglomorphicscorpionTroglotayosicushumiculum(Scorpiones,Troglotayosicidae),withrstdescriptionoftheadults.
AmericanMuseumNovitates,3691,1–19.
Ochoa,J.
A.
,OjangurenAflastro,A.
A.
,Mattoni,C.
I.
&Prendini,L.
(2011).
SystematicrevisionoftheAndeanscorpiongenusOrobothriurusMaury,1976(Bothriuridae),withdiscussionofthealtituderecordforscorpions.
BulletinoftheAmericanMuseumofNaturalHistory,359,1–90.
OjangurenAflastro,A.
A.
(2002).
Brachistosternusgalianoae(Scorpiones,Bothriuridae),unanuevaespeciedeBolivia.
RevistadelMuseoArgentinodeCienciasNaturales,4,104–109.
OjangurenAflastro,A.
A.
(2003a).
UnnuevoOrobothriurus(Scorpiones,Bothriuridae)delaregiondeAtacama,Chile.
RevistaIbericadeAracnologa,7,117–122.
OjangurenAflastro,A.
A.
(2003b).
LasespeciesandinasdeBrachistosternus(Leptosternus),conladescripciondetresnuevasespecies(Scorpiones,Bothriuridae).
RevistaIbericadeAracnologa,8,23–36.
OjangurenAflastro,A.
A.
(2004).
SistematicaydistribuciondeBrachistosternus(Leptosternus)intermediusLo¨nnberg(Scorpiones,Bothriuridae).
PhysisSectionC,59,29–35.
OrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1722012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176OjangurenAflastro,A.
A.
(2005).
EstudiomonogracodelosescorpionesdelaRepublicaArgentina.
RevistaIbericadeAracnologa,11,75–241.
OjangurenAflastro,A.
A.
&Mattoni,C.
I.
(2006).
AnewspeciesofBrachistosternusfromChileancentralAndes(Scorpiones;Bothriuridae).
StudiesonNeotropicalFaunaandEnvironment,41,1–8.
OjangurenAflastro,A.
A.
&Ramrez,M.
J.
(2009).
PhylogeneticanalysisofthescorpiongenusBrachistosternus(Arachnida,Scorpiones,Bothriuridae).
ZoologicaScripta,38,183–198.
OjangurenAflastro,A.
A.
&Scioscia,C.
L.
(2007).
NewdataaboutgenusBrachistosternusinChile(Scorpiones:Bothriuridae).
JournalofArachnology,35,102–113.
OjangurenAflastro,A.
A.
,Agusto,P.
,Pizarro-Araya,J.
&Mattoni,C.
I.
(2007).
TwonewscorpionspeciesofgenusBrachistosternus(Scorpiones:Bothriuridae)fromnorthernChile.
Zootaxa,1623,55–68.
OjangurenAflastro,A.
A.
,FernandezCampon,F.
,Lagos-Silnik,S.
&Mattoni,C.
I.
(2009).
ThegenusOrobothriurusMauryincentralArgentinawiththedescriptionofanewspeciesfromElNevadomountainchaininMendoza(Scorpiones:Bothriuridae).
Zootaxa,2209,28–42.
OjangurenAflastro,A.
A.
,Mattoni,C.
I.
,Ochoa,J.
A.
&Prendini,L.
(inpress).
Rumikiru,n.
gen.
(Scorpiones,Bothriuridae),anewscorpiongenusfromtheAtacamadesert.
AmericanMuseumNovitates.
Palma,R.
E.
,Marquet,P.
A.
&Boric-Bargetto,D.
(2005).
Inter-andintraspecicphylogeographyofsmallmammalsintheAtacamaDesertandadjacentareasofnorthernChile.
JournalofBiogeography,32,1931–1941.
Pefaur,J.
E.
(1981).
CompositionandphenologyofepigeicanimalcommunitiesintheLomasofsouthernPeru.
JournalofAridEnvironments,4,31–42.
Peretti,A.
V.
(2003).
Functionalmorphologyofspermatophoresandfemalegenitaliainbothriuridscorpions:genitalcourtship,coercionandotherpossiblemechanisms.
JournalofZoology,261,1–19.
Peretti,A.
V.
(2010).
Anancientindirectsexmodel:singleandmixedpatternsintheevolutionofscorpiongenitalia.
InJ.
L.
Leonard&A.
Cordoba-Aguilar(Eds)TheEvolutionofPrimarySexualCharactersinAnimals(pp.
218–248).
Oxford:OxfordUniversityPress.
Polis,G.
A.
&Sissom,W.
D.
(1990).
LifeHistory.
InG.
A.
Polis(Ed.
)TheBiologyofscorpions.
(pp.
161–223).
Stanford,California:StanfordUniversityPress.
Prendini,L.
(2000).
PhylogenyandclassicationoftheSuperfamilyScorpionoideaLatreille1802(Chelicerata,Scorpiones):anexemplarapproach.
Cladistics,16,1–78.
Prendini,L.
(2001).
SubstratumspecializationandspeciationinsouthernAfricanscorpions:theeffecthypothesisrevisited.
InV.
Fet&P.
A.
Selden(Eds)Scorpions2001.
InMemoriamGaryA.
Polis(pp.
113–138).
BurnhamBeeches,Bucks:BritishArachnologicalSociety.
Prendini,L.
(2003a).
AnewgenusandspeciesofbothriuridscorpionfromtheBrandbergMassif,Namibia,withareanalysisofbothriuridphylogenyandadiscussionofthephylogeneticpositionofLisposomaLawrence.
SystematicEntomology,28,149–172.
Prendini,L.
(2003b).
DiscoveryofthemaleofParabuthusmuelleri,andimplicationsforthephylogenyofParabuthus(Scorpiones:Buthidae).
AmericanMuseumNovitates,3408,1–24.
Sebrier,M.
,Lavenu,A.
,Fornari,M.
&Soulas,J.
-P.
(1988).
TectonicsandupliftincentralAndes(Peru,Bolivia,andnorthernChile)fromEocenetopresent.
Geodynamique,3,85–106.
Sissom,W.
D.
,Polis,G.
A.
&Watt,D.
D.
(1990).
Fieldandlaboratorymethods.
InG.
A.
Polis(Ed.
)TheBiologyofScorpions.
(pp.
445–461).
Stanford,CA:StanfordUniversityPress.
Song,H.
&Bucheli,S.
R.
(2010).
Comparisonofphylogeneticsignalbetweenmalegenitaliaandnon-genitalcharactersininsectsystematics.
Cladistics,26,23–35.
Stahnke,H.
L.
(1970).
Scorpionnomenclatureandmensuration.
EntomologicalNews,81,297–316.
Stahnke,H.
L.
(1972).
U.
V.
light,ausefuleldtool.
BioScience,22,604–607.
Strecker,M.
R.
,Alonso,R.
N.
,Bookhagen,B.
,Carrapa,B.
,Hilley,G.
E.
,Sobel,E.
R.
&Trauth,M.
H.
(2007).
TectonicsandclimateofthesouthernCentralAndes.
AnnualReviewsofEarthandPlanetarySciences,35,747–787.
Swofford,D.
L.
&Beagle,D.
P.
(1993).
PAUPphylogeneticanalysisusingparsimony.
Version3.
1User'smanual.
IllinoisNationalHistorySurvey,Champaign,Illinois,USA.
Swofford,D.
L.
&Maddison,W.
P.
(1987).
ReconstructingancestralcharacterstatesunderWagnerparsimony.
MathematicalBiosciences,87,199–229.
Swofford,D.
L.
&Maddison,W.
P.
(1992).
Parsimony,character-statereconstructions,andevolutionaryinferences.
InR.
L.
Mayden(Ed.
)Systematics,HistoricalEcology,andNorthAmericanFreshwaterFishes(pp.
187–223).
Stanford,CA:StanfordUniversityPress.
Trumbull,R.
,Riller,U.
,Oncken,O.
,Scheuber,E.
,Munier,K.
&Hongn,F.
(2006).
Thetime-spacedistributionofCenozoicvolcanisminthesouth-centralAndes:anewdatacompilationandsometectonicimplications.
InO.
Oncken,G.
Chong,G.
Franz,P.
Giese,H.
-J.
Go¨tze,V.
Ramos,M.
Strecker&P.
Wigger(Eds)TheAndes-ActiveSubductionOrogeny.
FrontiersinEarthSciences,Vol.
1,XXII(pp.
29–43).
Berlin,Heidelberg:SpringerVerlag.
Vachon,M.
(1952).
EtudesurlesScorpions.
Alger:InstitutPasteurd'Algerie.
Vachon,M.
(1973)[1974].
Etudedescaracte`resutilisespourclasserlesfamillesetlesgenresdescorpions(Arachnides).
1.
Latrichobothriotaxieenarachnologie.
Siglestrichobothriauxettypesdetrichobothriotaxiechezlesscorpions.
BulletinduMuseumNationald'HistoireNaturelle(Paris)Ser.
3,140,857–958.
Vuilleumier,B.
S.
(1971).
PleistocenechangesinthefaunaandoraofSouthAmerica.
Science,173,771–780.
Wheeler,W.
C.
(1995).
Sequencealignment,parametersensitivity,andthephylogeneticanalysisofmoleculardata.
SystematicBiology,44,321–331.
Appendix1CharacterlistListof65charactersscoredforcladisticanalysisof30bothriuridscorpionspecies,includingallspeciesofOrobothriurusMaury,1976,PachakutejOchoa,2004andRumikiruOjangurenAflastroetal.
,inpress.
DatamatrixinTable1.
Characterstatesarescored0–3,unknown()andinapplicable(-).
MorphologicalterminologyfollowsVachon(1974)fortrichobothrialnomenclature;VachonC.
I.
Mattonietal.
dOrobothriurusphylogenyandbiogeography2012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176173(1952)andPrendini(2000)forpedipalpcarinae,abbrevi-atedasfollows:internomedian(IM),dorsointernal(DI),dorsomedian(DM),dorsoexternal(DE),externomedian(EM),ventroexternal(VE),ventromedian(VM),ventroin-ternal(VI),dorsalpatellarprocess(DPP),ventralpatellarprocess(VPP),digital(D),dorsalsecondary(DS),dorso-marginal(DMA),external(E);amodiedversionofPren-dini(2000,2003b)usedbyOchoaetal.
(2010)fortergite,sterniteandmetasomalcarinae,abbreviatedasfollows:dorsolateral(DL),lateralsupramedian(LSM),lateralmed-ian(LM),lateralinframedian(LIM),ventrolateral(VL),ventrosubmedian(VSM),ventromedian(VM);Mattoni&Acosta(2005)formetasomalmacrosetae;Ochoa(2004)andMattoni&Acosta(2005)forhemispermatophore;Stahnke(1970)forothercharacters.
Charactersfrompreviousanalysesthatcorrespondpartiallyorentirelytothoseinthepresentmatrixareasfollows:O2004=Ochoa(2004);OA&R2009=OjangurenAflastro&Ramrez(2009);P2003=Prendini(2003a).
Pigmentationpattern0.
TergitesI–IV,pigmentation:entirelypigmented(0);pairedspotssublaterally,unpigmentedareamedially(1)[O2004:0;OA&R2009:1].
1.
TergiteVII,pigmentation:entirelypigmented(0);pairedspotssublaterally,unpigmentedareamedially(1);inapplicable(-).
2.
MetasomalsegmentsIIandIII,dorsalsurfaces,pig-mentation:absent,unpigmented(0);subtriangularspotmedially,maybedividedbyunpigmentedlinemedially(1).
3.
MetasomalsegmentsI–III,dorsalsurfaces,pigmenta-tionalongDLcarinae:absent,unpigmented(0);reticulatelines(1).
4.
MetasomalsegmentsIVandV,ventralsurfaces,VMstripe:contiguouswithVLstripeposteriorly(0);notcon-tiguouswithVLstripeposteriorly(1);absent(2).
[OA&R2009:5].
5.
Telsonvesicle,ventralandlateralsurfaces,colour-ation(#):similarto$,pigmented(0);differentfrom$,un-pigmented,withglandular,light-yellowcolouration(1).
Carapace6.
Anteriormargin,shape:sublinearorwithshallowmed-iannotch(0);withweakmedianprojection(epistome)(1).
[OA&R2009:7].
7.
Anteromedianlongitudinalsulcus,length(#):com-plete(0);vestigial(1).
[P2003:4;O2004:1;OA&R2009:8].
Chelicera8.
Movablenger,subdistalteeth,number:one(0);two(1).
[P2003:9;O2004:2;OA&R2009:6].
Pedipalps9.
Femur,length(#):greaterthanthreetimeswidth(0);90°(0);D-DMA-DIcarinaeforminganglepaessleri:Char.
14:10;23:12;45:01Orobothriurustamarugal:Char.
2:10;4:12;24:21;32:01Node2:Char.
19:10;52:10Node3:Char.
9:01;10:10;11:01;43:10;46:10;47:10Node4:Char.
7:01;33:01;34:01;35:01;50:10Node5:Char.
12:01;13:01;22:01;40:02;44:10Node6:Char.
52:01Node7:Char.
8:10;23:02Node8:Char.
30:01;48:01;51:12;59:01;64:01Node9:Char.
5:01;15:01;17:04;18:01;21:01;22:02;25:01;39:01;42:01;43:01;50:02Node10:Char.
24:10;27:10;46:01;47:01;63:01Node11:Char.
38:02;41:01Node12:Char.
40:01Node13:Char.
38:01Node14:Char.
22:01;23:01;30:01;39:01;51:012;60:01Node15:Char.
0:01;1:01;4:01Node16:Char.
3:10;61:12Node17:Char.
2:10;47:12Node18:Char.
24:12;49:01;56:01;62:01Node19:Char.
3:10;35:01Node20:Char.
34:01Node21:Char.
45:01Node22:Char.
6:01;14:01;15:01;31:10;32:10;33:01Node23:Char.
11:01;49:10Node24:Char.
43:10SupportingInformationAdditionalSupportingInformationmaybefoundintheonlineversionofthisarticle:AppendixS1.
Materialexamined.
Pleasenote:Wiley-Blackwellarenotresponsibleforthecontentorfunctionalityofanysupportingmaterialssup-pliedbytheauthors.
Anyqueries(otherthanmissingmaterial)shouldbedirectedtothecorrespondingauthorforthearticle.
OrobothriurusphylogenyandbiogeographydC.
I.
Mattonietal.
1762012TheAuthorsdZoologicaScripta2012TheNorwegianAcademyofScienceandLetters,41,2,March2012,pp160–176