The Egyptian fruit bat Rousettus aegyptiacus (Chiroptera: Pteropodidae) in the Palaearctic: Geographical variation and taxonomic status


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The Egyptian fruit bat Rousettus aegyptiacus (Chiroptera: Pteropodidae) in the Palaearctic: Geographical variation and taxonomic status

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The Egyptian fruit bat Rousettus aegyptiacus (Chiroptera: Pteropodidae) in the Palaearctic: Geographical variation and taxonomic status
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Biologia
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Benda, Petr
Vallo, Peter
Hulva, Pavel
Horáček, Ivan
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De Gruyter
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English

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Rousettus ( local )
Morphometrics ( local )
Mtdna ( local )
Taxonomy ( local )
Distribution ( local )
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serial ( sobekcm )

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Abstract:
Two metrically defined subspecies have traditionally been recognised within the Palaearctic distribution range of the Egyptian fruit bat Rousettus aegyptiacus; the larger R. a. aegyptiacus in Egypt and the northern part of the Middle East and the smaller R. a. arabicus in the southern Middle East. An extensive material of R. aegyptiacus from all parts of this area, i.e. the Levant (incl. Turkey and Cyprus), Egypt (incl. Sinai), northern Sudan, Yemen, Oman, Iran, and Pakistan, as well as comparative samples from its sub-Saharan range, were tested using both morphological and genetic approaches in order to revise the species’ taxonomic status. The results indicated two possible processes, depending on the method used. Genetic analysis of the mitochondrial genome (nd1 and cytb) indicated low variation (< 2.0% of genetic distance) and lack of geographical structure while morphometric analysis indicated significant metric differences. Two basic size morphotypes were found within the Palaearctic range, with a rather mosaic-like geographical distribution and a lack of clear size distinction between the two categories, though intermediate types were detected. Thus, we suggest that all Palaearctic populations of R. aegyptiacus represent one form, the nominotypical subspecies, which is uniform in genetic traits but plastic in metric traits.
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Biologia, Vol. 67, no. 6 (2012-10-19).

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Biologia 67 /6:12301244,2012 SectionZoology DOI:10.2478/s11756-012-0105-yTheEgyptianfruitbat Rousettusaegyptiacus (Chiroptera: Pteropodidae)inthePalaearctic:Geographicalvariation andtaxonomicstatusPetr Benda1,2,Peter Vallo3,Pavel Hulva2&Ivan Horek21DepartmentofZoology,NationalMuseum(NaturalHistory),Vclavsknm. 68,CZ– 11579 Praha 1 ,CzechRepublic; e-mail:petr benda@nm.cz2DepartmentofZoology,FacultyofScience,CharlesUniversity,Vinin 7 ,CZ– 12844 Praha 2 ,CzechRepublic3InstituteofVertebrateBiology,AcademyofSciencesoftheCzechRepublic,v.v.i.,Kvtn 8 ,CZ60365 Brno,Czech Republic Abstract: Twometricallyde“nedsubspecieshavetraditionallybeenrecognisedwithinthePalaearcticdistributionrange oftheEgyptianfruitbat Rousettusaegyptiacus ;thelarger R.a.aegyptiacus inEgyptandthenorthernpartoftheMiddle Eastandthesmaller R.a.arabicus inthesouthernMiddleEast.Anextensivematerialof R.aegyptiacus fromallparts ofthisarea,i.e.theLevant(incl.TurkeyandCyprus),Egypt(incl.Sinai),northernSudan,Yemen,Oman,Iran,and Pakistan,aswellascomparativesamplesfromitssub-Saharanrange,weretestedusingbothmorphologicalandgenetic approachesinordertorevisethespeciestaxonomicstatus.Theresultsindicatedtwopossibleprocesses,dependingon themethodused.Geneticanalysisofthemitochondrialgenome(nd1andcyt b )indicatedlowvariation( < 2.0%ofgenetic distance)andlackofgeographicalstructurewhilemorphometricanalysisindicatedsigni“cantmetricdierences.Twobasic sizemorphotypeswerefoundwithinthePalaearcticrange,witharathermosaic-likegeographicaldistributionandalack ofclearsizedistinctionbetweenthetwocategories,thoughin termediatetypesweredetected.Thus,wesuggestthatall Palaearcticpopulationsof R. aegyptiacus representoneform,thenominotypicalsubspecies,whichisuniformingenetic traitsbutplasticinmetrictraits. Keywords: Rousettus ;morphometrics;mtDNA;taxonomy;distributionIntroduction TheEgyptianfruitbat, Rousettusaegyptiacus (Geoffroy,1810),istheonlymemberofthePteropodidae familythathasthelargepartofitsdistributionrange inthePalaearcticregion(Fig.1),andtheonlyone whosetypelocalityliesinthePalaearctic.Consideringthedistributionrange, R . aegyptiacus ismostlyan Afro-tropicalspecies,whichreachesthesouthwestern partofthePalaearcticintheMiddleEast(Juste& Iba nez1993;Bergmans1994;Kwiecinski&Griths 1999).Insub-SaharanAfrica,itoccupiesareasaround theGulfofGuineafromSenegaltowesternAngola (includingsomeislandsintheGulf)andsavannahregionsofsouthandeastAfricafromtheCapetoEritrea.InthePalaearctic, R . aegyptiacus occursinEgypt andnorthernSudanaswellasinbroadareasalong theseacoastsoftheMiddleEast,fromsouth-western TurkeyandCyprusalongtheLevantineandArabian shoresuptosouthernIranandPakistan(Bergmans 1994;Bendaetal.2011).ThePalaearcticrange(here consideredasEgypt,northernSudanandthewhole MiddleEast,aswellassouthernPakistan)isreported tobeisolatedfromthesub-SaharanAfricanrangeby theRedSeaanddesertareasofnorth-easternAfrica. Thenorth-easternMediterraneanrepresentsthenorthernmarginofthespeciesdistributionrange(Harrison&Bates1991;Bergmans1994;Bendaetal.2006, 2011). ThespeciespresenceintheMiddleEasthastraditionallybeenseenassomethingofanenigmaasitis theonlyoshootofthefamilyfoundbeyondthetropics.Thespeciesoccurrencedependsuponcontinuous all-year-roundavailabilityoffruit,whichisstrictlyrelatedtoanthropogenicplantcultivationoverthevast majorityoftheMiddleEastandhasundergonenumerousdramaticspatialandtemporal”uctuations.Despite this,thespeciesisreportedasexhibitingclearpatterns ofgeographicphenotypevariationthatcorrespondwith thevariationpatterndisplayedovertherestofitsrange (Bergmans1994;Kwiecinski&Griths1999). R . aegyptiacus isregardedasapolytypicspecies, withfoursubspeciesrecognisedwithinitscontinentalAfro-Asianrange,i.e., R . a . aegyptiacus (Georoy, 1810), R . a . leachii (Smith,1829), R . a . unicolor (Gray, 1870),and R . a . arabicus Anderson,1902(Eisentraut 1959;Hayman&Hill1971;Bergmans1994;Koopman 1994;Kwiecinski&Griths1999;Simmons2005).In addition,Juste&Iba nez(1993)describedtwosubspeciesfromtwoislandsintheAfricanGulfofGuinea;c2012InstituteofZoology,SlovakAcademyofSciences

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Variationof Rousettusaegyptiacus inthePalaearctic 1231 R . a . princeps fromPrincipeand R . a . tomensis from S aoTomé. Traditionally,twosubspeciesoftheEgyptianfruit bathavebeenconsideredasmeetingintheMiddle East(Eisentraut1959;Bergmans1994),basedonevidenceoftwodistinctsizecategoriesoccurringintwo distinctranges:thelargerform, R . a . aegyptiacus ,living inEgyptandtheeasternMediterranean(typelocality: GreatPyramidofGiza,Egypt;restrictedbyAnderson1902),andasmallerform, R . a . arabicus ,livingin southernArabiaandsouthernpartsofIranandPakistan(typelocality:LahejnearAden,Yemen)(Eisentraut1959;Harrison1964;Corbet1978;Harrison& Bates1991;Bergmans1994;Koopman1994;Kwiecinski&Griths1999;Horáeketal.2000).Bendaetal. (2008),however,indescribingthe“rstevidenceof R . aegyptiacus populationsindesertoasesofSinai,found thatSinaiticfruitbatsconformedinbodysizealmost completelywithcomparativesamplesfromYemenand Iran( R . a . arabicus ),andonlypartlyoverlappedwith thosefromtheLevantandLowerEgypt( R . a . aegyptiacus ).Bendaetal.(2010)reportedthatthedimensionsofsouth-Jordanian R . aegyptiacus sampleswere intermediatebetweenthetwoMediterranean(Levantine)anddesert(Sinaitic)size-morphotypesand,therefore,theysuggestedthatacontinuousclineshiftinsize existedbetweentherespectiveregionsoftheeastern Mediterranean.Ontheotherhand,preliminarycomparisonsofmitochondrialgenes,thoughbasedonafew samplesonly(Bendaetal.2007),showedverysmall geneticdierencesbetween R . aegyptiacus populations overthewholeMiddleEast(fromCyprustoIran).This tendstoindicateadiscontinuousgeographicalarrangementofvariationcategories,ratherthantwometrically andgeographicallydistinctsubspecies sensu Bergmans (1994)revision.Hence,Bendaetal.(2008)speculated thatrearrangementsinbodysizecouldhaveoccurred overthecourseofjustafewgenerationsandrepresentadaptivechangesratherthanevidenceofphylogeneticdierencesbetweengeogr aphicallyseparatedpopulations. Overthelasttwodecades,numerousnewrecords of R . aegyptiacus havebeenobtainedfromtheMiddle Eastandnorth-eastAfrica(Bendaetal.2011),which providesanopportunitytore-examinepatternsofgeographicvariationandinterrelationshipsbetweenlocal populationsinthePalaearctic.Usingbothmorphometricandgeneticapproaches,thepresentpaperreports onthe“rstresultsofthisprocess,withparticularfocusonthetaxonomicarrang ementofthespecieswithin thispartofitsdistributionrange.Materialandmethods Morphologicalanalysis Wecomparedmorphological(morphometric)skullandforearmdatafrommuseumspecimenscoveringallpartsofthe Palaearcticdistributionrangeof R . aegyptiacus (Fig.1),includingmostofthetypematerialandseveralcomparative specimens/taxafromthespeciessub-Saharanrangeand fromitsOrientalpromontory(Pakistan).SeeAppendix1 foradetailedlistofthematerialexamined(seebelowfor abbreviationsused). Specimensweremeasuredwithamechanicalcalliper usingstandardmethodsaccordingtoe.g.Bendaetal. (2004).Statisticalanalysisofthemorphometricdatawas performedusingStatistica6.0software.Principalcomponentanalysis(PCA)wasusedasatestoftheimportance ofskullsizeand,inparticular,skulldimensionsforintraspeci“cvariation.Sexualdimorphismwastestedusing thet-testandone-wayanalysisofvariance(ANOVA).Clusteranalysis(unweightedpair-groupaverage,Euclideandistances)wasusedtoevaluatesizesimilaritiesamongrepresentativesofparticularpopulationswithinthedistribution range. Moleculargeneticanalysis Moleculargeneticanalysiswasbasedon79 R . aegyptiacus samplesfromthenorthernandsouthernpartsofthe Palaearcticrange(onetofoursamplespersite),with44 samplesfrom27sitesinthenorthernregions(Egypt,northernSudanandtheLevant,includingCyprus,Turkeyand Sinai)and35samplesfrom21sitesinthesouthernregions (Yemen,OmanandIran).SeveralsamplesfromAfrican sub-Saharanpopulationswerealsoadded(17samplesfrom specimensoriginatinginSenegal,Ghana,Gabon,Ethiopia, KenyaandMalawi).SeeAppendix2fordetaileddataand detailsoforiginforthematerialexamined. TotalgenomicDNAwasextractedfromthetissue samplesusingtheNucleospin96-welltissuekit(MachereyNagel,USA)ortheJetQuickDNAextractionkit(Genomed, Germany).Nicotine-amiddehydrogenase1(nd1)andcytochrome b (cyt b ),twomitochondrial(mt)genes,wereampli“edviaPolymeraseChainReaction(PCR)usingER65 andER66primers(Petitetal.1999)andL14724and H15915primers(Irwinetal.1991),respectively.EachPCR volumecontained12.5 µ lCombiPPPmix(TopBio,Czech Republic),2 µ lofa10 µ Msolutionofeachprimer,and 2.5 µ lofisolatedDNAsolution.ThePCRbeganwithinitial denaturationat94Cfor3min,followedby35cyclesofdenaturationfor40sat94C,annealingfor40sat58C(nd1) or50C(cyt b ),extensionfor90sat65C,witha“nalextensionat65Cfor5min.Ampli“edproductswerepuri“ed usingeithertheJetQuickPCRpuri“cationkit(Genomed, Germany)orcommerciallybyMacrogenInc.(Seoul,Korea).ThelattercompanyalsosequencedallPCRproducts withtherespectiveforwardprimeronanABI3137XLsequencerusingtheBigDyesequencingkit.Sequenceswere checkedforambiguitiesandeditedinSequencher(Gene Codes,USA),thenalignedmanuallyinBioEdit(Hall1999). Forreconstructionofphylogenetictrees,sequenceswerenarrowedtouniquehaplotypes.Sequencesof Rousettuslanosus Thomas,1906,and Lissonycterisangolensis (Bocage,1898) wereusedasanoutgroup(Appendix2).Phylogenetictrees werecomputedinPAUP*4.10b(SinauerAssociates,USA) usingmaximumparsimony(MP)andneighbour-joining (NJ).CharacterswereequallyweightedandunorderedduringMPanalysis,whiledistanceinNJanalysiswasbased ontheKimuratwo-parameter(K2p)evolutionarymodel (Kimura1980).K2pdistanceswerealsousedforexpressinggeneticdierenceamonghaplotypes.Supportforthe branchingpatternwasassessedthrough1000 × nonparametricbootstrappingduringMPanalysis.Relationshipsamong R . aegyptiacus sequenceswerealsoexploredusingthenetworkapproachinNetwork(Fluxus,USA)inordertorecoverstructureamongcloselyrelatedhaplotypesthatmay

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1232 P. Benda etal. Fig.1.Mapof Rousettusaegyptiacus recordsinthePalaearctic, sensu Bendaetal.(2011).Blackdots…sitesoforiginofthematerial examined,greydots…otherrecords. beinancestor-descendantrelationshipsandwhoseposition inphylogenetictreesremainedunresolved(Posada&Crandall2001). Abbreviations Dimensions: LAt=forearmlength(includingwrist);LCr =greatestlengthofskull;LCb=condylobasallength;LaZ =zygomaticwidth;LaI=widthofinterorbitalconstriction;LaP=widthofpostorbitalconstriction;LaN=neurocraniumwidth;AN=neurocraniumheight;CC=rostral widthbetweenuppercanines(incl.);M 2 M 2 =rostralwidth between2nduppermolars(incl.);CM 2 =lengthofupper tooth-rowbetweencanineand2ndmolar(incl.);LMd= condylarlengthofmandible;ACo=heightofcoronoidprocess;CM 3 =lengthoflowertooth-rowbetweencanineand 3rdmolar(incl.). Collections: BMNH=NaturalHistoryMuseum,London, UnitedKingdom;IVB=InstituteofVertebrateBiology, AcademyofSciencesoftheCzechRepublic,Brno,Czech Republic;JOC=JánObuchPrivateCollection,Blatnica, Slovakia;MSNG=CivilMuseumofNaturalHistoryGiacomoDoria,Genoa,Italy;NMP=NationalMuseum(NaturalHistory),Prague,CzechRepublic;ZFMK=Alexander KoenigZoologicalInstituteandMuseum,Bonn,Germany. Otherabbreviations: A=alcoholicpreparation;B=skin (balg);df=degreesoffreedom;f=female; F = F -valueof ANOVA;m=male;M=mean;max,min=maximumand minimumrangemargins; P =probability(signi“cance);PC =principalcomponent;S=skull;SD=standarddeviation; Sk=skeleton; T = t -valueof t -test. Results Morphometricanalysis Morphometriccomparisonsrevealedtwobasicsizemorphotypesamongthepopulationsamplesexamined(Table1).Thedimensionrangesofthesegroupsoverlappedonlypartially,withthelargestskullmeasurementsoverlappingbylessthan50%(Fig.2).Larger representatives(LAt86.6…98.1mm;LCr40.7…46.6mm) werefoundintwoseparateareas:theLevant(Turkey, Cyprus,SyriaandLebanon)andcontinentalEgypt (Cairoregion,theNilevalleyandWesternOases).The largestofall(LCr > 46.0mm)werefoundamong samplesfromtheWesternOasisofDakhlainEgypt. Smallerrepresentatives(LAt81.5…95.4mm;LCr38.0… 43.7mm)werealsofoundintwolargelyseparatedgeographicalareas:thesouthernMiddleEast(Yemen, Oman,IranandPakistan)andSinai.Smallestspecimens(LCr < 38.5mm)weredocumentedfromsouthcentralIran(Isin).Thedimensionsofsamplesfrom Jordanwereintermediatebetweenthesetwogroups (LAt86.4…96.5mm;LCr40.2…44.8mm).Comparativesamplesetsfromsub-SaharanAfrica(Ethiopiaand Senegal)werepositionedsimilarlytotheintermediate samplesfromJordan,althoughslightlylargerintheir externaldimensions(LAt88.3…100.7mm;LCr39.7… 44.2mm; n =12). Variationinsexualdimorphismwasnotedwithin

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Variationof Rousettusaegyptiacus inthePalaearctic 1233 Table1.Forearmandskulldimensions(inmm)andPC1valuesoftheexaminedsamplesetsof Rousettusaegyptiacus fromthe Palaearcticandresultsofsexualdimorphismanalysiswithinthesets. EgyptLevant n M minmaxSDdf FTPn M minmaxSDdf FTP LAt25 92.36 86.696.62.986231.691.301…16 94.53 90.098.12.321140.03…0.173… LCr45 44.34 41.4746.571.290283.881.970…19 43.51 40.6845.191.130160.430.654… LCb44 42.79 39.6444.861.253273.671.917…18 41.87 39.2443.571.129150.460.677… LaZ44 27.02 24.2329.271.270282.241.498…19 26.88 24.6329.261.050162.021.420… LaI24 8.63 7.739.510.514140.080.292…18 8.37 7.939.080.30115 4.962.227 * LaP25 7.87 6.878.870.555152.571.603…16 7.78 7.228.760.42413 5.862.420 * LaN44 17.45 16.8218.340.400282.251.501…19 17.31 16.8317.930.323160.090.300… AN45 13.26 12.3114.280.532281.391.177…19 13.17 11.6413.930.546160.010.078… CC44 8.83 8.219.710.43328 8.762.959 *18 8.89 8.429.680.40116 5.432.329 * M 2 M 2 40 13.30 12.3514.080.426270.930.963…17 12.90 12.2913.500.380150.120.353… CM 2 44 16.87 15.6118.040.53527 5.252.290 *19 16.54 15.1517.620.64216 5.762.400 * LMd45 34.47 31.9736.320.997282.501.581…19 33.87 31.9835.320.955161.031.012… ACo45 16.03 14.5417.670.76728 6.392.529 *18 15.13 13.2716.630.937150.040.202… CM 3 44 18.38 17.0819.850.622280.400.632…18 18.20 17.5619.200.464151.491.222… PC145 –0.980 …2.0390.2620.59128 4.48–2.117 *19 –0.542 …1.5110.4580.593164.412.099… SinaiJordan n M minmaxSDdf FTPn M minmaxSDdf FTP LAt14 91.56 87.095.42.358123.261.805…11 91.73 86.496.53.29093.041.742… LCr13 41.27 38.9143.661.32911 4.872.208 *10 42.44 40.2444.821.4818 14.003.742 * LCb13 39.55 37.5341.471.173114.742.176…10 40.80 38.6742.841.4278 17.454.177 ** LaZ13 24.95 23.4827.330.920114.052.014…10 25.85 24.8926.870.7198 21.274.612 ** LaI13 7.87 7.198.590.403112.541.594…10 8.30 7.839.170.37980.000.045… LaP13 7.39 6.617.940.360111.221.106…9 7.87 7.448.710.35870.80…0.896… LaN13 16.47 16.0217.910.530112.391.546…10 15.45 6.5417.713.47680.030.162… AN13 12.53 11.5613.240.453113.711.926…10 13.18 12.4514.570.65280.160.397… CC13 8.25 7.828.840.29611 6.402.530 *10 8.46 7.729.020.4608 9.723.118 * M 2 M 2 13 12.38 11.8413.300.42511 5.152.269 *10 12.62 12.1113.120.3808 11.323.365 * CM 2 13 16.02 14.9116.990.58311 6.362.522 *10 16.09 15.3117.070.5878 25.305.030 ** LMd13 32.12 30.0833.821.04811 7.372.714 *10 32.70 30.7534.371.2378 11.113.333 * ACo13 14.89 14.1816.170.61411 11.103.332 *10 15.38 14.0816.560.96985.162.272… CM 3 13 17.29 16.0818.440.69211 6.242.498 *10 17.35 14.9818.851.0778 8.162.856 * PC113 0.532 …0.9361.4180.626112.02…1.421…10 0.124 …0.8921.1610.64180.01…0.101… SouthArabiaIranandPakistan n M minmaxSDdf FTPn M minmaxSDdf FTP LAt35 90.37 85.294.82.41633 9.053.008 **19 88.42 81.592.92.97317 4.612.146 * LCr29 40.80 38.4842.941.06127 27.895.281 ***18 39.95 38.0042.071.20016 67.108.191 *** LCb29 39.15 36.6441.471.10427 23.064.803 ***18 38.33 36.1440.491.27116 43.256.576 *** LaZ29 24.57 23.1326.570.82427 24.064.905 ***19 24.49 23.4226.420.82617 22.874.782 *** LaI29 7.90 7.068.650.379270.500.707…19 7.73 7.258.730.363171.771.332… LaP29 7.52 6.638.630.511270.21…0.459…19 7.75 6.978.980.490171.41…1.188… LaN29 16.41 15.5817.040.44527 14.073.751 **18 16.49 15.7417.190.43616 28.515.340 *** AN29 12.46 11.9013.290.37027 7.002.646 *18 12.25 11.6113.230.40616 6.742.595 * CC29 8.14 7.588.820.290273.651.912…19 8.64 7.6817.962.279170.41…0.644… M 2 M 2 28 12.13 11.6512.740.275260.18…0.419…16 12.08 11.3412.830.43714 12.493.534 ** CM 2 29 15.81 14.8216.700.46627 19.144.375 ***19 15.56 14.3016.540.66917 47.736.909 *** LMd29 31.70 29.6333.370.84327 22.524.745 ***19 31.30 29.6832.891.01617 81.279.015 *** ACo29 14.42 12.8815.580.75027 6.872.621 *19 13.49 12.3814.360.602174.252.061… CM 3 29 17.22 16.3818.230.49027 24.244.923 ***19 16.93 15.7518.320.68117 13.363.655 ** PC128 0.720 …0.0961.6120.454260.080.291…19 1.038 …0.0831.9510.790170.060.241… Explanations: F … F valueofANOVA; T … t -valueof t -test; P …signi“cance).SeetheAbbreviationssectionforanexplanationofthe dimensionabbreviationsused. thePalaearctic R . aegyptiacus samplesets(Table1). WhilelargerbatsoftheLevantandEgypthadlesspronounceddimorphism,observedmainlyinskullwidth (LaI,LaP,CC)anduppertooth-rowlength(CM 2 ), highlysigni“cantsexual dimorphismwasdetectedin smallerbatsfromthesouthernMiddleEastandSinai, mainlyinskulllengthdimensions(LCr,LCb,CM 2 , LMd,CM 3 )andanumberofskullwidthmeasurements (LaZ,LaN,CC,M 2 M 2 ).Dimorphisminthemajority ofsamplesets,however,wasnothighlysigni“cant(see thetestresultsforPC1inTable1andcomparefactor loadingsofparticulardimensionsforbothsexesinTable2).Sexualdimorphism,therefore,doesnotappear tobeasigni“cantfactoraectingtheseparationofsamplesintothegeographicallylimitedsizemorphogroups observed. Themutualpositionsofthegeographicalsetswere indicatedthroughPCAofthe13cranialdimensions (Fig.3).Samplesalongthe1stPC(aectedmainly bysizecharacters)wereclu steredintothreegroups,

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1234 P. Benda etal. Fig.2.Bivariateplotof Rousettusaegyptiacus samplesfromthePalaearctic:greatestlengthofskullagainstthelengthofupper tooth-row.Symbolsassociatedwith × representmales,symbolswith+representfemales. Fig.3.Bivariateplotof Rousettusaegyptiacus samplesoffromthePalaearctic:resultsofprincipalcomponentanalysisforallcranial dimensions(“rstandsecondfactors,principlecomponent).Symbolsassociatedwith × representmales,symbolswith+represent females. withlargerbatsfromEgyptandtheLevant(PC1…2.0… 0.5),smallerbatsfromthesouthernMiddleEastand Sinai(PC1…0.9…2.0),andintermediatebatsfromJordan(PC1…0.9…1.2).Alongthe2ndPCaxis(presumably,lessaectedbysizecharacters),however,there appearedtobenogeographicalvariation(seeTable2). Thepercentagesofvariance,aswellasthefactorloadingsforeachsex,wereverysimilar,suggestingsimilar levelsofsex-biasedin”uenceaectingtheseparationof samplesintogeographicalmorphogroups. UPGMAclusteranalysisofcranialdimensions from45localitiesoforiginforPalaearctic R . aegyptiacus samples,andfortheeightcomparativesamples fromsub-SaharanAfricaandPakistan(calculatedfrom

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Variationof Rousettusaegyptiacus inthePalaearctic 1235 Table2.Factorloadingsandpercentageofvarianceresultingfromprincipalcomponentanalysisof13cranialdimensionsforallsamples examined(seealsoFig.3)andofparticularsexgroups. AllsamplesMalesFemales Dimension \ FactorPC1PC2PC1PC2PC1PC2 LCr…0.965…0.037…0.961…0.028…0.9640.038 LCb…0.860…0.053…0.956…0.041…0.723…0.043 LaZ…0.908…0.034…0.867…0.004…0.9150.048 LaI…0.7760.101…0.7340.117…0.7180.271 LaP…0.0570.964…0.2290.887…0.0980.751 LaN…0.5490.270…0.5330.448…0.4420.064 AN…0.7850.012…0.751…0.014…0.7840.105 CC…0.410…0.122…0.8540.003…0.1440.698 M 2 M 2 …0.8420.163…0.8680.128…0.8390.051 CM 2 …0.886…0.016…0.886…0.123…0.825…0.146 LMd…0.959…0.021…0.947…0.047…0.9530.001 ACo…0.826…0.205…0.777…0.270…0.7700.236 CM 3 …0.846…0.074…0.749…0.264…0.832…0.118 %var61.5228.52164.2889.07555.5709.532 meanvaluesofalldimensionsfromallspecimensfrom therespectivesite),revealedsimilarpatternofgeographicalvariation(Fig.4).Aswiththeabovecomparison,thetwobasicsizegroupsandoneintermediategroupwereindicated;however,clusteranalysis betterillustratedtherelationshipsobserved(seeFigs 4A,B).SixlocalitysamplesfromEgypt,theonlysamplefromTurkey,two(ofthree)samplesfromCyprus, three(of“ve)samplesfromLebanon,one(oftwo)samplefromSyria,andone(ofthree)samplefromJordanwereallplacedwithin theclusteroflargerbats (LCr43.5…45.3mm,LaZ26.2…28.7mm).Inaddition, thetypespecimenof Eleutheruraunicolor (= Rousettusaegyptiacusunicolor )fromGabon(LCr46.2mm, LaZ27.3mm)wasalsoclusteredwithinthisgroup. AllthreelocalitysamplesfromSinai,ten(oftwelve) samplesfromYemen[includingthetypespecimenof Rousettusarabicus (YEM7)],“ve(ofsix)samplesfrom Oman,bothsamplesfromIran,andbothsamplesfrom Pakistanwereclusteredamongstthesmallerbats(LCr 39.4…41.8mm,LaZ23.7…26.6mm).Inaddition,acluster(LCr41.4…42.5mm,LaZ23.2…24.7mm)containing thetypespecimenof Cynopterusangolensis (= Lissonycterisangolensis )fromAngola,thetypespecimen of Rousettuslanosus fromUganda,andasampleof R.lanosus fromEthiopiawereallclassedasasister grouptotheclusterofsmallerEgyptianfruitbats. Finally,onesample(ofthree)fromCyprus,one(of two)samplesfromSyria,two(ofthree)samplesfrom Jordan,two(of“ve)samplesfromLebanon,two(of twelve)samplesfromYemen,andone(ofsix)samplefromOmanformedaclusterofintermediatebats (LCr41.9…43.4mm,LaZ24.6…27.6mm).Alsoclusteredwithinthisgroupwerethecomparativesamplesof R . aegyptiacus fromlocalitiesinsub-Saharan Africa,onesampleof R . a . leachii fromEthiopia(LCr 41.4…42.9mm,LaZ24.2…26.9mm)andonesampleof R . a . unicolor fromSenegal(LCr39.7…44.2mm,LaZ 23.2…28.5mm).Aseparatecluster(LCr37.2…38.8mm, LaZ20.1…23.4mm)containingreferencesamplesof R . leschenaulti fromPakistanand Lissonycterisangolensis fromSierraLeone(typespecimenof Rousettussmithii ) andUganda(includingthetypespecimenof Rousettus angolensisruwenzorii )wasrankedasasistergroupto theclustercontainingalllocalitysamplesof R.aegyptiacus . Moleculargeneticanalysis Weobtained83nd1sequencesand68cyt b sequences for R . aegyptiacus fromthespeciesdistributionrange. Thesequenceswereassembledintoa520bpalignmentofnd1andan867bpalignmentofcyt b .Nd1 sequencescorrespondedto17uniquehaplotypesand cyt b sequencesto30uniquehaplotypes(Appendix2). ThesehaplotypesweresubmittedtoGenBankunder accessionnumbersJX274443…JX274462fornd1(N1… N20)andJX274463…JX274497forcyt b (C1…C35),respectively. Inthend1alignment,25positionswerevariable withninesitesparsimonyinformative.MPsearchanalysisyieldedoneshortesttree(160steps)withvery shortbrancheswithintheingrouparrangedinacomblikepattern,indicatingshallowphylogeneticstructure. Asidefrom100%bootstrapsupportforthewhole R . aegyptiacus ingroup,onlytwointernalgroups(which mostlycontainedtheMiddleEasternhaplotypes)were signi“cantlysupported,i.e., > 70%support(Fig.5).In thecyt b alignment,80positionswerevariablewith36 sitesparsimonyinformative.MPsearchanalysisyielded nineshortesttrees(279steps)withveryshortbranches withintheingroup.SeveralhaplogroupswereconsistentlypresentineachrecoveredMPtreeandsigni“cantlysupportedbybootstrap(Fig.6). R . aegyptiacus fromAfricawererepresentedbytwohaplogroupsand asolitaryhaplotype,theremaininghaplogroupcontainingsamplesfromtheMiddleEast.Thetopology ofNJtreesforbothmarkersbasicallyagreedwiththe respectiveMPtreesexceptf orminorrearrangements forterminaltaxa;theformer,however,wereusedfor presentationpurposesduetoclearerstructure(Fig.5).

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1236 P. Benda etal. Fig.4.Clusteranalysisoflocalitiesfor Rousettusaegyptiacus andcomparativetaxasamples;AÂ…resultsofclusteranalysis(unweighted pair-groupaverage,Euclideandistances,calculatedfrommeanvaluesofallcranialdimensionsfromallspecimensfromtherespective site;*averagevaluesformultiplesamplesfromonesite;+typespecimens;seeAppendix1forexplanationsofacronyms);BÂ…metric distributionofPalaearcticsamplesseparatedbyclusteranalysis(coloursofsymbolsaccordingtodivisionsinA);CÂ…geographical distributionofPalaearcticsamplesseparatedbyclusteranalysis(coloursofsymbolsaccordingtodivisionsinA).

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Variationof Rousettusaegyptiacus inthePalaearctic 1237 Fig.5.Neighbour-joiningtreeandmedian-joiningnetworkdepictingphylogeneticrelationshipsamongthe Rousettusaegyptiacus samplesbasedonnd1sequences.Explanations:NorthMiddleEastÂ…Cyprus,Turkey,Syria,Lebanon,Jordan,andSinai;North-east AfricaÂ…continentalEgyptandnorthernSudan;SouthMiddleEastÂ…Yemen,OmanandIran;thedashedlinesarenotproportionalto thenumberofsubstitutions(notedattheline);numbersatthetreenodesindicatethebootstrapsupportfrommaximumparsimony analysis. Fig.6.Neighbour-joiningtreeandmedian-joiningnetworkdepictingphylogeneticrelationshipsamongthe Rousettusaegyptiacus samplesbasedoncyt b sequences.ForexplanationsseeFig.5. Geneticdivergencebetween R . aegyptiacus andthe outgrouptaxarangedfrom15.4Â…16.9%forbothmarkers.Within R . aegyptiacus ,nd1divergencesranged upto2.0%anddidnotprovidemuchdistinctionto theshallowstructureofthephylogenetictree.Onthe otherhand,cyt b divergencesreachedvaluestwicethose fornd1,rangingupto4.2%.Amongthecyt b haplogroups(seeFig.6),thatcontainingfourhaplotypes, fromMalawiandKenya,dieredby2.1Â…3.3%fromthe otherhaplogroupandthesolitaryhaplotypefromsubSaharanAfrica,andby2.8Â…4.2%fromtheMiddleEasternhaplogroup.TheMiddleEasternhaplogroupfurtherdieredby1.8Â…2.7%and1.3Â…1.6%fromtheformertwosub-Saharanunits.VariabilitywithintheMid-

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1238 P. Benda etal. dleEasternhaplogrouprangedfrom0.1…0.6%andup to1.2%and2.0%,withinthetwosub-Saharanhaplogroups. Thereconstructednd1haplotypenetworkdisplayedastar-likepatternforthemostfrequenthaplotypes(N1andN2extractedfrommorethan85%of specimens,plus“veadditionalhaplotypes…N3…N7) coveringallfourmaingeographicdivisions,i.e.,the northern(Turkey,Cyprus,theLevantandSinai)and southernparts(Yemen,OmanandIran)oftheMiddleEast,andEgyptandsub-SaharanAfrica(Senegal, Ghana,Gabon,EthiopiaandMalawi)(Fig.5).Mostof thelessfrequenthaplotypesformedasecondgroupwith amorecomplicatedstructur e,includingclosedloops andmissinghaplotypes. Thecyt b networkdisplayedasimilarstar-likepatternforhaplotypesfromtheMiddleEastandnortheastAfrica(EgyptandnorthernSudan)(Fig.6).Contrarytothend1network,however,cyt b haplotypes fromsub-SaharanAfrica(Senegal,Ghana,Gabon, Ethiopia,KenyaandMalawi)wereplacedseparately, creatingtwodistinctclustersandasolitaryhaplotype, i.e.,theclusteringcorrespondedwiththerecoveredphylogenetictree.Asseenfromboththephylogenetictree andthenetwork,substantialdiversityispresentinsubSaharanAfrica,whileMiddleEasternandnorth-east Africansamplesformacompactgroupwithcloselyrelatedhaplotypes,threebeingpresentathighfrequency throughouttheareasampled(61.8%ofthesamplesexamined).Onlytwocyt b haplotypesdisagreewiththe clearseparationofPalaearcticandsub-Saharansamples,haplotypeC6fromEthiopiarepresentingalineageofitsownclusteredclosetotheMiddleEastern lineage,andtheC7haplotypefromYemenclustered withinthesub-Saharanlineage(composedofsamples fromMalawi,Gabon,Ethiopia,GhanaandSenegal). Discussion Previousstudieshaveconcludedthatvariationinbody sizeisthemostpronouncedaspectofgeographicvariationin Rousettusaegyptiacus inthePalaearcticand that,inthenorthernpartofthespeciesrange,this ischaracterizedbytheappearanceoftwodistinctsize groupsrepresentingseparate subspecies:alarger-sized form( R . a . aegyptiacus )inthenorthernMiddleEast andEgyptandasmallerform( R . a . arabicus )inthe southernMiddleEast(seeEisentraut1959;Hayman& Hill1971;Bergmans1994;Koopman1994;Kwiecinski &Griths1999;Simmons2005).Although,ingeneral, theresultsofourstudycon“rmthisview,re“ningof geographicscalerevealspatternsthattendtocontradictit,withincongruentdistributionoftherespective sizegroupsand/orapredominanceofintermediatesize characteristicsoveraconsid erablepartofthespecies Palaearcticrange,stretchingfromCyprustoOman(see Fig.4C).ThesituationinthesouthernLevantsuggestsaclineshiftinsizebetweenthelargerMediterraneanmorphotype(Turkey,CyprusandLebanon)and thesmallerdesertmorphotype[southernJordan,Sinai and,presumably,southernIsrael(Zelenova&Yosef 2003;Bendaetal.2008,2010)].Incontrast,thesmall Sinaiticmorphotypedirectlyabutsalargemorphotype (thelargestofitsrepresentatives)incontinentalEgypt, whereaverysharpstepinsizeisevident.Afurthersize shiftmightbeexpectedalongtheHijazRangeinwesternArabia,fromJordantoYemen,whereindividuals ofintermediatebodysize(inthepresentsense)have previouslybeenreported(seedatabyNader1975). However,verylimitedmaterialhasbeenexaminedfrom thisextensivearea(butseethediscussionbyBergmans 1994:103).Despitegeneralagreementwithdierencein bodysizebetweenthemarginalpopulationswithinthe speciesPalearcticrange(EgyptandnorthernLevant vs .southernArabia,IranandPakistan),theprevailing patternofgeographicsizevariationtendstosuggest amosaicpatternratherthancontinuousdistribution oftwomoreorlesscategoricallydelimitedandbroadly distributedphenotypes.Asidefromthepopulationsin continentalEgyptandtheeasternmostpartsofthe MiddleEast(whichonly“tthemodelwell),thesituationisobviouslymorecomplicatedinotherregions, appearingtoincludenumerouslocalspeci“citiessuch astheeectsoflocalhabitatconditions(cf.Bendaet al.2008).Inconclusion,ourresultsprovidelittlesupportforthetraditionalsubspeci“carrangementofthe speciesinthePalaearctic,asituationfurthersupported bytheresultsofgeneticanalysis. Ingeneral,wefoundverylowlevelsofgeneticvariationinthemtmarkersstudiedandalackofanyrobust phylogeographicstructurethroughoutthedistribution rangeof R . aegyptiacus (includingthesub-Saharanregions).Identicalhaplotypeswerefoundinverydistantpartsoftherange,e.g.thend1haplotypeN1 (foundin23specimensor27.7%ofsamplesanalysed) appearedinseveralpopulationsoverthePalaearctic range(Lebanon,Jordan,Sinai,EgyptandYemen)as wellasinSenegal.Similarly,haplotypeN2wasfound inextremelyhighnumbers ,appearingin36samples (52.9%)coveringthewholePalaearcticrange(allcountriesoforiginexceptSyria)aswellassub-Saharan GabonandMalawi.Othernd1haplotypes,diering fromeachotherbyasingleorfewsubstitutionsonly (seeFig.5),alsoshowedquiteextensivedistribution, suchasN3(Lebanon,Jordan,YemenandOman)and N4(EgyptandYemen). ConsideringthemutualpositionsofthePalaearctichaplotypes,thepatternofgeneticvariationdemonstratedbyanalysisofcyt b haplotypesdieredonly slightlyfromthatofnd1(seeFig.6).Threehaplotypesweremostwidespread,whileanumberofadjacentvariantsdierbyoneortwosubstitutionsonly. TheC1haplotypewasobservedin15samples(22.1% ofsamplesanalysed)andfoundoveralmostthewhole oftheMiddleEast(Cyprus,Lebanon,Jordan,Egypt, YemenandOman),whileC2wasfoundin12samples (17.6%)fromJordan,SinaiandIran.Themostwidely distributedcyt b haplotypewasC3,foundin15samplescollectedfromnorth-easternAfrica(NSudanand Egypt),easternMediterranean(Turkey,Syria,Lebanon

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Variationof Rousettusaegyptiacus inthePalaearctic 1239 andJordan)andinYemen.Althoughclearseparation ofthePalaearcticandsub-Saharansamplesisobvious withintheresultsofcyt b analysis(unlikethend1results),twohaplotypesdisagreewiththisarrangement: oneEthiopiansamplethatclustersclosetothosefrom theMiddleEastandoneYemenisamplethatgroups withthesub-Saharansamples. Theabove“ndingssuggestthreepossibleexplanationsfortheevolutionaryhistoryof R . aegyptiacus : (1)thegenesexamined(andespeciallynd1)arehighly conservativeandnotinformativeforthehistoryofthe species;(2)thereisanunstructuredgene”owoverthe wholerange,mediatedbylong-distancemigrations(at leastinfemales)interconnect ingduringhistoricaltime eventhemostdistantpart softherange(e.g.,SenegaltoLebanon,MalawitoTurkey,CyprustoOman, etc.);or(3)thecontemporaryPalaearcticrangewas colonisedrelativelyrecentl yfromageographicallylimitedpopulation,possiblyofAfro-tropicalorigin. Thedierenceinphylogeographicsignalforthe twomarkerssuggestsputativevalidityofhypothesis (1),atleastinthecaseofthend1gene.However, manyotherphylogeneticstudiesofbatshaveconvincinglydemonstratedaconsiderabledegreeofwithin-and between-speciesvariationinthend1gene(seeforexampleMayeretal.2007;Bendaetal.2004andBenda &Gvoždík2010),oftenevenhigherthanthatforthe morecommonlyusedcyt b gene.Therelevanceofthis argument,andthelowlevelofvariationfoundinboth markers,meansthatonecannotexcludethepossibilitythatgenotypicvariationinpteropodidbatsdiffersfromthatinmicrobatsingeneral,asisthecase forgenomesize(Smith&Gregory2009).Inconnectionwiththis,itisworthmentioningthatGoodman etal.(2010),whenstudyinggenogeographicvariation in Rousettusmadagascariensis Grandidier,1928,one ofthesistertaxaof R . aegyptiacus ,alsofoundnophylogeographicsignalincyt b sequences,andveryshallowbetween-populationdis tances(notexceeding1.1%) throughoutthe1,600kmdistancebetweenMadagascaranditsneighbouringislands.Similarly,Chenetal. (2010)foundunexpectedlylowvariationinmtDNAin fruitbatsofthegenus Pteropus Brisson,1762colonisingislandsintheIndianOcean.Inbothcases,the extensivepopulationmixingandcontinuousgene”ow hypothesis,whichcouldexplainlowgenotypicvariation,wasrefutedforlackofanydirectsupport,i.e., nolong-distancemigrationswereobservedandthegeographicdistancesbetweenparticularsamplesitesexceededbyalargedegreetheassumeddispersalcapacityofthetaxaconcerned.Campbelletal.(2004: 774),whofoundconsiderableincongruencebetween patternsofmolecularandmorphologicalvariationin the Cynopterusbrachyotis (M¨ uller,1838)speciescomplex,commentedontheunexpectedlylowgeographic variationinmtDNAinthiscladeusingthefollowing words:wecannotdeterminewhethertheabsenceof geneticstructureintheSundalineageissolelyaconsequenceoftheeectexpectedwhensourcepopulationsundergorapiddemographicexpansion,orifvariationhasbeenfurtherreducedby“xationofapositivelyselectedmitochondrialhaplotypeviaaselective sweep[ ... ].Thus,withoutcomparisonofnuclearmarkers,wecannotruleoutthepossibilitythatthislineagemayhaveasigni“cantlylongerandmorecomplexevolutionaryhistorythanthatinferredfrommitochondrialhaplotypedata.ŽThesurprisinglylowgenetic variationfoundinPalaearctic R.aegyptiacus couldbe commentedonusingidenticalwords.Analternativeexplanationforthelackofphylogeographicsignalinthe Mediterraneancouldbebasedontheeectsofrecent expansionpromotedbypost-Neolithicspreadofculturalfruittrees.Inthecaseofthesub-Saharanrange, however,suchanexplanationwouldnecessarilyhave torelyonothermechanismsdrivingextensiverange expansionand/orenormousvagility,i.e.,qualitiesnot knownforthespeciesand,withregardtoitscavedwellingspecialization,quiteimprobable. Incontrasttosomeotherf ruitbatspecies,such as Eidolonhelvum (Kerr,1792)fromtheAfro-tropics, therefore,thereisnodirect supportforhypothesis(2) withregardto Rousettusaegyptiacus asregularmass long-distancemigrationsh aveneverbeenobservedfor thespecies,despiteitsconsiderablecapacityformovement(Tsoaretal.2011).Suchasedentarylifehistory for R . aegyptiacus isalsosupportedbythemorphologicalrecord(seeabove). Consequently,hypothesis(3)appearstobethe mostplausibleexplanation.Thecyt b phylogeographic signalappearstoconformtoasingleinvasionfrom AfricaandsubsequentmultipleradiationsintheMiddle East,followedbyepisodicchangesamongpopulations ofnorth-easternAfricaandsouth-westernArabia(as the“ndingsoftheYemenicyt b haplotypeinEthiopia andAfricanhaplotypeintheMiddleEastsuggest;see Fig.6).Thelattertworegionscanbelookeduponas Afro-Arabiantransitionareasthatcouldserveasgatewaysfortheoriginalinvasionastheycurrentlyrepresentthemostcloselysituatedpartsofthesub-Saharan andPalaearcticpartsofthespeciesrange(Bergmans 1994;Kwiecinski&Griths1999). Asdemonstratedelsewhere(seethereviewby Bendaetal.2007),theoccurrenceof R . aegyptiacus intheMediterraneanarborealzone,aswellasin oasesoftheSaharo-Sindianeremiczone,isstrictlycontrolledbyfruitsupplyinwinter.Inadditiontothe autochthonouscarob( Ceratoniasiliqua L.,1758)and datepalm( Phoenixdactylifera L.,1753),fruitavailabilityincludesalienplantssuchaschinaberry(Melia azedarach L.,1753)(Korineetal.1999).Thewinter survivalof R . aegyptiacus inthisregion,therefore,is closelydependentuponanthropogenicagriculturalproduction;colonisationofthepresentMediterraneanpart oftherangebeinglargelycontingentonanthropiccrop productioninthepost-NeolithicHoloceneperiod(cf. Bergmans1994).Indeed,itappearshighlyprobable thatmostofthespeciespresentMediterraneanrange wascolonisedduringtheHoloceneperiod.QuestionsregardingtheactualdynamicsoftheHoloceneexpansion, includingwhether R . aegyptiacus representstheapo-

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1240 P. Benda etal. choricelementoftheHoloceneinallregionswithinits Palearcticrangeorwhether someareaswerecolonised byresidentlocalpopulationspriortotheHoloceneradiation,cannotbeansweredatthepresenttime.Up tillnow,nofossilrecordfor R . aegyptiacus olderthan theHoloceneisavailableintheMediterraneanLevant (Tchernov1988)andfurtherstudiesusingmoresensitivegeneticmarkersareurgentlyneeded. Thisstudysuggeststhatthecurrenttaxonomicarrangementof Rousettusaegyptiacus inthePalaearctic shouldberevisedandthetraditionalsubspeci“cdivisionrefused.Followingfromthis,allPalaearctic R . aegyptiacus populationsshouldnowbeconsideredas representingoneform,thenominotypicalsubspecies, whichisuniforminitsgenetictraitsbutplasticin itsmetrictraits.Unfortunate ly,theexcitingquestions regardingtheorigin,historyandevolutionarydynamicsofthisuniquebiogeographicphenomenon,i.e.,the PalaearcticoshootoftheAfro-tropicalrangeof R . aegyptiacus ,cannotbeansweredde“nitivelyatthe presenttime.Theseissuesneedtobere-investigated usinganextendedlineoffurtherevidenceandwehope thatthepresentsurveyofgeographicvariationwithin thespecieshasprovidedareliableplatformforsuch studies.Acknowledgements Foraccesstothemuseumspecimensundertheircare,we thankG.Doria(MSNG),J.Gaisler(MUB),R.Hutterer (ZFMK),P.JenkinsandL.Tomsett(BMNH),P.Koubek (IVB),andJ.Obuch(JOC).WefurtherthankCh.Drosten andhisteam(UniversityofBonnMedicalCentre,Germany) foracquiringthe R . aegyptiacus samplesfromGhanaandE. Leroyandhisteam(InternationalCentreforMedicalResearchinFranceville,Gabon)foracquiringthesamplesfrom Gabon.ThisstudywassupportedbygrantsfromtheCzech ScienceFoundati on(#206/09/0888),t heGrantAgencyof theAcademyofSciencesoftheCzechRepublic(#IAA 601110905),andtheMinistryofCultureoftheCzechRepublic(#DKRVO00023272). ReferencesAndersonJ.1902.ZoologyofEgypt:Mammalia.Revisedand completedbyW.E.deWinton,F.Z.S.HughRees,Ltd.,London,374pp. BendaP.,Abi-SaidM.,BartonikaT.,BilginR.,FaizolahiK., LuanR.K.,NicolaoH.,ReiterA.,ShohdiW.M.,UhrinM. &HoráekI.2011. Rousettusaegyptiacus (Pteropodidae)in thePalaearctic:listofrecordsandrevisionofthedistribution range.Vespertilio 15 :3…36. BendaP.,AndreasM.,KockD.,LuanR.K.,MunclingerP., NováP.,ObuchJ.,OchmanK.,ReiterA.,UhrinM.&WeinfurtováD.2006.Bats(Mammalia:Chiroptera)oftheEastern Mediterranean.Part4.BatfaunaofSyria:distribution,systematics,ecology.ActaSoc.Zool.Bohem. 70: 1…329. BendaP.,DietzC.,AndreasM.,HotovýJ.,LuanR.K.,Maltby A.,MeakinK.,TruscottJ.&ValloP.2008.Bats(Mammalia: Chiroptera)oftheEasternMediterraneanandMiddleEast. Part6.BatsofSinai(Egypt)withsometaxonomic,ecological andecholocationdataonthatfauna.ActaSoc.Zool.Bohem. 72: 1…103. BendaP.&GvoždíkV.2010.Taxonomyofthegenus Otonycteris (Chiroptera:Vespertilionidae:Plecotini)asinferredfrom morphologicalandmtDNAdata.ActaChiropterol. 12: 83… 102.DOI:http://dx.doi.org/10.3161/150811010X504617 BendaP.,HanákV.,HoráekI.,HulvaP.,LuanR.&RuediM. 2007.Bats(Mammalia:Chiroptera)oftheEasternMediterranean.Part5.BatfaunaofCyprus:reviewofrecordswith con“rmationofsixspeciesnewfortheislandanddescription ofanewsubspecies.ActaSoc.Zool.Bohem. 71: 71…130. BendaP.,KieferA.,HanákV.&VeithM.2004.SystematicstatusofAfricanpopulationsoflong-earedbats,genus Plecotus (Mammalia:Chiroptera).FoliaZool. 53 (Monograph1) : 1… 47. BendaP.,LuanR.K.,ObuchJ.,ReiterA.,AndreasM.,Bakor P.,BohnenstengelT.,EidE.K.,—evíkM.,ValloP.& AmrZ.S.2010.Bats(Mammalia:Chiroptera)oftheEasternMediterraneanandMiddleEast.Part8.BatsofJordan: fauna,ecology,echolocation,ectoparasites.ActaSoc.Zool. Bohem. 74: 185…353. BergmansW.1994.TaxonomyandbiogeographyofAfricanfruit bats(Mammalia,Megachiroptera).4.Thegenus Rousettus Gray,1821.Beaufortia 44(4): 79…126. CampbellP.,SchneiderC.J.,AdnanA.M.,ZubaidA.&KunzT. H.2004.PhylogenyandphylogeographyofOldWorldfruit batsinthe Cynopterusbrachyotis complex.Mol.Phylogenet. Evol. 33(3): 764…781.DOI:10.1016/j.ympev.2004.06.019 ChenJ.,RossiterJ.S.,FlandersJ.R.,SunY.,HuaP.,MillerButterworthC.,LiuX.,RajanK.E.&ZhangS.2010.Contrastinggeneticstructureintwoco-distributedspeciesofold worldfruitbat.PLoSONE 5(11): 1… 9, art.no:e13903.DOI: 10.1371/journal.pone.0013903 CorbetG.B.1978.TheMammalsofthePalaearcticRegion: ATaxonomicReview.BritishMuseum(NaturalHistory)& CornellUniversityPress,London&Ithaca,314pp.ISBN: 0801411718,9780801411717 EisentrautM.1959.DerRassenkreis Rousettusaegyptiacus E. Geo.Bonn.Zool.Beitr. 10(3–4): 218…235. GoodmanS.M.,ChanL.M.,NowakM.D.&YoderA.D.2010. PhylogenyandbiogeographyofwesternIndianOcean Rousettus (Chiroptera:Pteropodidae).J.Mammal. 91(3): 593…606. DOI:10.1644/09-MAMM-A-283.1 HallT.A.1999.BioEdit:auser-friendlybiologicalsequencealignmenteditorandanalysisprogramforWindows95/98/NT. NucleicAcidsSymp.S. 41: 95…98. HarrisonD.L.1964.TheMammalsofArabia.VolumeI.Introduction,Insectivora,Chiroptera,Primates.ErnestBennLtd., London,192pp. HarrisonD.L.&BatesP.J.J.1991.TheMammalsofArabia.SecondEdition.HarrisonZoologicalMuseum,Sevenoaks,354pp. ISBN:0951731300 HaymanR.W.&HillJ.E.1971.Part2.OrderChiroptera,pp. 1…73.In:MeesterJ.&SetzerH.W.(eds),TheMammals ofAfrica.AnIdenti“cationManual,SmithsonianInstitution Press,Washington,483pp.ISBN:0874741165 HoráekI.,HanákV.&GaislerJ.2000.BatsofthePalearcticregion:ataxonomicandbiogeographicreview,pp.11…157.In: Wo loszynB.W.(ed.),ProceedingsoftheVIIIthEuropean BatResearchSymposium,Vol.I.ApproachestoBiogeographyandEcologyofBats,ChiropterologicalInformationCenter,InstituteofSystematicsandEvolutionofAnimalsPAS, Kraków,273pp.ISBN:8385222839 IrwinD.M.,KocherT.D.&WilsonA.C.1991.Evolutionofthe cytochromebgeneofmammals.J.Mol.Evol. 32(2): 128… 144.DOI:10.1007/BF02515385 JusteJ.&IbanezC.1993.Geographicvariationandtaxonomy of Rousettusaegyptiacus (Mammalia:Megachiroptera)inthe islandoftheGulfofGuinea.Zool.J.Linn.Soc. 107(2): 117…129.DOI:10.1111/j.1096-3642.1993.tb00217.x KimuraM.1980.Asimplemethodforestimatingevolutionary ratesofbasesubstitutionsthroughcomparativestudiesof nucleotidesequences.J.Mol.Evol. 16(2): 111…120.DOI: 10.1007/BF01731581 KoopmanK.F.1994.Chiroptera:Systematics,pp.1…217.In:NiethammerJ.,SchliemannH.&StarckD.(eds),Handbuch derZoologie,BandVIII,Mammalia,Teilband60,Walterde Gruyter,Berlin&NewYork,vii+224pp.ISBN:3111809269, 9783111809267

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Variationof Rousettusaegyptiacus inthePalaearctic 1241KorineC.,IzhakiI.&AradZ.1999.IstheEgyptianfruit-bat Rousettusaegyptiacus apestinIsrael?Ananalysisofthe batsdietandimplicationsforitsconservation.Biol.Cons. 88(3): 301…306.DOI:10.1016/S0006-3207(98)00126-8 KwiecinskiG.G.&GrithsT.A.1999. Rousettusegyptiacus . Mammal.Species 611: 1…9. MayerF.,DietzC.&KieferA.2007.Molecularspeciesidenti“cationboostsbatdiversity.Front.Zool. 4: 4.DOI: 10.1186/1742-9994-4-4 NaderI.A.1975.Onthebats(Chiroptera)oftheKingdom ofSaudiArabia.J.Zool.Lond. 176(3): 331…340.DOI: 10.1111/j.1469-7998.1975.tb03205.x PetitE.,ExcoerL.&MayerF.1999.Noevidenceofbottleneck inthepostglacialrecolonizationofEuropebythenoctule bat( Nyctalusnoctula ).Evolution 53(4): 1247…1258.DOI: 10.2307/2640827 PosadaD.&CrandallK.A.2001.Intraspeci“cgenegenealogies: treesgraftingintonetworks.TrendsEcol.Evol. 16(1): 37… 45.DOI:10.1016/S0169-5347(00)02026-7 SimmonsN.B.2005.OrderChiroptera,pp.312…529.In:WilsonD.E.&ReederD.M.(eds),MammalSpeciesofthe World,ATaxonomicandGeographicReference,ThirdEdition,Volume1,TheJohnHopkinsUniversityPress,Baltimore,xxxv iii+743pp.ISBN: 0801882214 SmithD.L.&GregoryT.R.2009.Thegenomesizesofmegabats (Chiroptera:Pteropodidae)areremarkablyconstrained.Biol. Letters 5(3): 347…351.DOI:10.1098/rsbl.2009.0016 TchernovE.1988.Thebiogeographicalhistoryofthesouthern Levant,pp.159…250.In:Yom-TovY.&TchernovE.(eds), TheZoogeographyofIsrael,TheDistributionandAbundanceataZoogeographicalCrossroad,MonographiaeBiologicae 62. DrWJunkPublishers,Dordrecht,600pp.ISBN: 9061936500,9789061936503 TsoarA.,NathanR.,BartanY.,VyssotskiA.,DellOmoG.& UlanovskiN.2011.Large-scalenavigationalmapinamammal.Proc.Natl.Acad.Sci. 108(37): E718…E724.DOI: 10.1073/pnas.1107365108 ZelenovaN.&YosefR.2003.BatsintheEilatregion(Israel), spring2002.Nyctalus(N.F.) 9(1): 57…60. ReceivedSeptember25,2011 AcceptedJuly23,2012Appendix1.Listofmaterialexaminedduringmorphologicalanalysis.Acronymsfortherespectivesitesusedinthecluster analysisappearinbrackets(Fig.4). Rousettusaegyptiacus (Georoy,1810) Cyprus :1m(NMP90435[S+A]),AndrolikouGorge,2kmSWProdromi[CYP1],20April2005,leg.P.Benda&V. Hanák;…1ind.(NMP90399[S]),Akamas,BathsofAphrodite[CYP2],10April2005,leg.P.Benda&V.Hanák;…1m (NMP91274[S+A]),Akamas,SmigiesTrail,MagnesiaMine[CYP3],27March2005,leg.I.Horáek,P.Hulva&R.Luan. Egypt :1m(NMP92582[S+A]),Aswan[EGY1],24January2010,leg.P.Benda,R.Luan&I.Horáek;…8m,6f,1ind. (IVB1…3,5…10,13…16,18[S+B],MUB1.1.106[S]),Cairo,SultanHamidMosque[EGY2],23April1969,June1971,leg.J. Gaisler&J.Groschaft;…3m,7f(ZFMK63.267…63.272,63.274[S+B],63.275…63.277[S]),Cairo,SultanHassanMosque [EGY3],11May1951,leg.H.Hoogstraal;…1m(NMP92570[S+A]),Bawiti,BahariyaOasis[EGY5],18January2010, leg.P.Benda,R.Luan&I.Horáek;…2m,1f,2inds.(ZFMK62.199,62.200[S],94.499,94.501[S+B],94.502[S+Sk]), [Lower]Egypt(undef.)[EGY4],coll.M¨ ohres,17August1994,ded.FlughafenD¨ usseldorf;…13inds.(NMP91817…91820[S], 92101[S+A]),Qasr,DakhlaOasis[EGY6],17April2002,leg.P.Munzlinger&P.Nová,21January2010,leg.P.Benda, R.Luan&I.Horáek;…1m,1f(NMP90527,90528[S+A]),Sinai,AinElFurtaga[SIN1],16September2005,leg.M. Andreas,P.Benda,J.Hotový&R.Luan;…1m(NMP90520[S+A]),Sinai,AinHudra[SIN2],14September2005,leg.M. Andreas,P.Benda,J.Hotový&R.Luan;…5m,6f(NMP90501,90510[Sk],90502…90509,90511[S+A]),Sinai,Feiran [SIN3],10September2005,leg.M.Andreas,P.Benda,J.Hotový&R.Luan. Ethiopia :3m,5f(NMP92170…92176[S+A],JOCunnumbered[S+Sk]),GiloRiverbridge,5kmSofTepi[ETH1],8May 2003,leg.P.Benda&J.Obuch. Gabon :1f(BMNH62.8.26.1.[S+B],typeof Eleutheruraunicolor Gray,1870),AfriqueOccidentale(Gabon)[GAB1], purch.Verreaux. Iran :5m,5f(NMP48377…48386[S+A]),Espakeh[IRN1],10April2000,leg.P.Benda&A.Reiter;…3m,6f(NMP 40467[S+B]),Isin[IRN2],29April&2May1977,leg.B.Pražan. Jordan :1m(NMP92558[S+A]),Kufranja,IraqalWahajCave[JOR1],26May2009,leg.P.Benda,A.Reiter&J. Obuch;…1m,1f(NMP92362,92411[S+A]),WadiasSir,IraqalAmirCave[JOR2],10October2008,10May2009,leg. P.Benda,A.Reiter&J.Obuch;…1m,7f(NMP92430,92431,92433,92434,92436,92438,92440,92442[S+A]),Wadi Dhana,4kmENEofFeinanEcolodge[JOR3],14May2009,leg.P.Benda,A.Reiter&J.Obuch. Lebanon :1m(AUBM021[S]),Antelias[LEB1],19March1960,leg.J.E.Stencel;…1m,1f(NMP91799,91910[S+A]), Antelias[LEB1],KassaratCave,25January2007,25Januar y2008,leg.P.Benda,R.erný,I.Horáek,R.Luan&M. Uhrin;…1f(AUBM006[S]),cave4kmSEofBeitMeri[LEB2],4October1959,leg.R.E.Lewis;…1m,1f(NMP 91904,91905[S+A]),DahrelMghara,MgharetelAaonamiecave[LEB3],19January2008,leg.P.Benda,I.Horáek,R. Luan&M.Uhrin;…2f(NMP93697,93699[S+A]),JeitaCave[LEB4],20March2009,leg.T.Bartonika,P.Benda,I. Horáek&R.Luan;…2m,1f(NMP91765,91766,91899[S+A]),Tarabulus,MtalalAzraqCave[LEB5],21January 2007,18January2008,leg.P.Benda,R.erný,I.Horáek,R.Luan&M.Uhrin. Oman :1f(NMP92733[S+A]),AinJarziz[OMA1],27October2009,leg.P.Benda,A.Reiter&M.Uhrin;…1m,1f (NMP92679,92680[S+A]),AlNakhar[OMA2],22October2009,leg.P.Benda,A.Reiter&M.Uhrin;…1f(NMP92654 [S+A]),DhahirAlFawaris[OMA3],21October2009,leg.P.Benda,A.Reiter&M.Uhrin;…1m(NMP92778[S+A]),wadi 7kmWofDibab[OMA4],2November2009,leg.P.Benda,A.Reiter&M.Uhrin;…1m(NMP92651[S+A]),Khutwa [OMA5],20October2009,leg.P.Benda,A.Reiter&M.Uhrin;…1m(NMP92756[S+A]),Taiq[OMA6],30October 2009,leg.P.Benda,A.Reiter&M.Uhrin.

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1242 P. Benda etal. Pakistan :2m(BMNH20.1.17.1.,20.1.17.2.[S]),Karachi,Sind[PAK1],leg.C.B.Tycehurst;…2m,1f(BMNH19.11.7.1., 19.11.7.2.,19.11.7.4.[S]),Panjgur,Baluchistan[PAK2],leg.J.E.Hotson. Senegal [SEN1]:1m,2f(IVBS172…174[S+B]),Dakar,22October2004,leg.J.ervený&P.Koubek;…1m(IVBS1267 [S+A]),Niokolo,19August2006,leg.J.ervený&P.Koubek. Syria :1m,1f(NMP48865,48866[S+A]),Talshhab[SYR1],25May2001,leg.M.Andreas,P.Benda,A.Reiter&D. Weinfurtová;…1m,1f(NMP48264,48265[S+A]),YaarOdenforest(GolanHeights)[SYR2],18July1999,leg.P.Benda. Turkey :1m(ZFMK65.205[S+B]),DermustluK¨ oy,H¨ ohlebeiAntakya[TUR1],2January1952,leg.H.Kumerloeve. Yemen :3m,2f(NMPpb3112…pb3116[S+A]),5kmWofHammamAli[YEM1],27October2005,leg.P.Benda;…2f (NMPpb3056,pb3057[S+A]),AlKhuraybah,WadiDawan[YEM2],19October2005,leg.P.Benda;…4m,2f(NMP pb3628…3630,3632,3633[S+A],pb3631[A]),AssalaatMashgab,SofTaiz[YEM3],26October2007,leg.P.Benda&A. Reiter;…1m(NMPpb3758[S+A]),Halhal,10kmNEHajja[YEM4],2November2007,leg.P.Benda&A.Reiter;…2m, 1f(NMPpb2959…pb2961[S+B]),Hawf[YEM5],12October2005,leg.P.Benda;…2f(NMPpb3118,pb3119[S+B]),Jebel Bura,WofRiqab[YEM6],30October2005,leg.P.Benda;…1m(BMNH95.6.1.47.[S+B],holotypeof Rousettusarabicus Anderson,1902),Lahej,Aden;shotnearSultansgarden[YEM7],21March1895,leg.G.W.Yerbury;…2m(NMPpb2943, pb2944[S+A]),Maarib[YEM8],9October2005,leg.P.Benda;…1m(NMPpb2956[S+A]),Sah,WadiHadramawt [YEM9],11October2005,leg.P.Benda;…1f(NMPpb3159[A]),WadiAlLahm,WofAlMahwit,1October2005,leg. P.Benda;…1m(NMPpb2917[S+A]),WadiDhahr,15kmNofSanaa[YEM10],6October2005,leg.P.Benda;…2f (NMPpb3089,pb3090[S+A]),WadiMaytam,12kmSEofIbb[YEM11],26October2005,leg.P.Benda;…1m(NMP pb3728[S+A]),WadiZabid,SEofAlMawkir[YEM12],30October2007,leg.P.Benda&A.Reiter. Rousettusleschenaultii (Desmarest,1820) Pakistan :1ind.(BMNH69.484[S]),Lahore,OldTomb,WPakistan[PAK3],2February1968,leg.T.J.Roberts;…2m, 1f(BMNH67.1105,67.1106[S]),Lahore,TombofAliMordanKhan[PAK3],WPakistan,16November1966,leg.T.J. Roberts. Rousettuslanosus Thomas,1906 Ethiopia :5m,2f(NMP92181…92187[S+A]),BaroRiverbridge,15kmNofMasha[ETH2],9May2003,leg.P.Benda &J.Obuch. Uganda :1m(BMNH6.4.1.2.[S+A],holotypeof Rousettuslanosus Thomas,1906),Ruwenzori[UGA2]. Lissonycterisangolensis (Bocage,1898) Angola :1f(BMNH97.8.6.1.[S+A],typeof Cynopterusangolensis Bocage,1898),Quihula,Benguella[ANG1]. SierraLeone :1f(BMNH8.9.11.1.[S+B],holotypeof Rousettussmithii Thomas,1908),SierraLeone[SLE1],leg.Canon F.C.Smith. Uganda :4f(MSNG15303…d[S+B]),Arcip.Sesse:Maiba[UGA1],1908,leg.E.Bayon;…2m(MSNG13616a,b[S+B]), Bukasa(Iss.Sesse)[UGA1],15June1908,leg.E.Bayon;…2m,1f(BMNH6.12.4.1.[S+B],holotypeof Rousettusangolensis ruwenzorii Eisentraut,1965,MSNG6414,6415[S+B]),RuwenzoriEast[UGA1],5500ft.,8&12March1906,leg.R.R. Dent. Appendix2.Listofmaterialexaminedduringmoleculargeneticanalysis;nd1,cyt b =haplotypesoftherespectivegenes. nd1cyt b VoucherCountrySiteCoordinates haplotypeGenBankhaplotypeGenBank Acc.No.Acc.No. Rousettusaegyptiacus N2JX274457……NMP93850TurkeySayk¨ oy,WofTarsus36 57 N,34 47 E N2…C3JX274479NMP93851TurkeySayk¨ oy,WofTarsus36 57 N,34 47 E N2…C3…NMP93852TurkeyHarbiye36 09 N,36 08 E N2…C1JX274474NMP90435CyprusProdromi,AndrolikouGorge35 00 N,32 23 E N2………NMP91274CyprusNeoHorio,MagnesiaMine35 03 N,32 20 E N17JX274446……NMP48865SyriaTalshhab32 42 N,35 58 E N11JX274452……NMP48867SyriaTalshhab32 42 N,35 58 E N13JX274450C3…NMP48264SyriaYaarOden,Golan33 12 N,35 46 E N16JX274447C13JX274482NMP91909LebanonAamchite,SalehCave34 09 N,35 40 E N2………NMP91799LebanonAntelias,Kassaratcave33 55 N,35 36 E N3JX274462C1…NMP91910LebanonAntelias,Kassaratcave33 55 N,35 36 E N2…C1…NMP93697LebanonJeitaCave33 57 N,35 39 E N2…C3…NMP93699LebanonJeitaCave33 57 N,35 39 E N2…C3…NMP91765LebanonTarabulus,MtalalAzraqCave34 25 N,35 50 E N1JX274456C1…NMP91766LebanonTarabulus,MtalalAzraqCave34 25 N,35 50 E ……C3…NMP91899LebanonTarabulus,MtalalAzraqCave34 25 N,35 50 E N1…C1…NMP93712LebanonWadiJilo33 14 N,35 19 E N1……… biopsy IsraelBeitOren32 43 N,35 01 E N2…C2JX274468 biopsy JordanJufatal-Qafrayn31 53 N,35 37 E N7JX274458C2… biopsy JordanJufatal-Qafrayn31 53 N,35 37 E N3…C14JX274483NMP92558JordanKufranja,IraqalWahajCave32 19 N,35 43 E

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Variationof Rousettusaegyptiacus inthePalaearctic 1243 Appendix2.(continued) nd1cyt b VoucherCountrySiteCoordinates haplotypeGenBankhaplotypeGenBank Acc.No.Acc.No. N2…C1… biopsy JordanNahla32 17 N,35 49 E N1………NMP47975JordanTabaqatFahl32 27 N,35 37 E N2…C2… biopsy JordanWadialWalah31 33 N,35 44 E N1…C3…NMP92362JordanWadiasSir,IraqalAmir32 13 N,35 53 E N2…C3…NMP92411JordanWadiasSir,IraqalAmir32 13 N,35 53 E N2…C2…NMP92440JordanWadiDhana,nearFeinan30 39 N,35 32 E N1………NMP92430JordanWadiDhana,nearFeinan30 39 N,35 32 E N1………NMP90527EgyptSinai,EinElFurtaga29 03 N,34 33 E N2…C2…NMP90528EgyptSinai,EinElFurtaga29 03 N,34 33 E N1…C2…NMP90520EgyptSinai,EinKhudra28 54 N,34 25 E N1………NMP90503EgyptSinai,WadiElFeiran28 42 N,33 40 E ……C30JX274496NMP90504EgyptSinai,WadiElFeiran28 42 N,33 40 E N1…C2…NMP90508EgyptSinai,WadiElFeiran28 42 N,33 40 E N1…C2…NMP90509EgyptSinai,WadiElFeiran28 42 N,33 40 E N1…C3…NMP92582EgyptAswan24 07 N,32 54 E N2…C1…NMP92570EgyptBahariya,Bawiti28 21 N,28 52 E N16…C3… biopsy EgyptCairo,Zamalek,FishGarden30 03 N,31 13 E N4JX274461C3… biopsy EgyptCairo,Zamalek,FishGarden30 03 N,31 13 E N2…C1… biopsy EgyptCairo,Zamalek,FishGarden30 03 N,31 13 E N2…C3…NMP92101EgyptDakhla,Qasr25 42 N,28 53 E N1…C3…NMP92102EgyptDakhla,Qasr25 42 N,28 53 E ……C27JX274486NMP93677SudanFerqa20 54 N,30 35 E ……C3…NMP93678SudanFerqa20 54 N,30 35 E N3………NMPpb3056YemenAlKhuraybah,WadiDawan15 09 N,48 26 E N2………NMPpb3057YemenAlKhuraybah,WadiDawan15 09 N,48 26 E N1…C1…NMPpb3629YemenAssala,Mashgab13 21 N,43 57 E N1………NMPpb3630YemenAssala,Mashgab13 21 N,43 57 E N2…C1…NMPpb3631YemenAssala,Mashgab13 21 N,43 57 E N1…C12JX274481NMPpb3758YemenHalhal,NEofHajja15 44 N,43 37 E N2………NMPpb3113YemenHammamAli14 41 N,44 07 E N5JX274460……NMPpb3115YemenHammamAli14 41 N,44 07 E N3…C8JX274476NMPpb2959YemenHawf16 40 N,53 05 E N1…C1…NMPpb2960YemenHawf16 40 N,53 05 E N2…C9JX274477NMPpb2961YemenHawf16 40 N,53 05 E N1…C10JX274478NMPpb3118YemenJebelBura,WofRiqab14 52 N,43 25 E N2…C1…NMPpb3119YemenJebelBura,WofRiqab14 52 N,43 25 E N1………NMPpb2943YemenMaarib15 24 N,45 16 E N2…C1…NMPpb2944YemenMaarib15 24 N,45 16 E N10JX274453……NMPpb2956YemenSah,WadiHaramawt15 41 N,48 52 E N4…C3…NMPpb3159YemenWadiAlLahm,WofAlMahwit15 26 N,43 29 E N6JX274459……NMPpb2917YemenWadiDhahr,NofSanaa15 27 N,44 08 E N1…C7JX274475NMPpb2918YemenWadiDhahr,NofSanaa15 27 N,44 08 E N2………NMPpb3089YemenWadiMaytam,SEofIbb13 52 N,44 18 E N1…C11JX274480NMPpb3728YemenWadiZabid,SEofAlMawkir14 10 N,43 30 E N3………NMP92733OmanAinJarziz17 06 N,54 05 E N2………NMP92735OmanAinTabruq17 06 N,54 20 E N2…C4JX274497NMP92679OmanAlNakhar23 12 N,57 13 E N2………NMP92680OmanAlNakhar23 12 N,57 13 E N2…C16JX274485NMP92654OmanDhahirAlFawaris23 39 N,56 39 E N3…C1…NMP92714OmanHagarir16 42 N,53 09 E N16…C15JX274484NMP92651OmanKhutwa24 19 N,56 08 E N2…C1…NMP92756OmanTaiq17 09 N,54 37 E N2…C4…NMP92778OmanWadiDibab23 04 N,58 59 E N2…C2…NMP48379IranEspakeh26 48 N,60 10 E N2…C2…NMP48381IranEspakeh26 48 N,60 10 E N14JX274449……NMP48382IranEspakeh26 48 N,60 10 E N2…C2…NMP48380IranEspakeh26 48 N,60 10 E ……C2…NMP48383IranEspakeh26 48 N,60 10 E N2………IVBS1316SenegalMako12 51 N,12 21 W N1…C26JX274495IVBS1267SenegalNiokolo13 04 N,12 43 W N1………IVBS1000SenegalWassadou12 53 N,11 51 W N12JX274451C22JX274487IVBpv113GabonBelingaMts00 59 N,13 12 E ……C23JX274488IVBpv114GabonBelingaMts00 59 N,13 12 E ……C24JX274489IVBpv115GabonBelingaMts00 59 N,13 12 E N2…C25JX274490IVBpv116GabonBelingaMts00 59 N,13 12 E ……C20JX274491IVBpv051GhanaBuoyem07 43 N,01 59 W N15JX274448C21JX274492IVBpv052GhanaBuoyem07 43 N,01 59 W

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1244 P. Benda etal. Appendix2.(continued) nd1cyt b VoucherCountrySiteCoordinates haplotypeGenBankhaplotypeGenBank Acc.No.Acc.No. N9JX274454C5JX274469NMP92170EthiopiaGiloRiver,SofTepi07 07 N,35 26 E N13Â…C6JX274470NMP92175EthiopiaGiloRiver,SofTepi07 07 N,35 26 E N8JX274455Â…Â…NMP92176EthiopiaGiloRiver,SofTepi07 07 N,35 26 E Â…Â…C28JX274493NMPrs458KenyaMt.Elgon01 02 N,34 47 E Â…Â…C29JX274494NMPrs459KenyaMt.Elgon01 02 N,34 47 E N2Â…C17JX274465NMPmw128MalawiMulanje-Chitakali16 02 S,35 31 E Â…Â…C18JX274466NMPmw131MalawiMulanje-Chitakali16 02 S,35 31 E N13Â…C19JX274467NMPmw194MalawiMpalanganga15 27 S,35 15 E Rousettuslanosus N18JX274444C31JX274471NMP92182EthiopiaBaroRiver,NofMasha07 52 N,35 29 E N19JX274445C32JX274472NMP92183EthiopiaBaroRiver,NofMasha07 52 N,35 29 E Â…Â…C33JX274473NMP92186EthiopiaBaroRiver,NofMasha07 52 N,35 29 E Lissonycterisangolensis N20JX274443C34JX274463NMP90938MalawiZombaPlateau15 21 S,35 17 E N20Â…C35JX274464NMP90939MalawiZombaPlateau15 21 S,35 17 E


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