The following manuscript was published as short communication in Mammalian Biology 78 (pp. 383-386). The manuscript was peer-reviewed and I analysed the data. I wrote the manuscript that was revised by myself and all co-authors. More details on my contribution to this collaborative work are given below in the form Anlage 1 of the Ausführungsbestimmungen für Dissertationen im Promotionsfach Biologie, die bereits veröffentlichte Teile oder eingereichte Manuskripte enthalten.
Anlage 1 Titel der Publikation/ des Manuskripts: Name des/der jeweiligen Autors/Autoren/Autorin* (1) Entwicklung und Planung (2) Durchführung der einzelnen Untersuchungen/ Experimente (3) Erstellung der Daten-sammlung und Abbildungen (4) Analyse/Interpretation der Daten (5) übergeordnete Einleitung/ Ergebnisse/Diskussion Bei 2, 3 und 4 bitte kurze inhaltliche Angaben der jeweiligen Anteile, bei 1 und 5 reichen prozentuale Angaben*Mehrfacheintragungen möglich Datum/OrtDatum zustimmende Bestätigung der vorgenannten Angaben Unterschrift Promovend/PromovendinUnterschrift Betreuer/Betreuerin
Erklärung über Anteile der Autoren/Autorinnen an den einzelnen Kapiteln der Promotionsarbeit
Als Autoren/Autorinnen werden solche Personen bezeichnet, die an der Arbeit in Bezug auf die genannten Punkte in einer Weise mitgewirkt haben, dass sie für die ausgewiesenen Passagen (mit) verantwortlich sind. Personen, die an der Arbeit mitgewirkt haben, jedoch nicht in diese Kategorie fallen, sollten in der Danksagung Erwähnung finden.Was hat der/die Promovierende bzw. was haben die Co-Autoren/Autorinnen beigetragen#
Phylogenetic and population genetic analyses suggest a potential species boundary between Mountain Gazella gazella and Arabian GazellesG. arabica in the Levant (annes Lerp:% Co-Autoren:% (annes Lerp und Co-Autoren: Amplifizieren der Mikrosatelliten Co-Autoren: Sequenzierung des Cytochrom-b-Gens und der mitochondrialen Kontrollregion (annes Lerp: Veröffentlichung der Sequenzen bei Genbank, Erstellen der Abbildung (annes Lerp: phylogenetische Analyse mit BEAST, Analyse der Mikrosatelliten mit STRUCTURE und GENET)X (annes Lerp und Co-Autoren: )nterpretation der Daten (annes Lerp:% Co-Autoren:%
Martin Plath Markus Pfenninger, Martin Plath Torsten Wronski, Martin Plath, Markus Pfenninger
Anne Schröter en
- 9 5
-MammalianBiology78 (2013) 383–386
ContentslistsavailableatSciVerseScienceDirect
Mammalian Biology
jo u r n al h om ep a g e :w w w . e l s e v i e r . c o m / l o c a t e/ m a m b i o
Shortcommunication
Phylogenetic and population genetic analyses suggest a potential species
boundary between Mountain (Gazella gazella) and Arabian Gazelles (G. arabica) in the Levant
HannesLerpa,∗,TorstenWronskib,c,MartinPlatha,AnneSchrötera,MarkusPfenningerd
aEvolutionaryEcologyGroup,Goethe-UniversityFrankfurt,Max-von-Laue-Str.13,60438FrankfurtamMain,Germany
bZoologicalSocietyofLondon,ConservationPrograms,Regent’sPark,LondonNW14RY,UnitedKingdom
cKingKhalidWildlifeResearchCentre,SaudiWildlifeAuthority,P.O.Box61681,Riyadh11575,SaudiArabia
dMolecularEcologyGroup,BiodiversityandClimateResearchCentre(BiK-F)bySenckenbergGesellschaftfürNaturforschungandGoethe-University, 60325FrankfurtamMain,Germany
a r t i c l e i n f o
Articlehistory:
Received24October2012 Accepted26November2012 Available online 28 December 2012
Keywords:
Antilopinae Bovidae Crypticspecies Gazella Israel
a b s t r a c t
Twocrypticlineagesof‘MountainGazelles’havebeenreportedbasedonmolecularphylogeneticanalyses usingmaternallyinherited(mitochondrial)sequencemarkers,namelyGazellagazellaintheLevantand G.arabicasouthoftheAravaValleyintotheArabianPeninsula.Here,weprovidearigoroustestforthe existenceoftwodistinctlineagesbasedonbi-parentallyinherited(nuclearmicrosatellite)markers.Our studyconfirmstwogeneticallydistinctclustersintheLevantanddetectednogene-flowbetweenthem.
Divergencetime(inferredfromacytochromeb-basedphylogeny)wasapproximatelyoneMYA.Treating andbreedingbothlineagesseparatelyinfutureconservationandcaptivebreedingprogrammesishighly recommended.
© 2012 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
‘MountainGazelles’inhabitbrokenorundulatingterraininthe LevantandtheArabianPeninsula.Theywereformerlydescribed asasinglespecies[Gazellagazella(Pallas,1766)]withanumber of presumedsubspeciesofdoubtful taxonomicvalidity (Groves 1996).A recent phylogenetic analysisby Wronski etal. (2010) suggested that gazelles originating fromthe Golan Heights are reciprocallymonophyletictorepresentativesfromsouthernIsrael andtheArabianPeninsula,alludingtotheexistenceoftwo cryp-tic species. Bärmann et al. (in press) found the lectotype skin of G. arabica(Lichtenstein, 1827) tobe nested within the Ara-bianclade of MountainGazelles (here calledArabianGazelles), hencethename G.arabica isavailable forthis taxon(provided that thelectotype skull – belongingtoG. gazella –is excluded fromthetype).However,theresultsobtainedbyWronskietal.
(2010)werebasedonmitochondrialDNAonly,andintheorythe spatialdistributionpatternofmaternalhaplolinescouldmerely reflectfemalephilopatryalongwithmale-biaseddispersal.To over-comelimitationsofmtDNA-basedphylogenyweusedbi-parentally inherited,nuclearmicrosatellitelociformulti-locusgenotypingof
∗Correspondingauthor.
E-mailaddress:hannes.lerp@gmx.de(H.Lerp).
bothtaxaandforunravellingpossiblegene-flowbetweenthem, whilefocussingonIsraelasaregionwherebothtaxasupposedly co-occur(Mendelssohnetal.1997).
We analysed47 specimens ofboth putativetaxaoriginating fromtheLevantandthewesternArabianPeninsula(Fig.1A,Suppl.
material).Twenty-onespecimensfromWronskietal.(2010)were re-analysed (i.e. re-sequenced); all others were obtained from theSteinhardtNationalCollection, Tel-Aviv(18specimens)and othercollectors(eightspecimens).Forphylogeneticanalysis,we sequencedthecytochromebgeneof39specimensfollowingLerp et al. (2011) while including G. dorcas and the genera Nanger andEudorcasinthefinalalignment(Genbankaccessionnumbers aregiveninFig.1B).BayesiananalysiswasperformedinBEAST 1.6.1(DrummondandRambaut2007);nooutgroupwasdefined beforehand. jModelTest 0.1.1 (Posada 2008) identified HKY+Ŵ asthebestfittingsubstitutionmodel. Weusedmolecularclock dataestimates inferred forG. dorcas(Lerpet al. 2011)and ran MC3simulationswith107generations,discardingthefirst10%of therunsasburn-in.Tocomparesequence divergencewithinG.
gazella,withinG.arabica,andbetweenbothtaxa,respectively,we calculatedpairwiseKimura2-Parameterp-distances(K2P)using MEGA5(Tamuraetal.2011)andconductedaMantel-Testwith K2Pvalues(multipliedby100,seeTobeetal.2010)asthe depen-dentvariableandabinaryindependentvariablethatdifferentiated 1616-5047/$–seefrontmatter© 2012 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
http://dx.doi.org/10.1016/j.mambio.2012.11.005
384 H.Lerpetal./MammalianBiology78 (2013) 383–386
Fig.1. Phylogenybasedoncytochromebsequences.(A)Sampleorigin.(B)Bayesiananalysiswasperformedwith47sequenceswiththeHKY+Ŵsubstitutionmodel.Only PP-values≥0.95arereported.Nodebarsrepresentthe95%credibilityintervalsofdivergencetimesofstatisticallysupportedphylogeneticsplits.OnepresumedG.arabica sample(*)fromtheAravaValleyinIsraelclusteredwithinG.dorcasandseemstohavebeenmisidentified.
between comparisons within species (‘0’) and comparisons betweenspecies(‘1’). Thisanalysiswasconducted usingFSTAT 2.9.3.2(http://www2.unil.ch/popgen/softwares/fstat.htm).
Supplementary material related to this article found, in theonlineversion,athttp://dx.doi.org/10.1016/j.mambio.2012.11.
005.
Forthepopulationgeneticanalysesweamplified microsatel-litelociBM302,BM415,BM4505(Bishopetal.1994);CSSM043 (Mooreetal.1994);INRA40(Vaimanetal.1994);MCM38(Hulme etal.1994);OarFCB304 (Buchananand Crawford1993),RM088 (Kossarek etal. 1995); SRCRSP-6(Bhebheet al.1994);TEXAN6 (Burns et al. 1995a) and TEXAN19 (Burns et al. 1995b) with dye-labelled forward primers (Cy5 and IRD700 obtained from Metabion; Dy-751 from Biomers) for visualization on a Beck-manCoultercapillarysequencerCEQ2000.Differentdye-labelled primerpairswerearrangedinthreemultiplexreactionsusingthe Type-itMicrosatellitePCRKit®(Qiagen,Hilden,Germany).
WeusedGENETIX4.05.2(Belkhiretal.2004)tovisualizegenetic differencesbymeansofamultidimensionalfactorial correspon-denceanalysis(FCA).STRUCTURE2.3.3(Pritchardetal.2000)was employedtoidentifythenumberofgeneticallydistinctclusters(K) accordingtoEvannoetal.(2005)usingtheweb-basedtool STRUC-TUREHARVESTER0.6.8(EarlandvonHoldt2011).Foreachvalue ofK,fiveiterationswererunwithaburn-inperiodof105 gener-ations,followedby106generationsforvaluesofK=1through10.
Eachsimulationwasperformedusinganancestrymodel incorpo-ratingadmixture,amodelofcorrelatedallelefrequencies,butno priorinformationonsampleorigins.
Ourphylogeneticanalysisuncoveredhighposterior probabil-ities(i.e.PP=1)for thesplit betweenMountain(G.gazella)and
ArabianGazelles(G.arabica),confirmingthefindingsofWronski etal.(2010)withnewsequences(Fig.1B).Thedivergencetimeof bothtaxawasestimated1.73to0.86MYA,i.e.intheCalabrian (mid-dlePleistocene).SamplesoriginatingfromcentralIsraelclustered togetherwiththosefromtheGolanHeights(i.e.G.gazella).One pre-sumedG.arabicasamplefromtheAravaValleyclusteredwithin G.dorcas.Bothspeciesoccursympatricallyinthisregion,sothe sampleseemstohavebeenmisclassifiedandwasexcludedfrom subsequentanalyses.Nofurthergeneticstructurewasdetectablein bothtaxaalongtheirdistributionranges(Fig.1B),whichisofspecial interestinthecaseofG.arabicaasGrovesandGrubb(2011)claimed thatthreedifferentspecieswouldexistwithinthesampledrange.
Mean(±s.d.)K2P-values(×100)wereratherlowforcomparisons withintaxa(G.gazella:0.167±0.173;G.arabica:0.856±0.694)but higherbetweentaxa2.270(±0.205);theMantel-Testfoundthis differencetobestatisticallysignificant(rP=0.989,P<0.001).
Populationgeneticanalysessupportgeneticdistinctivenessof bothtaxa.FCAclearly clusteredsamplesintotwogroups corre-spondingwithG.gazellaandG.arabicaeventhoughnopriorsample informationwasprovided (Fig.2A).The uppermosthierarchical levelofpopulationdifferentiationinferredfromSTRUCTUREwas K=2(Fig.2B).Allindividualsincludedwereassignedtoagenetic clusterwithanestimatedgroupmembershipofQ>0.9(Fig.2C),i.e.
norecurrentgene-flowcouldbedetected.
Ourphylogeneticanalysisofcytochromebsequencesandour populationgeneticanalysessupportthehypothesisoftwo recipro-callymonophyleticlineageswithinpresumed‘MountainGazelles’
(Wronskiet al.2010).Thelackofrecurrentgene-flow between both suggests complete (atleast recent) reproductive isolation in thewild. In the light of theIntegrative Species Concept (de
H.Lerpetal./MammalianBiology78 (2013) 383–386 385
-1.5 -1.0 -0.5 0.0 0.5 1.0 1.5
-1.0 -0.5 0.0 0.5 1.0 1.5
Axis 1
Axis 2
A
Δ K
K
0 1 2 3 4 5 6 7 8 9 10 11
0 50 100 150 200 250 300 350 400
-2,000 -1,900 -1,800 -1,700 -1,600 -1,500
0.0 0.2 0.4 0.6 0.8 1.0
Percentage population assignment
Gazella gazella Gazella arabica
B
C
ln Pr(X|K)
Fig.2. Numberofgeneticallydistinctgroupsinthedataset.(A)FactorcorrespondenceanalysisofallelefrequenciesobtainedfromGENETIX(blacksquaresandgreydiamonds representingG.arabicaandG.gazella,respectively).(B)EstimatedlnPr(X|K)(greydiamonds)andK(blackcircles)asafunctionofKinferredfromSTRUCTUREresults.(C) PercentagepopulationassignmenttoinferredgeneticclustersforK=2.
Queiroz2007)theseresultscanbeseenasevidenceforseparately evolving metapopulation lineages (sensu de Queiroz 2007), althoughfuturestudiesshouldincludenuclearsequencemarkers tofurtherevaluateifthesemetapopulationsrefertogoodspecies.
However,pairwisegeneticdistances(i.e.K2P-values)withinboth taxa clearly indicate intraspecific variation, whereas distances betweenthetwotaxaareofamagnitudethatcanbeinterpretedas interspecific(Tobeetal.2010).InthecaseofG.arabicaour samp-lingschemeonlycomprisesspecimensfromthewesternArabian Peninsulaandwecan,therefore,onlyspeculateaboutthe taxo-nomicpositionofpopulationsoccurringinOmanandtheUnited ArabEmirates.
Ourresultsareinlinewithmorphologicaldifferencesand pale-ontologicalfindings.Besidedifferentbodysizeandfurcolouration, both taxa show distinct skull and horn features (see Wronski etal.2010).Furthermore,mammalianfaunasfromlatePliocene and Pleistocene beds in the Levant comprise remnantsof two distinctGazellaspecies (Sickenberg,1975;Tchernov, 1988).The commonancestorsupposedlyimmigratedintotheLevantacross theSaharo-SaheliandesertbeltduringthelatePliocene(Tchernov 1986).WithinthePalaearctictheLevantwasacentreofadaptive radiationsasit wasa crossroadof bioticinterchanges between AfricaandEurasiaandcomprised extremebiogeographical het-erogeneityduetotheadmixtureofPalaearctic,Palaeotropicand Saharo-Arabianelements(Tchernov1988).Here,immigratingtaxa (suchasGazella)underwentextensivespeciationespeciallyduring thelateNeogene(Tchernov1988).Someofthespeciesthatevolved intheLevantinvadedareassuchassouthernTurkeyandthe Ara-bianPeninsula(Tchernov1988).In thelate PleistoceneTurkish gazellesdisappearedandthemorehumid-adaptedG.gazellasettled inthenorthernLevantandsouthernTurkey(Kasparek1986),while themorearid-adaptedG.arabicacolonizedtheArabianPeninsula.
Thedivergencebetweenbothtaxa wasfurtherpromotedwhen thelocalclimatereacheditsextantconditionsinthemiddleto
latePalaeolithicum(GibbardandvanKolfschoten2004;Royetal.
2004).Inparticular,the500mm-isohyetseparatestheLevant eco-logically fromNegev Desert and SinaiPeninsula (Yom-Tov and Ilani 1987), correspondingalsotodifferences invegetationand soilcharacteristics(Abdulsalametal.1988).Inthenorth(Eastern MediterraneanRegion)highandsubdued,heavilykarstified moun-tainsprevail,withconiferousforestsandshrublandsrepresenting thetypicalG.gazellahabitat.TheclimateisMediterraneanwith winterprecipitationofupto1800mm.Furthersouth(Western&
SouthernArabianRegion),whereG.arabicaoccurs,highvolcanic massifsandmountainchainswithxeromorphicshruband wood-landsprevail.Theclimateis tropicalwithtworainyseasons(at leastinthesouth)and precipitationbetween100and 450mm.
Therefore,wehypothesizeanecologicalspeciesboundarybetween southernG.arabicaadaptedtounpredictablerainfallpatternsand poorneedle-and feather-leavedshrubs (Vesey-Fitzgerald1952) andnorthernG.gazellaadaptedtoMediterraneanclimateenabling amorebroad-leaveddietincludingconsiderablequantitiesofgrass (Baharav1981,1983).
Intermsofconservationandre-introductioneffortstheadvice ofbreedingbothtaxaseparately(Wronskietal.2010)wasfully justified.Today,thesituationforbothtaxaiscritical.Thetotal num-berofG.gazellaisestimatedat3000individualsandhasdrastically declinedwithinthelasttwodecades(IUCN2008),whereasthetotal numberofG.arabicaleftontheArabianPeninsulaisestimatedat approx.11,000individuals(IUCN2008).
Acknowledgements
We would like to thank H. H. Prince Bandar bin Saud bin Mohammed al Saud for his support to conduct phylogenetic researchonwildlifein theKingdom.Weacknowledgethe sup-port rendered by the Steinhardt National Collection of Natural History,ZoologicalMuseumatTel-AvivUniversityandother
col-386 H.Lerpetal./MammalianBiology78 (2013) 383–386
lectors.Financialsupportcamefrom“LOEWE–Landesoffensivezur Entwicklungwissenschaftlich-ökonomischerExzellenz”ofHesse’s MinistryofHigherEducation,Research,andtheArts.
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Chapter 6
by Hannes Lerp, Martin Plath, Torsten Wronski, Eva V. Bärmann, Anna Malczyk, Revina-Rosa Resch, Bruno Streit and Markus Pfenninger