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Studies on Neotropical Fauna and Environment Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713817190 Molecular phylogeny of Megacephalina Horn, 1910 tiger beetles (Coleoptera: Cicindelidae)

First Published on: 13 June 2007 To cite this Article: Zerm, Matthias, Wiesner, Jürgen, Ledezma, Julieta, Brzoska, Dave, Drechsel, Ulf, Cicchino, Armando C., Rodríguez, Jon Paul, Martinsen, Lene, Adis, Joachim and Bachmann, Lutz , 'Molecular phylogeny of Megacephalina Horn, 1910 tiger beetles (Coleoptera: Cicindelidae)', Studies on Neotropical Fauna and Environment, 1 - 9 To link to this article: DOI: 10.1080/01650520701409235 URL: http://dx.doi.org/10.1080/01650520701409235

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The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 Tetracha eCeca xca auae,UiesddNcoa eMrdlPaa a e lt,Argentina, Plata, del Mar Plata, del Mar de Nacional Universidad Naturales, y Exactas Ciencias de nttt eeoaod netgcoe Cientı Investigaciones de Venezolano Instituto Rcie ac 07 cetd1 pi 2007) April 19 accepted 2007; March 8 (Received Norway Oslo, Oslo, of University noooi,MsoNe epfMrao at rz Bolı Cruz, Santa Mercado, Kempff Noel Museo Entomologia, orsodne .Zr,Toia clg,MxPac-nttt o vltoayBooy otah15 40 Plo 24302 165, online Postfach 1744-5140 Biology, print/ISSN the Evolutionary 0165-0521 for ISSN Max-Planck-Institute 2002). with Ecology, Tropical Zerm, al., taxon M. Correspondence: 1998; et pantropical Galian a Barraclough, 2001; & is & Vogler, (Vogler Megacephalina & Vogler cicindelids of Pearson 1996; clades group basal Pearson, diverse most the the as (Cicindelidae) within species 1910 100 Horn, over subfamily well Megacephalina with beetle subtribe relationships the tiger phylogenetic identified the the on within studies Recent Introduction non- independently The and America secondarily tree. colonized The phylogenetic then plains. the and salt Keywords: on margins coastal river and/or mapped habitats. inland to and coastal to confined taxa and/or moved first species uplands likely the Australian flooded most to The were uplands. assigned groups in non-flooded were American inhabit observed types species African are habitat basal groups Additionally, resulting group. species The monophyletic subgeneric outgroups. monophyletic of as Several placement well-supported used taxon. the is were paraphyletic classification traditional lineages system. South the from mostly classification the derived deviation 40 current and of than 18S the dendrogram. more nuclear with species the of the agreement in specimens cicindelid on in basal 60 based basically Three placed includes Megacephalina are dataset of taxa. trees The phylogeny Australian phylogenetic genes. molecular III a and 1910, oxidase provide in cicindelid American cytochrome back we basal who, and study the Horn 16S present to of contrast mitochondrial the diverse in most In is monospecific the This taxa). the subtribe. is taxa: other the genus-level species within two 100 genera only eight than conceded recognize more taxonomists by most Today, represented groups. Megacephalina subtribe pantropical The Abstract 1 ZERM MATTHIAS beetles tiger 1910 Cicindelidae) Horn, (Coleoptera: Megacephalina of phylogeny Molecular ARTICLE ORIGINAL article iFirst 1–9, Environment and 2007, Fauna Neotropical on Studies O:10.1080/01650520701409235 DOI: Tetracha OCI ADIS JOACHIM DRECHSEL rpclEooy a-lnkIsiuefrEouinr ilg,Plo Biology, Evolutionary for Max-Planck-Institute Ecology, Tropical / Aniara and Aniara Aniara ihntelte h monophyletic the latter the Within . 5 RAD .CICCHINO C. ARMANDO , utai,ccnei,habitat, cicindelid, Australia, , lofr oohltcgop h oiinof position The group. monophyletic a form also 1 UZBACHMANN LUTZ & 1 ,JU ¨ GNWIESNER RGEN Pseudotetracha # 07Tyo Francis & Taylor 2007 fcs eeul,and Venezuela, ´ficas, and Australicapitona Megacephala Aniara 8 6 2 O ALRODRI PAUL JON , UIT LEDEZMA JULIETA , sobrina ´via, and uhr seHbr 94frachronological these a genus reunited genus monospecific of Horn the for within Walther variety 1994 taxa 1910, a In Huber, by synopsis). established (see or genus were the authors at America, level subgroups Over several subgenus years South relationships controversial. 150 still last the the are in However, Megacephalina Africa. within diversity and Australia species highest h itrtxnto taxon sister the , 4 eaehl sensu Megacephala oohltcgop hlgntcte,South tree, phylogenetic group, monophyletic , als lrd,USA, Florida, Naples, carolina 8 aua itr uem eateto Zoology, of Department Museum, History Natural n Germany, ¨n, omamnpyei utainclade. Australian monophyletic a form Metriocheila and Aniara ´ brasiliensis GUEZ 2 on( eu hc ntdteseven the united which genus (a Horn ean neti.Tems striking most The uncertain. remains ofbr,Germany, Wolfsburg, within eaehl sensu Megacephala Megacephala 3 AEBRZOSKA DAVE , Aniara Pseudotetracha 5 7 Megacephala Asuncio EEMARTINSEN LENE , Tetracha n emn.Eal [email protected] Email: Germany. ¨n, ldstgte ersn a represent together clades hc hsrepresented thus which n Paraguay, ´n, and edrn h atra latter the rendering , 7 etod Ecologı de Centro , Grammognatha Phaeoxantha xetfrthe for except on Today, Horn. 3 Dep. Phaeoxantha 4 6 ULF , Facultad and are ´a, 8 , , Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 h egahcldsrbto ftegnr n h ubro pce i aetee)i ie codn oWiesner’s to according given modified). is and parentheses) extended (in but species (1994, of World Huber number following the the Megacephalina and of subtribe genera Beetles the the of of classification distribution genus-level geographical used The Currently (a) 1. Figure Ro see as (but described species initially to American was from rise known which taxon ‘‘ a amber, giving is there Baltic Europe north Further region. towards euphratica Grammognatha major moved two had one assumed Accordingly, intercontinental bridges. postulated land (1910) across routes Horn migration scenario his Megacephala of origin geographical Megacephala the Africa considered He charac- ters. morphological and considerations graphical of taxonomists history many by supported is 1a). (Figure this and World tion) the of Beetles Wiesner’s Tiger to subtribe according the within Megacephalina recognized genera eight are there al. et Zerm M. eaehl carolina Megacephala on(90 rpsdadtie evolutionary detailed a proposed (1910) Horn eaehl sensu Megacephala stetxnsms neta ieg.In lineage. ancestral most taxon’s the as n codnl ocie h African the conceived accordingly and 19;udt npeaain.()Gnslvlrltosisacrigt h rsn study. present the to according relationships Genus-level (b) preparation). in update (1992; ’ otmoayNorth contemporary a ’’, 19;udt nprepara- in update (1992; nteMediterranean the in onbsdo biogeo- on based Horn Megacephala shan 1999). ¨schmann, hcls of Checklist lineage oalneotropical all to suturalis Tetracha Tetracha ol nyb nesodi iwo h hoyof theory the of view in understood (2001) be of distribution only Vogler the could & that Pearson state explicitly outdated. 2 Figure in therefore were summarized as the are tectonics ideas plate developed his and of Horn forward, idea that put the mind before in well keep scenario to has One Australian the into developed branch neotropical the Australian into oped neotropical rise gave the branch One to Australia. both to which eventually branches, it two made into split lineage the point forming America Southern Megacephala ooie s‘‘ as ‘‘north- this colonized scenario (1910) ern’’ Horn’s to According Megacephala usqetysra ot forming south spread subsequently Australicapitona ieg oe ietyfo fiato Africa from directly moved lineage Tetracha hc oncniee ancestral considered Horn which , Phaeoxantha ieg oe etadand westward moved lineage Tetracha ’esenNrhAmerica. North eastern ’’ h other The . ercagermaini Tetracha Metriocheila pce.Tesecond The species. htfrhrdevel- further that hcls fteTiger the of Checklist Pseudotetracha Megacephala Megacephala n the and tthis At . . Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 2. .TeAmerican The 3. African The 1. (1910): Horn to by implicit forward are put that scenario hypotheses the several the allows of set within testing data the obtained genera the recognized Furthermore, subtribe. currently Megacephalina and of includes 1910 monophyly relating dataset questions the address The we to and taxa genes. 40 than (coxIII) more the on III cytochrome and based oxidase 16S is mitochondrial the that 18S, nuclear Megacephalina of phylogeny here. addressed are and valid still Horn are of (1910) implications continents. systematic some southern which Nevertheless, the megacontinent disconnected Gondwana ancient sequentially the break- of the to up specifically refer They drift. continental routes. dispersal evolution the of scenario of biogeographical (1910) Horn’s 2. Figure aao h oeua phylogeny: molecular the respective on the of type taxa habitat the mapped we Finally, 6. 5. 4. o.Within not. whereas habitats wetland Phaeoxantha Neither 7. ntepeetsuyw rvd molecular a provide we study present the In eaehl sensu Megacephala Grammognatha within clade ancestral taxa. Pseudotetracha of taxon sister Phaeoxantha clade. the within Australicapitona Phaeoxantha within Tetracha carolina Tetracha taxon. Tetracha. Tetracha pce,freape ooie mostly colonized example, for species, ersn h otdrvdspecies derived most the represent Tetracha / Australicapitona Megacephala onwt yohtclintercontinental hypothetical with Horn saprpyei ao with taxon paraphyletic a is Pseudotetracha. and stesse ao of taxon sister the is sacsrladteneotropical the and ancestral is hr r losome also are there Tetracha and Australicapitona and eaehl sensu Megacephala Megacephala Metriocheila ersnstemost the represents samonophyletic a is Phaeoxantha / Metriocheila Tetracha sensu pce did species Tetracha. er firme terra r sister are included Horn. Horn, sthe is nor and n 5 (pri- gene 18S and nuclear subsequently the 5 were i.e. mers: PCR, genes by three MN, amplified the of Minneapolis, of Sections Systems, protocol USA). (Gentra the kit following Puregene extracted was DNA sequencing DNA and amplification PCR (2001). al. et Zerm al. and et McCairns (1997) to (1997), Sumlin according (1999), of al. types assigned et Pearson habitat (accession were The species I). GenBank Table Megacephalina in from listed are retrieved numbers were outgroup these outgroups. of species sequences coxIII as and 16S used 18S, The were Megacephalina), subtribe discrepans pce eeaaye.Treccnei species, cicindelid Three analyzed. sub- Neocollyris or were species American species three of 31 specimens Australian, and some African/Eurasian, 60 nine different total, representing For In from Megacephalina collected. subgenera. specimens were subspecies, these localities biogeo- and the of cover species to range order in graphical selected were species AGAACTTGRCC-3 oIIgn pies 5 (primers: gene coxIII Fgr a eeicue nteaaye favailable. if analyses the For in Megacephalina included of were genera 1a) (Figure eight the I. of Table species in listed Several are study this in included species The sampling Taxon methods and Materials and derived monophyletic is preference are ancestral. habitat or adaptations observed the these whether not whether question the or address we and species, upland a sdfretbihn utpesequence multiple establishing (Applied 1997) for chemistry al., used et BigDye (Thompson was the ClustalX CA, Biosystems). using City, Foster PCR automatic USA) Biosystems, ABI310 and QIAquick (Applied an Germany), sequencer on the Hilden, sequenced of (Qiagen, subsequently kit protocol on purification purified differed provided the were be products protocols following PCR can obtained PCR regions and The request. The species conserved between genes. slightly the highly the available on publicly within study targeting of this alignments sequences, of sequence purpose of the basis for specifically TATCTACCTCATC-3 h iohnra 6 ee(rmr:5 (primers: gene 16S mitochondrial the CATCGAGGTCGCAA-3 GTATTTTGACTGTGC-3 Phaeoxantha 9 -TTGTCTCAAAGATTAAGCCATGC-3 9 -CCGTTACAACCATGGTAGGCG-3 ta r nydsatyrltdt the to related distantly only are (that sp., dnohiaconfusa Odontocheila , Tetracha 9 n 5 and ’ 9 9 hlgn fMegacephalina of Phylogeny -CTGTAGAAYTWGGR- .Pieswr designed were Primers ). 9 and n h mitochondrial the and ) 9 9 -TACTATATAAAT- n 5 and Pseudotetracha 9 and -TAATCCAA- Pentacomia 9 -CCGA- ,many 9 ), 9 Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 2Bai,Aaoa,na Manaus: near Amazonas, Brazil, 18 Genus Rio Pantanal, Grosso, Rı Mato Cruz, Brazil, Santa 17 Ichilo, Bolivia, 16 Genus Rı Cruz, Santa Cordillera, Bolivia, 15 Rı Carrasco, Bolivia, 13 12 11 10 9 8 7 6 5 4 3 Genus 2 20 19 Genus Genus 14 site Collecting Genus Genus name Species study. this in used specimens beetle Tiger I. Table al. et Zerm M. 1 22 21 25 24 23 28 26 9Prga,Dp Boquero Dep. Paraguay, 29 Genus 27 1848) regalis regalis Megacephala citernii regalis Megacephala 1822) Dejean, euphratica Grammognatha lindemannae Phaeoxantha limata Phaeoxantha klugii Phaeoxantha cruciata Phaeoxantha (Brulle bifasciata aequinoctialis Phaeoxantha 1825) (Dejean, aequinoctialis aequinoctialis Phaeoxantha nigricollis Metriocheila suoercacorpulenta Pseudotetracha 1895) blackburni Pseudotetracha 1841) australasiae Australicapitona hexnh wimmeri Phaeoxantha naasepulcralis Aniara suoercahelmsi Pseudotetracha 1997) cuprascens Pseudotetracha suoercaoleadorsa Pseudotetracha mendacia Pseudotetracha ion Pseudotetracha 1907) brevisulcata affinis Tetracha pulchra Pseudotetracha suoercawhelani Pseudotetracha Australicapitona Megacephala Phaeoxantha Pseudotetracha Grammognatha Metriocheila Aniara Tetracha ,1837) ´, oe1938 Hope oe1838 Hope hmo 1857 Thomson huor1850 Chaudoir aril 1902 Latreille Fbiis 81 rzl mzns erMnu:Reserva Manaus: near Amazonas, Brazil, 1801) (Fabricius, luiu,1894 Fleutiaux, Smi,19)Asrla .. .Al-ae d /3D129 Q507DQ152153 DQ152037 DQ152096 4/03 Rd. Auld-Wapet L. W.A., Australia, 1997) (Sumlin, oshlk 1850 Motschulsky Cadi,15)Bai,Aaoa,na aas Rio Manaus: near Amazonas, Brazil, 1850) (Chaudoir, uln 1992 Sumlin, Pry 80 rzl aoGos,SnoAde Rio Andre, Santo Grosso, Mato Brazil, 1830) (Perty, Bakun 82 utai,WA,Lk nen(S Annean Lake W.A., Australia, 1892) (Blackburn, (Brulle Rih,14)Blva arso at rz Rı Cruz, Santa Carrasco, Bolivia, 1842) (Reiche, Mnl 98 oii,Crac,Ccabma Rı Cochambamba, Carrasco, Bolivia, 1958) (Mandl, Bon 89 utai,WA,L oni SSnsoe /3D129 Q500DQ152156 DQ152040 DQ152099 4/03 Sandstone) (S Noondie L. W.A., Australia, 1869) (Brown, Smi,19)Asrla .. rnSoeLgo /3D120 Q501DQ152157 DQ152041 DQ152100 3/03 Lagoon Stone Iron S.A., Australia, 1992) (Sumlin, Smi,19)Asrla .. .Bre WMnis /3D129 Q509DQ152155 DQ152039 DQ152098 4/03 Menzies) (W Barlee L. W.A., Australia, 1992) (Sumlin, Smi,19)Asrla .. aeBlad(4k NE km (24 Ballard Lake W.A., Australia, 1997) (Sumlin, Hr,10)Asrla .. 3k utnSg /3D129 Q504DQ152150 DQ152034 DQ152093 3/03 Spgs Curtin E km 33 N.T., Australia, 1907) (Horn, (Sumlin, (Fleutiaux, Mnl 94 rzl mzns erManaus: near Amazonas, Brazil, 1964) (Mandl, Ltele& (Latreille (Boheman, Hr,11)Kna atr rvneHl /1D128 Q500DQ152148 DQ152030 DQ152089 5/01 Hola Province Eastern Kenya, 1912) (Horn, ,13)Prga,Dp ocpin a Carlos, San Concepcion, Dep. Paraguay, 1837) ´, (Horn, (Hope, osaa rnitw 20 Q500DQ152031 DQ152090 12/01 DQ152147 DQ152029 DQ152088 5/02 Francistown Botswana, Atlit Plain, Coastal North Israel, Sossego Bom Sitio Grande Volta Praia Cuiaba: Cacares Seca, Carne Praia Paraguay, Marchantaria de Ilha Solimoes, Grande Praia Rio Negro, Manaus: near Amazonas, Brazil, Paciencia de Ilha Rio Solimoes, Manaus: near Amazonas, Brazil, Blanquito i Apa Rio utai,WA,L ery(EKlori)40 Q502D123 DQ152149 DQ152033 DQ152092 4/03 Kalgoorlie) (SE Lefroy L. W.A., Australia, iha ,ofAm od(ETownsville) (SE Road Aims N off km 1, 5 Highway E, km 2 Queensland, Australia, Cesarsama Grande Praia Negro, Rio Ducke utai,WA,L or,2 mSE Find km 24 Payne Moore, L. W.A., Australia, Menzies) Meekathara) rv nr Rı Entre Colo Prov. Dep. Urquiza, Arroyo Argentina, ´os oCsraa1/2D127 Q508DQ152136 DQ152018 DQ152077 12/02 Cesarsama ´o n aPti 10 Q512D124 DQ152159 DQ152043 DQ152102 11/02 Patria La ´n, oSrt 00 Q502D122 DQ152141 DQ152023 DQ152082 10/02 Surutu ´o ´o oSc /2D128 Q501DQ152139 DQ152021 DQ152080 3/02 Seco ´o ´o ´n, Collecting 08 Q505D122 DQ152144 DQ152026 DQ152085 DQ152143 DQ152025 DQ152084 10/82 DQ152142 DQ152024 DQ152083 6/99 9/98 DQ152140 DQ152022 DQ152081 10/97 DQ152137 DQ152019 DQ152078 6/98 DQ152135 DQ152017 DQ152076 10/97 DQ15213 DQ152016 DQ152075 12/02 00 Q509D122 DQ152138 DQ152020 DQ152079 10/02 10 Q501DQ152032 DQ152091 11/02 20 Q507D122 DQ152146 DQ152028 DQ152087 DQ152145 DQ152027 DQ152086 12/02 3/98 08 Q504D121 DQ152133 DQ152015 DQ152074 10/82 /3D129 Q505DQ152151 DQ152035 DQ152094 4/03 /3D129 Q508DQ152154 DQ152038 DQ152097 4/03 /3D129 Q506DQ152152 DQ152036 DQ152095 4/03 /2D120 Q502DQ152158 DQ152042 DQ152101 2/02 date 8 6 coxIII 16S 18S eBn numbers GenBank Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 59 60 al .Continued. 1. Table ihpeiul salse lgmnsta a be can three congruent that alignments and all established straightforward previously of was with sequences genes the marker Aligning alignments. a DQ152166 DQ152050 DQ152109 11/02 Ravelo, Cruz, Santa Cordillera, Bolivia, California Baja Mexico, 43 42 41 40 site Collecting 39 38 37 36 30 name Species Outgroups 44 Colo Dep. Urquiza, Arroyo Argentina, Carlos, San Concepcion, Dep. Paraguay, Pirizal: Pantanal, Grosso, Mato Brazil, 33 32 31 45 34 35 46 7Blva odlea at rz Camiri Cruz, Santa Cordillera, Bolivia, 47 8Agnia rooUqia e.Colo Dep. Urquiza, Arroyo Argentina, 48 4Pet io1/2D122 Q507DQ152183 DQ152067 DQ152126 11/02 Rico Puerto Rio Manaus: near Amazonas, Brazil, 58 57 56 55 54 53 52 51 9Prga,Dp ocpin at oi 00 Q511D126 DQ152178 DQ152062 DQ152121 10/02 Sofia Santa Concepcion, Dep. Paraguay, 50 49 http://www.bio.ic.ac.uk/research/tigerb/benzpaper.htm etcmadiscrepans Pentacomia confusa Odontocheila Neocollyris ercathomsoniana Tetracha ercavirginica Tetracha ercafulgida Tetracha 1916) floridana Tetracha distinguenda Tetracha 1997) Wiesner, chacoensis Tetracha 1834) chilensis carolina Tetracha carolina carolina Tetracha brasiliensis Tetracha ercafuliginosa Tetracha ercanicaraguensis Tetracha camposi Tetracha ercacarolina Tetracha ercapilosipennis Tetracha ercaspixi Tetracha spinosa Tetracha sparsimpunctata Tetracha 1851) sommeri sobrina Tetracha sobrina sobrina Tetracha 1835) punctata sobrina Tetracha 1905) pseudodistinguenda Tetracha p ni,GnakL62 F395AF438930 AF438905 L76822 GenBank India, sp. (Brulle Ku,13)Blva Andre Bolivia, 1834) (Klug, (Brulle Hr,10)Eudr aai rct /1D120 Q508DQ152164 DQ152048 DQ152107 2/01 Crucita Manabi, Ecuador, 1900) (Horn, (Linne (Linne Ln Mutchler, & (Leng Bts 84 iaau 10 Q516D125 DQ152173 DQ152057 DQ152116 11/02 Nicaragua 1874) (Bates, Dja,12)Blva eBn 784A480 AF438928 AF438903 L76824 GenBank Bolivia, 1825) (Dejean, Swd & (Sawada Kry 88 rzl aoGos,Pnaa,Przl Baia Pirizal: Pantanal, Grosso, Mato Brazil, 1818) (Kirby, ,13)Blva lrd,SnaCu,Rı Cruz, Santa Florida, Bolivia, 1837) ´, Hr,19)Blva eBn 783A480 AF438927 AF438902 L76833 GenBank Bolivia, 1893) (Horn, Mnl 98 cao,Psaa uo aeo /1D121 Q509DQ152175 DQ152059 DQ152118 2/01 Canelos Puyo, Pastaza, Ecuador, 1958) (Mandl, Dja,13)Prga,Dp t.Hys rn Chaco Trans Hayes, Pte. Dep. Paraguay, 1831) (Dejean, Hr,11)Blva lrd,SnaCu,Pmarne10 Q511D127 DQ152188 DQ152072 DQ152131 1/03 Pampagrande Cruz, Santa Florida, Bolivia, 1915) (Horn, Jhsn 93 iaau 10 Q517D125 DQ152174 DQ152058 DQ152117 11/02 Nicaragua 1993) (Johnson, ,13)Bai,Aaoa,na aas Rio Manaus: near Amazonas, Brazil, 1837) ´, ,16)UA eBn 781L42918 L76821 GenBank USA, 1766) ´, ,16)UA ass1/2D123 Q503DQ152189 DQ152073 DQ152132 11/02 Kansas USA, 1766) ´, Dja,13)Vnzea agrt sad Playa Island, Margarita Venezuela, 1831) (Dejean, (Chaudoir, Mnl 91 oii,O atetbn at rz Rı Cruz, Santa Santiesteban, O. Bolivia, 1961) (Mandl, (Castelnau, (Linne (Laporte, (Horn, ,16)UA ea 10 Q518D124 DQ152165 DQ152049 DQ152108 11/02 Texas USA, 1766) ´, S,Foia1/2D121 Q505DQ152171 DQ152055 DQ152114 11/02 DQ152167 DQ152051 DQ152110 Florida USA, laguna de 2/01 Orilla Boquero Dep. Paraguay, Crucita Manabi, Ecuador, iha,k 415 km Highway, rv nr Rı Entre Prov. Apa Rio Novo Retiro Fazenda ubaa‘‘Forestal’’ Quebrada Coqueiro de rla rı orillas rv nr Rı Entre Prov. oios lad Paciencia de Ilha Solimoes, Eisenhower Puente Pirai Praia Grande Cuiaba: Volta Rio Andre, Santo Brazil, Grande Praia Negro, Marchantaria de Ilha Rio Solimoes, Manaus: near Amazonas, Brazil, Parguito aau,RoVerde Rio Lolita, Yaragui, Hayes, Pte Dep. Paraguay, oParapeti ´o sIban ´s ´os ´os e,SnaCruz, Santa ˜ez, n aPti /3D121 Q502DQ152168 DQ152052 DQ152111 5/03 Patria La ´n, meilClee odn UK. http://www.bio. London, as College, Imperial such Vogler, A. by websites ic.ac.uk/research/tigerb/benzpaper.htm from retrieved oCue /3D123 Q501DQ152187 DQ152071 DQ152130 1/03 Cruces ´o ´n, ´n, ´o Collecting /3D121 Q504DQ152170 DQ152054 DQ152113 4/03 /3D121 Q503DQ152169 DQ152053 DQ152112 1/03 00 Q515D124 DQ152162 DQ152046 DQ152105 DQ152161 DQ152045 DQ152104 10/02 6/99 /2D121 Q506DQ152172 DQ152056 DQ152115 4/02 DQ152160 DQ152044 DQ152103 8/00 /3D120 Q507DQ152163 DQ152047 DQ152106 2/03 /3D121 Q500DQ152176 DQ152060 DQ152119 4/03 19 Q519D127 DQ152186 DQ152070 DQ152129 DQ152185 DQ152069 DQ152128 11/97 DQ152184 DQ152068 DQ152127 5/03 6/99 DQ152181 DQ152065 DQ152124 DQ152180 8/98 DQ152064 DQ152123 11/97 /2D122 Q506DQ152182 DQ152066 DQ152125 2/02 /3D122 Q503DQ152179 DQ152063 DQ152122 3/03 /3D122 Q501DQ152177 DQ152061 DQ152120 2/03 date hlgn fMegacephalina of Phylogeny 8 6 coxIII 16S 18S eBn numbers GenBank a Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 oa ubr niaewl-upre ugnrcgop setx) rosidct h curneo orteho h ih mandible right uplands). the non-flooded sequences. on t, teeth flats; 18S four salt and of inland occurrence s, 16S habitats; the coastal coxIII, indicate habitat c, Arrows the of floodplains; text). as shores, well (see consisting as lake groups brackets set and within subgeneric in rivers given well-supported data of are indicate margins I combined numbers Table (r, from Roman the taxa codes respective running of Specimen the nodes. of inference respective type the Bayesian in at indicated by are probabilities reconstruction Posterior given Phylogenetic 3. is Figure tree resulting inference The Bayesian subsequent was reconstruction. the model for GTR applied This then sites. certain invariant a a of and with the proportion heterogeneity model as rate GTR well distributed a as gamma always separately preferred was MrModeltest genes optimal dataset The three combined set. estimate the data for to the model First, for used model was substitution 2002) analyses. (Nylander, for used subsequently phylogenetic was alignment obtained The 1910 Megacephalina subtribe Horn, the in relationships in Phylogenetic in listed deposited are the were to numbers bp 422 accession I. sequences Table and The All gene GenBank. 18S 16S gene. the the to to coxIII relate bp 364 bp 359 all gene, bp excluded); 1145 of total (primers in alignment from long and The straightforward was determined. specimens genes three be the could genes For tralasiae regalis specimens. regalis obtained 18S, Megacephala 58 were nuclear genes the coxIII for of and 16S sets mitochondrial data sequence Complete data molecular of Analysis Results their on based assessed frequencies. were observed clades trees individual the 000 of 30 From burn-in. 20% as resulting initial discarded taking were The (8000) trees generations. chains, the 100 and Markov every generations samples 000 Metropolis-coupled 000 4 analyses inference four under gamma Bayesian performed a sites. were with propor- invariant certain of model a tion and GTR heterogeneity preferred a rate The distributed always methods. was probabilistic model & the for (Posada models with optimal 3.06 find use to Modeltest used was of 1998), Crandall, version simplified 2002), (Nylander, a MrModeltest reconstructions (Huelsenbeck 2001). 3.04b Ronquist, inference & MrBayes Bayesian with (Kumar conducted 2.1 2001). were version al., MEGA et with performed were analyses Parsimony Maximum and Neighbor-joining analyses Phylogenetic al. et Zerm M. Tetracha nytesqecso h 6 n 18S and 16S the of sequences the only P., . Phaeoxantha posteriori a Ps., ; and Pseudotetracha utaiaioaaus- Australicapitona rbblte for probabilities . h utiu eaehln,terslso the of follows: results as summarized the be can analyses Megacephalina, phylogenetic subtribus the the outgroup observed. in were genera the the differences within species minor among of grouping only basal Otherwise, very species. positioned was The exception, shown). American (not one topologies South tree respectively. with same the yielded, basically Maximum analyses analyses, and and Parsimony Neighbor-joining Neighbor-joining the Parsimony Nevertheless, in support Maximum and bootstrap inference Bayesian low after probabilities posterior to close clade own its only forms study the is which 18S and Megacephala 16S regalis 1. the Figure regalis in depicted M. of not therefore consisting are and sequences set data reduced and of exception the dendrogram. the the of genera with Aniara recognized Megacephalina currently subtribus All 3. Figure N N N N N ihrsett h urnl eonzdgnr of genera recognized currently the to respect With low by supported only are clades some However, h hlgntcrltosisof relationships phylogenetic The oehrwt oeafnt oteAustralian the to affinity clade. some Pseudotetracha with together klugii-lindemannae-wimmeri consist- clades (1) is, of distinct prob- ing three that posterior comprises low It group by ability. supported monophyletic only a however, form species eai regalis regalis Megacephala Grammognatha Grammognatha Phaeoxantha and analyses). Neighbor-joining Parsimony the the in Maximum clade this and different of the for position Tetracha/Aniara above see (but the clade Phaeoxantha to affinity species, Metriocheila Aniara aa ntetree. the in basal onaprpyei taxon. paraphyletic a Horn Tetracha sepulcralis .australasiae A. and oe 98 h only the 1938: Hope, .nigricollis M. pce hc makes which species Tetracha ,isnestedwithinthecladeofall aequinoctialis-cruciata taxon, hmo,15:teonly the 1857: Thomson, huor 80 the 1850: Chaudoir, aril,10:teAfrican the 1902: Latreille, rue oehrwt h second the with together grouped Metriocheila and species, oshlk,15:teonly the 1850: Motschulsky, .rglscisternii regalis M. Australicapitona ommnpyei rusin groups monophyletic form eeol eue iha with deduced only were .rglscisternii regalis M. om t w ld with clade own its forms .euphratica G. . species (2) , Aniara eaehl sensu Megacephala Pseudotetracha .rglsregalis regalis M. . pce nthis in species .australasiae A. limata .nigricollis M. Phaeoxantha Metriocheila species, sfound is , n (3) and cluster . M. A. Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 hlgn fMegacephalina of Phylogeny Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 N N N .Zr tal. et Zerm M. swl sa mrcncaecomprising clade American an as well Australian as an of comprising monophyly clade the provides for also study evidence current strong Our tree. the in positions Tetracha genera African the represent good that in Megacephala species the are i.e. 3) (Figure system. genes classification this The 18S with and agreement coxIII genera. nuclear Megacephalina 16S, mitochondrial eight the the the on based of of species (see relationships consists Huber extensions/modifications phylogenetic and of some 1a), that Figure with follows the (1994) of Megacephalina classification genus-level subtribe used currently The Discussion lsiiaini etil h lcmn of placement the within certainly is classification foeo two of (2002). one al. of that et found of Galian on authors by phylogeny (based The data) molecular levels rRNA the 18s taxonomic the full-length with higher on at consistent study were Cicindelidae a is markers of three This results the separately. when even observed analyzed was analyses, finding this all that in noted be can It taxon. sexpected, As h otsrkn eito rmtetraditional the from deviation striking most The .camposi T. fulgida chacoensis brasiliensis carolina chilensis of in consisting genus groups found the species were monophyletic makes Four 18S clade paraphyletic. and the 16S within the of Tetracha analyses the sequences. on only but Pseudotetracha the of species, positioning one by represented occur Australicapitona groups probability. the species posterior within high well-supported by Several supported is that Pseudotetracha Pseudotetracha guenda .pilosipennis T. i o omwl-upre ldswt other with clades well-supported Tetracha form not did Tetracha , , Metriocheila and , 2 the (2) , .thomsoniana T. .floridana T. oe 88 h etn of nesting the 1838: Hope, species. n 4 the (4) and , , and ld ossigof consisting clade Pseudotetracha .spinosa. T. Megacephala .affinis T. aigtelte paraphyletic a latter the making , .rglsciternii regalis M. .sobrina T. , sntbsdo opeedt set, data complete a on based not is pce om oohltcclade monophyletic a forms species Pseudotetracha Grammognatha luiu,19:teAustralian the 1894: Fleutiaux, .distinguenda T. uln 92 h eu sonly is genus the 1992: Sumlin, Tetracha carolina Australicapitona and , and T.fuliginosa Aniara oeohrseissc as such species other Some fulgida , Phaeoxantha , ..seis hs nthis in Thus, species. s.l. 1 the (1) : .spixi T. .nicaraguensis T. clade. .sparsimpunctata T. ld ossigof consisting clade and .australasiae A. .brasiliensis T. a h itrtaxon sister the was r on tbasal at found are , ld ossigof consisting clade and , Australicapitona .pseudodistin- T. and and ld ls to close clade .euphratica G. . sobrina .sepulcralis A. .virginica T. .carolina T. Tetracha 3 the (3) , Aniara Aniara and The . clade , T. T. T. , , , .mendacia P. ieaue ae nmrhlgclcharacters morphological on describes (1997) Sumlin Based literature. Pseudotetracha the test to comprehensive hypothesis. required be development. of would datasets a interpretation morphological incidental; independent be analytical just proper may an congruence this of However, mandible considered right the are on teeth four the the that sense makes morphology. it virginica different that a believe therefore with may but One teeth four has also the as well as ofre ntemlclrpyoey carset described McCairns (1997) phylogeny. al. molecular the in a of confirmed relative as considers closest also study) (1997) the Sumlin group present analyses. polyphyletic our a the in is it in However, complex. included species not species ruswr bevd hs rusaecongruent the are considerations: groups morphological with These observed. were groups characters. ethological or ecological logical, of of branch placement puzzling study et saprnl h lsoopi tt within state plesiomorphic mandible, three the the with right Tetracha apparently in mandible is the documented right teeth A been on literature. never teeth taxonomic has this three four although having of for morphologists instead by together grouped eaehl yHr 11)aefliidb the incorpora- the by of that unlikely falsified tion very is are It (1910) results. of present Horn scenario evolutionary by the Megacephala to implicit hypotheses African species. separated true as treated be thus should the to close found other is the but with group not does species: be may true subspecies two considered sequences coxIII the and 16S to related closely as oleodorsa here) considered not species h rsn eut,i atclrtemnpyyof topology. monophyly the tree particular in observed results, present the The of substantially evolution altered the Pseudotetracha for central iia iuto a bevdi h genus the chilensis in carolina observed with group was Phaeoxantha situation similar h ruigo eea pce within species several of grouping The Within ihteecpino h aa oiino the of position basal the of exception the With mitochondrial the 18S, nuclear the to According sobrina Megacephala Megacephala / ercagermaini Tetracha osntcutrtgte ihteseisof species the with together cluster not does Aniara hc sas o orbrtdb u data. our by corroborated not also is which Tetracha , Tetracha : carolina .aqiotai aequinoctialis aequinoctialis P. and oee,i o ossetwt the with consistent not is however, , .aqiotai bifasciata aequinoctialis P. h ih adbeof mandible right The . nHr’ cnro ol have would scenario, Horn’s in carolina swl as well as .cuprascens P. ..wsprpyei swl.The well. as paraphyletic was s.l. sntcrooae ymorpho- by corroborated not is .helmsi P. and eea oohltcspecies monophyletic several and .whelani P. sobrina .blackburni P. and brasiliensis Aniara Grammognatha ercacrln chilensis carolina Tetracha .aqiotai bifasciata aequinoctialis P. nti td,aspecies a study, this in aogwt oeother some with (along brasiliensis .carolina T. a ela another as well (as ld of clade ihnaderived a within Phaeoxantha hc sas not also is which ldsadthat and clades , .corpulenta P. sobrina Tetracha .virginica T. .virginica T. pce,all species, ldsare clades subspecies Both . osnot does .ion P. clade and .A T. T. P. , Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 a-lnkIsiuefrEouinr Biology, Junk, Evolutionary Wolfgang for thank We Max-Planck-Institute specimens. of collection Valdivia, Montan Alejandra Jose Garcia, Rossy Gutierrez, Marcos Ivan Teresa Werner, Aramayo, Frank Orbach, Onore, Giovanni Eylon to grateful are We Acknowledgments independently habitats. coastal and and/or uplands secondarily non-flooded colonized species American sepulcralis is some it that interpretation this likely to According confined but margins. mainly river plains are to salt however, coastal species, American and/or the species inland to species African moved Australian have the (4) basal uplands, and non-flooded The plains, inhabit salt (1) uplands. floodplains, inland and (3) i.e. non-flooded shores, habitats, types, lake coastal and (2) habitat rivers of different we margins phyloge- purpose four this the For to 3. considered Figure on in presented mapped tree also netic were study under conclusions. detailed more for this allow for not sampling does the future study detail, phylogeo- more in However, intraspecific patterns such graphic address biogeography. to topology from need the tree studies separated This reflects is (D). also that specimen group Ecuadorian the one form Paraguay nis ahohr(.Zr,proa observation). personal Zerm, (M. other each adults laboratory, the of in kept When species. different wimmeri all either P. that of system. specimens however, sampled river implies, This different specimens distribution. the a of the grouping of the in mind, part in Pantanal this the is Having to close which Both Paraguay basin Rio was (Brazil). and 10 I; the (Table Specimen Manaus at Bolivia basin. collected Amazon near the 9 eastern to 13 belong areas in and and 8 collected specimens 14 were 3) Figure specimens phylogeo- the interesting Within Phaeoxantha, emerge. two patterns level in graphic intraspecific clades the subgeneric at the results as present well Tetracha relationships as the supragenus-level ancient of 1b) (Figure these on the classification reflect of should future based break-up A Megacephalina the continent. Megacephalina during of Gondwana evolution place strongly early took clades, the Australian that and suggest American the both h aia rfrne fteMgcpaiataxa Megacephalina the of preferences habitat The h eodisac eae to relates instance second The limited, is set data our of resolution the Although pcmn rmBlva retn and Argentina Bolivia, from Specimens . .klugii P. klugii , Phaeoxantha n oalse xetsome extent lesser a to and and ersn h aeo,alternatively, or, same the represent oadMtha ate o the for Wantzen Matthias and ˜o ld elcsterbiogeographical their reflects clade .lindemannae P. and .klugii P. Pseudotetracha Tetracha , edl ouaewith copulate readily .lindemannae P. ercapilosipen- Tetracha species, klugii Phaeoxantha Fgr 3). (Figure ld of clade ´Luis Aniara and References emM dsJ aranW mrmM,FneaC de. CR Fonseca MA, Amorim W, Paarmann J, Adis M, Zerm Sandlaufka der Verzeichnis 1992. J. Wiesner tiger the of phylogeny molecular A 1996. DL. Pearson AP, Vogler in shifts Reconstructing 1998. TG. F, Barraclough Jeanmougin AP, Vogler F, Plewniak TJ, Gibson JD, Thompson to Additions 1997. WD. Sumlin rmteRsac oni fNra grant 146515/ knowledge the of Norway towards Biosystematics’’, contribution 91st of for J.W.: 420); Council and Centre Research the help (‘‘National from support the their grants acknowledges from for by L.B. MPI supported the Max-Planck-Society. was the and M.Z. at Reusch support. staff Thorsten thanks laboratory M.Z. support. Germany/Plo Ro model the testing MODELTEST: evolution, 1998. KA. the Crandall D, Posada beetles: Tiger 2001. AP. Vogler DL, Pearson of beetles tiger http:// The 1999. from: DW. Brzoska JF, Available Guerra DL, 1.1. Pearson MrModeltest 2002. J. Nylander carsR,FetgR oeHA Rose R, MEGA2: Freitag 2001. RF, M. McCairns Nei IB, Bayesian Jakobsen K, MRBAYES: Tamura 2001. S, Kumar F. Ronquist JP, Huelsenbeck coast west the from Tetracha of species new A 1994. RL. Huber Subfam. Carabidae, Fam. 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Evol Biol Mol beetles. tiger in chromosomes ihnts(oepea.Ccneia:Bull Cicindelidae: (Coleoptera). notes with n(P)frlgsi n financial and logistic for (MPI) ¨n . hlgn fMegacephalina of Phylogeny eaehl,Nickerlea Megacephala, ´ Cicindela coadFD 1997. FJD. McDonald , netb Taxon Invertebr . frdrWelt. der ¨fer and ,