This article was downloaded by:[Max Planck Inst & Research Groups Consortium] [Max Planck Inst & Research Groups Consortium] On: 17 June 2007 Access Details: [subscription number 770286779] Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK 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 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article maybe used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. 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 Studies on Neotropical Fauna and Environment 2007, 1–9, iFirst article ORIGINAL ARTICLE Molecular phylogeny of Megacephalina Horn, 1910 tiger beetles (Coleoptera: Cicindelidae) MATTHIAS ZERM1,JU¨ RGEN WIESNER2, JULIETA LEDEZMA3, DAVE BRZOSKA4, ULF DRECHSEL5, ARMANDO C. CICCHINO6, JON PAUL RODRI´GUEZ7, LENE MARTINSEN8, JOACHIM ADIS1 & LUTZ BACHMANN8 1Tropical Ecology, Max-Planck-Institute for Evolutionary Biology, Plo¨n, Germany, 2Wolfsburg, Germany, 3Dep. Entomologia, Museo Noel Kempff Mercado, Santa Cruz, Bolı´via, 4Naples, Florida, USA, 5Asuncio´n, Paraguay, 6Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata, Mar del Plata, Argentina, 7Centro de Ecologı´a, Instituto Venezolano de Investigaciones Cientı´ficas, Venezuela, and 8Natural History Museum, Department of Zoology, University of Oslo, Oslo, Norway (Received 8 March 2007; accepted 19 April 2007) Abstract The pantropical subtribe Megacephalina represented by more than 100 species is the most diverse of the basal cicindelid groups. Today, most taxonomists recognize eight genera within the subtribe. This is in contrast to Horn who, back in 1910, conceded only two genus-level taxa: the monospecific Aniara and Megacephala sensu Horn (a genus which united the seven Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 other taxa). In the present study we provide a molecular phylogeny of Megacephalina based on the nuclear 18S and the mitochondrial 16S and cytochrome oxidase III genes. The dataset includes 60 specimens of more than 40 mostly South American and Australian taxa. Three cicindelid species of derived lineages were used as outgroups. The resulting phylogenetic trees are basically in agreement with the current classification system. Megacephala and Grammognatha are placed basal in the dendrogram. Pseudotetracha and Australicapitona form a monophyletic Australian clade. Phaeoxantha, Tetracha and Aniara also form a monophyletic group. The position of Metriocheila remains uncertain. The most striking deviation from the traditional classification is the well-supported placement of Aniara within Tetracha, rendering the latter a paraphyletic taxon. Several monophyletic subgeneric species groups are observed in Pseudotetracha, Phaeoxantha and Tetracha/Aniara. Within the latter the monophyletic sobrina, carolina and brasiliensis clades together represent a monophyletic group. Additionally, habitat types were assigned to the taxa and mapped on the phylogenetic tree. The basal African species inhabit non-flooded uplands. The Australian species moved to inland and/or coastal salt plains. The American groups were most likely first confined to river margins and then colonized secondarily and independently non- flooded uplands and/or coastal habitats. Keywords: Aniara, Australia, cicindelid, habitat, Megacephala, monophyletic group, phylogenetic tree, South America Introduction highest species diversity in South America, Australia and Africa. However, the relationships Recent studies on the phylogenetic relationships within Megacephalina are still controversial. Over within the tiger beetle subfamily (Cicindelidae) the last 150 years several subgroups at the genus or identified the subtribe Megacephalina Horn, 1910 subgenus level were established by a variety of with well over 100 species as the most diverse group authors (see Huber, 1994 for a chronological within the basal clades of cicindelids (Vogler & synopsis). In 1910, Walther Horn reunited these Pearson, 1996; Vogler & Barraclough, 1998; taxa within the genus Megacephala except for the Pearson & Vogler, 2001; Galian et al., 2002). monospecific genus Aniara which thus represented Megacephalina is a pantropical taxon with the the sister taxon to Megacephala sensu Horn. Today, Correspondence: M. Zerm, Tropical Ecology, Max-Planck-Institute for Evolutionary Biology, Postfach 165, 24302 Plo¨n, Germany. Email: [email protected] ISSN 0165-0521 print/ISSN 1744-5140 online # 2007 Taylor & Francis DOI: 10.1080/01650520701409235 M. Zerm et al. there are eight genera recognized within the subtribe According to Horn’s (1910) scenario this ‘‘north- Megacephalina according to Wiesner’s Checklist of ern’’ Megacephala lineage moved westward and Tiger Beetles of the World (1992; update in prepara- colonized as ‘‘Tetracha’’ eastern North America. tion) and this is supported by many taxonomists Tetracha subsequently spread south forming (Figure 1a). Tetracha suturalis, which Horn considered ancestral Horn (1910) proposed a detailed evolutionary to all neotropical Tetracha species. The second history of Megacephala sensu Horn based on biogeo- Megacephala lineage moved directly from Africa to graphical considerations and morphological charac- Southern America forming Tetracha germaini. At this ters. He considered Africa the geographical origin of point the lineage split into two branches, which both Megacephala and accordingly conceived the African made it eventually to Australia. One branch gave rise Megacephala as the taxon’s most ancestral lineage. In to the neotropical Phaeoxantha that further devel- his scenario Horn (1910) assumed two major oped into the neotropical Metriocheila and the migration routes across postulated intercontinental Australian Australicapitona. The other Megacephala land bridges. Accordingly, one Megacephala lineage branch developed into the Australian Pseudotetracha. had moved towards Europe giving rise to One has to keep in mind that Horn developed the Grammognatha euphratica in the Mediterranean scenario well before the idea of plate tectonics were region. Further north there is a taxon known from put forward, and his ideas as summarized in Figure 2 Baltic amber, which was initially described as are therefore outdated. Pearson & Vogler (2001) ‘‘Megacephala carolina’’, a contemporary North explicitly state that the distribution of Megacephala American species (but see Ro¨schmann, 1999). could only be understood in view of the theory of Downloaded By: [Max Planck Inst & Research Groups Consortium] At: 05:32 17 June 2007 Figure 1. (a) Currently used genus-level classification of the subtribe Megacephalina following Huber (1994, but extended and modified). The geographical distribution of the genera and the number of species (in parentheses) is given according to Wiesner’s Checklist of the Tiger Beetles of the World (1992; update in preparation). (b) Genus-level relationships according to the present study. Phylogeny of Megacephalina upland species, and we address the question whether or not these adaptations are monophyletic and whether the observed habitat preference is derived or ancestral. Materials and methods Taxon sampling The species included in this study are listed in Table I. Several species of the eight genera of Megacephalina (Figure 1a) were included in the analyses if available. For Phaeoxantha, Tetracha and Pseudotetracha,many species were selected in order to cover the biogeo- graphical range of these subgenera. For some Figure 2. Horn’s (1910) biogeographical scenario of the evolution species and subspecies, specimens from different of Megacephala sensu Horn with hypothetical intercontinental localities were collected. In total, 60 specimens of dispersal routes. Megacephalina representing nine Australian, three African/Eurasian, and 31 American species or sub- continental drift. They refer specifically to the break- species were analyzed. Three cicindelid species, up of the ancient Gondwana megacontinent which Neocollyris sp., Odontocheila confusa and Pentacomia