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New Morphological Aspects and Phylogenetic Considerations Of Neotropical Entomology ISSN: 1519-566X journal homepage: www.scielo.br/ne SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY New Morphological Aspects and Phylogenetic Considerations ofCicindis Bruch (Coleoptera: Carabidae: Cicindini) S Roig-Juñent, S Sallenave, FA Agrain Lab de Entomología, Instituto Argentino de Investigaciones de las Zonas Áridas – IADIZA, CCT-CONICET, Mendoza, Argentina Keywords Abstract Cladistics, Cicindis horni, morphology, phylogeny Cicindis Cicindis horni Correspondence Bruch is a monospecific genus of carabid beetles endemic to Federico A Agrain, Lab de Entomología, Argentina. In this contribution, Bruch is re-described, Instituto Argentino de Investigaciones de las with addition of new morphological features of male internal sac, Zonas Áridas – IADIZA, CCT-CONICET Mendoza, Cicindis CC 507, 5500 Mendoza, Argentina; saroig@ female genital tract and elytral closure. New information on the mendoza-conicet.gov.ar species’ habitat and distribution is also provided. The phylogenetic placement and relationships of within the family Carabidae Edited by Roberto A Zucchi – ESALQ/USP are discussed on the basis of a cladistic analysis. Terminal taxa included representatives of all subfamilies of Carabidae and Received 27 October 2010 and accepted 13 supertribes of Carabinae, with a major samplingCicindis of those taxa December 2010 considered to be closely related to Cicindini by previous authors. The phylogenetic analysisCicindis shows the basal position of in a clade that includes Ozaeninae, Omophronini, Scaritinae and Conjuncta. A close relationship of with Ozaenini + Metriini is supported by the particular closure of the procoxa and the ventral position of the oviduct with respect to the spermatheca. Introduction Cicindis such as Cnemalobini (Roig-Juñent 1993), Notiokasini (Kavanaugh & Nègre 1983), and Cicindini (Kavanaugh The monotypic genus Bruch constitutes one & Erwin 1991) which are related to holarctic or tropical of the several enigmatic carabid beetles endemic to carabids. Archaeocindisthe southern regions of South America. It is classified Beyond the particular pattern of distribution of the within the tribe Cicindini together with the genus tribe Cicindini, with one speciesCicindis in SouthArchaeocicindis America and Chaudoir. Southern southamerican carabid other in Iran, the unusual combination of morphological beetles (as other austral American insects groups) are characters exhibited by and phylogentically related with the carabid fauna from other had led taxonomists to propose appreciably different regions of the world. Southern southamerican carabids Cicindishypotheses about its relationship with other carabid such as zolines, migadopines, and broscines are related groups. When Bruch (1908) described the genus to groups occurring in other austral continents (Jeannel , he considered it to be related to Nebriini and 1938, 1967, Darlington Jr 1965,Systolosoma Roig-Juñent & Cicchino Omphronini. Other classification schemes considered 2001). Other membersPycnochila of the southern South America this genus to be a unique taxon within the tribe Cicindini, fauna such as trachypachids ( Solier) and related to Ozaenini and Metriini (Bänninger 1925, Bruch omines (the genus Motschulsky ofet theal tribe 1927, Kryzhanovsky 1976, Reichardt 1977). Erwin & Megacephalini) are relictual lineages related to groups Sims (1984) classified Cicindini within the supertribe also occurring in North America (Roig-Juñent 2008). Nebriitae, subfamily Carabinae, along with the tribes NeotropIn southern Entomol South 40(3): America 331-344 there© 2011 are Sociedade also Pangean Entomológica taxa do BrasilNebriini, Notiokasini, Opisthiini, and Notiophilini. Later,331 Morphology and Phylogeny of Cicindis Roig-Juñent et al Erwin (1985) hypothesized that Cicindini were closely Washington D.C, USA (Terry Erwin) (USNM). related to the tribe Notiophilini. Finally, Kavanaugh & Dissections were made following the techniques used Erwin (1991) modified Kryzhanovsky’s classification in previous contributions of Carabidae (Roig-Juñent scheme by elevating Cicinditae to the supertribe level 2000). Drawings were made with camera lucida adapted and placing it taxonomically between Nebriitae and to a stereomicroscope. Elytral structures were examined Elaphritae. and photographed under a compound microscope. Kavanaugh (1998) presented a phylogenetic A tranverse section of the elytron was made using a analysis includingOmophron both genera of the tribe Cicindini,Cicindela microtome after inclusion of the elytron in paraplast and proposed thatOmus the tribe is the sister group of a clade solution. Scanning electron microscope pictures were Scaphinotuscomprising Latreille (Omophronini),Carabus taken using a LEO 1450 VP microscope. Terminology used L. (Cicindelini), Eschscholtz (Megacephalini), follows previous revisions (Deuve 1988, 1993, Liebherr Latreille (Cychrini), and L. (Carabini). & Will 1998, Roig-Juñent 1998, 2000, Roig-Juñent & Because representatives of Ozaeninae and other carabid CicchinoCladistic 2001). analyses subfamilies such as Psydrinae were not included in Kavanaugh’s analysis, the relationships of Cicindini with these taxa were not tested. Cicindis In our analyses we included representatives of all Liebherr & Will (1998) in a phylogenetic analysis using the supertribes of Carabinae and of the other carabid characters from female genitalia found as part of Cicindissubfamilies, especially those for which previous authors a polytomy with Migadopini, Amblytelina, Carabidae proposed closer phylogenetic relationships with Limbata, and a monophyletic group conformed by . Online Supplementary Material Siagonini, Cychrini, Pamborini, Carabini, and Cicindelini. For the cladistic analysis, a total of 50 adult Liebherr & Will (1998) considered Cicindini in a middle morphological characters ( level grade because it posses gonocoxal rami, but lacks 2) were scored for 27 species belonging to six subfamilies harpalidian type of abdomen. These authors also pointed and 20 tribes. These species represent all the subfamilies out the absence of accessory spermathecal gland. proposed by Erwin & Sims (1984)i e 10 and1 20 of the 86 tribes. Representatives of the tribe Cicindini have not Characters in the text are referred to by number and their been included in other phylogeneticet al analyses using states appear in superscript ( . ). etmorphological al (e.g. Beutel 1998, Kavanaugh 1998), or A representative species of the family Trachypachidae, molecular data (Maddison 1998, 1999, 2009, Balke regarded as the sister taxon of Carabidae in previous 2005). Cicindis horni Bruch, such as works (Erwin 1985, Beutel 1998, Kavanaugh 1998, Roig- The main objectives of this paper are to describe new Juñent 1998), was used to root the tree. morphological features of Morphological characters used in this analysis male and female internal structures and the particular correspond to those proposed for the higher classification closure of the elytra, and to performCicindis a preliminary of Carabidae in previous studies (Sloane 1923, Jeannel cladistic analysis based on adult morphology in order to 1941, 1955, Bell 1967, Erwin 1985, Nichols 1985, Deuve explore the phylogenetic placement of within the 1993, Baehr 1998, Liebherr & Will 1998, Roig-Juñent & family Carabidae. Cicchino 2001). All characters were considered to be Material and Methods Online Supplementary Material 3 non-additive (unordered). The data matrix is presented asData the analysis . C horni The description of the morphological variability of . et al is based on examination of 25 males and 14 females.Online Phylogenetic analyses were performed using parsimony SupplementarySeveral specimens Material of 25 other1 carabid and trachypachid software TNT (Goloboff 2003). The data set was species were studied for the cladistic analysis (See analyzed using two procedures: (a) equally weighted ). Material for this study was character analysis, and (b) implied weighting method borrowed from entomological collections of the following (Goloboff 1993), exploring the topologies obtained with institutions: Instituto Argentino de Investigaciones de different K (constant concavity) values from K = 1 to Zonas Áridas Mendoza, Argentina (Sergio Roig-Juñent) K = 6. All analyses were conducted using a traditional (IADIZA), Museo Argentino de Ciencias Naturales, heuristic search on the base of Wagner trees, 100 random “Bernardino Rivadavia,” Buenos Aires, Argentina (Arturo addition sequences, followed by the tree-bisection Roig- Alsina) (MACN), Museo de Ciencias Naturales de La reconnection (TBR) swapping algorithm, saving 100 Plata, La Plata, Argentina (Alberto Abramovich) (MLPA), trees per replicate. Branch robustness was assessed University of Nebraska State Museum, USA (Brett Ratcliffe) by standard Bootstrapping and Jackknifing (removal 332(UNSM), National Museum of Natural History, Smithsonian,Neotrop probability Entomol 40(3): = 36),331-344 with © 2011500 Sociedadereplicates, Entomológica searching doamong Brasil Roig-Juñent et al Morphology and Phylogeny of Cicindis trees with traditional search for the equally weighted analysis. Bremer support and symmetric resampling (change probability = 33) were used as support values for implied weighting analyses since neither of these two measures is distorted by weight. All support numbers are given as relative values. Redescription Characters not described in Kavanaugh & Erwin (1991) Systematicare provided. remarks C horni The new material of . shows some interesting morphological
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