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Neotropical Entomology ISSN: 1519-566X Journal Homepage 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, F 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 Alejandro Agrain, L a b Argentina. In this contribution, Bruch is re-described, de Entomología, Instituto Argentino de with addition of new morphological features of male internal sac, Investigaciones de las Zonas Áridas – IADIZA, Cicindis CCT-CONICET Mendoza, CC 507, 5500 female genital tract and elytral closure. New information on the Mendoza, Argentina; saroig@mendoza- species’ habitat and distribution is also provided. The phylogenetic conicet.gov.ar placement and relationships of within the family Carabidae are discussed on the basis of a cladistic analysis. Terminal taxa Edited by Roberto A Zucchi – ESALQ/USP included representatives of all subfamilies of Carabidae and supertribes of Carabinae, with a major samplingCicindis of those taxa Received 27 October 2010 and accepted 13 considered to be closely related to Cicindini by previous authors. The December 2010 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 xx(X): America xxx-xxx there© 2011 are Sociedade also Pangean Entomológica taxa do BrasilNebriini, Notiokasini, Opisthiini, and Notiophilini. Later,1 Morphology and Phylogeny of Cicindis Roig-Juñent et al Erwin (1985) hypothesized that Cicindini were closely History, Smithsonian, Washington D.C, USA (Terry Erwin) related to the tribe Notiophilini. Finally, Kavanaugh & (USNM). Erwin (1991) modified Kryzhanovsky’s classification Dissections were made following the techniques used scheme by elevating Cicinditae to the supertribe level in previous contributions of Carabidae (Roig-Juñent and placing it taxonomically between Nebriitae and 2000). Drawings were made with camera lucida adapted Elaphritae. to a stereomicroscope. Elytral structures were examined Kavanaugh (1998) presented a phylogenetic and photographed under a compound microscope. analysis includingOmophron both genera of the tribe Cicindini,Cicindela A tranverse section of the elytron was made using a and proposed thatOmus the tribe is the sister group of a clade microtome after inclusion of the elytron in paraplast Scaphinotuscomprising Latreille (Omophronini),Carabus solution. Scanning electron microscope pictures were L. (Cicindelini), Eschscholtz (Megacephalini), taken using a LEO 1450 VP microscope. TerminologyRoig-Juñent used & Latreille (Cychrini), and L. (Carabini). Cicchinofollows previous 2001 revisions (Deuve 1988, 1993, Liebherr Because representatives of Ozaeninae and other carabid & Will 1998, Roig-Juñent 1998, 2000, subfamilies such as Psydrinae were not included in Cladistic analyses). Kavanaugh’s analysis, the relationships of Cicindini with these taxa were not tested. Cicindis Liebherr & Will (1998) in a phylogenetic analysis using In our analyses we included representatives of all characters from female genitalia found as part of the supertribes of Carabinae and of the other carabid a polytomy with Migadopini, Amblytelina, Carabidae Cicindissubfamilies, especially those for which previous authors Limbata, and a monophyletic group conformed by proposed closer phylogenetic relationships with Siagonini, Cychrini, Pamborini, Carabini, and Cicindelini. Online Supplementary Material Liebherr & Will (1998) considered Cicindini in a middle For the cladistic analysis, a total of 50 adult level grade because it posses gonocoxal rami, but lacks morphological characters ( harpalidian type of abdomen. These authors also pointed 2) were scored for 27 species belonging to six subfamilies out the absence of accessory spermathecal gland. and 20 tribes. These species represent all the subfamilies Representatives of the tribe Cicindini have not proposed by Erwin & Sims (1984)i e 10 and1 20 of the 86 tribes. been included in other phylogeneticet al analyses using Characters in the text are referred to by number and their etmorphological al (e.g. Beutel 1998, Kavanaugh 1998), or states appear in superscript ( . ). molecular data (Maddison 1998, 1999, 2009, Balke A representative species of the family Trachypachidae, 2005). Cicindis horni Bruch, such as regarded as the sister taxon of Carabidae in previous The main objectives of this paper are to describe new works (Erwin 1985, Beutel 1998, Kavanaugh 1998, Roig- morphological features of Juñent 1998), was used to root the tree. male and female internal structures and the particular Morphological characters used in this analysis closure of the elytra, and to performCicindis a preliminary correspond to those proposed for the higher classification cladistic analysis based on adult morphology in order to of Carabidae in previous studies (Sloane 1923, Jeannel explore the phylogenetic placement of within the 1941, 1955, Bell 1967, Erwin 1985, Nichols 1985, Deuve family Carabidae. 1993, Baehr 1998, Liebherr & Will 1998, Roig-Juñent & Material and Methods CicchinoOnline 2001). Supplementary All characters Material were 3considered to be non-additive (unordered). The data matrix is presented C asData the analysis . horni The description of the morphological variability of . is based on examination of 25 males and 14 females.Online et al SupplementarySeveral specimens Material of 25 other1 carabid and trachypachid Phylogenetic analyses were performed using parsimony species were studied for the cladistic analysis (See software TNT (Goloboff 2003). The data set was ). Material for this study was analyzed using two procedures: (a) equally weighted borrowed from entomological collections of the following character analysis, and (b) implied weighting method institutions: Instituto Argentino de Investigaciones (Goloboff 1993), exploring the topologies obtained with de Zonas Áridas Mendoza, Argentina (Sergio Roig- different K (constant concavity) values from K = 1 to Juñent) (IAZA), Museo Argentino de Ciencias Naturales, K = 6. All analyses were conducted using a traditional “Bernardino Rivadavia,” Buenos Aires, Argentina (Arturo heuristic search on the base of Wagner trees, 100 random Roig- Alsina) (MACN), Museo de Ciencias Naturales addition sequences, followed by the tree-bisection de La Plata, La Plata, Argentina (Alberto Abramovich) reconnection (TBR) swapping algorithm, saving 100 (MLPA), University of Nebraska State Museum, USA trees per replicate. Branch robustness was assessed 2(Brett Ratcliffe) (UNSM), National Museum of NaturalNeotrop byEntomol standard xx(X): Bootstrappingxxx-xxx © 2011 Sociedade and Jackknifing Entomológica (removal do Brasil Roig-Juñent et al Morphology and Phylogeny of Cicindis probability = 36), with 500 replicates, searching among 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) areSystematic provided. remarks C horni The new material of . shows some interesting morphological
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