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(Insecta: Coleoptera) in the Cis Taurus Species-Group

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(Insecta: Coleoptera) in the Cis Taurus Species-Group 71 (3): 181 – 210 20.12.2013 © Senckenberg Gesellschaft für Naturforschung, 2013. Review of the Neotropical Ciidae (Insecta: Coleoptera) in the Cis taurus species-group Ester H. Oliveira *, 1, 4, Cristiano Lopes-Andrade 2 & John F. Lawrence 3 1 Programa de Pós-Graduação em Entomologia, Departamento de Entomologia, Universidade Federal de Viçosa, 36570-900,Viçosa, Minas Gerais, Brazil — 2 Departamento de Biologia Animal, Universidade Federal de Viçosa, 36570-900, Viçosa, Minas Gerais, Brazil; Cristiano Lopes-Andrade [[email protected]] — 3 Australian National Insect Collection, CSIRO Ecosystem Sciences, GPO Box 1700, Canberra, ACT 2601, Australia; John F. Lawrence [[email protected]] — 4 Senckenberg Natural History Collections Dresden, Königsbrücker Landstrasse 159, 01109 Dresden, Germany; Ester H. Oliveira* [[email protected]] — * Corresponding author Accepted 26.xi.2013. Published online at www.senckenberg.de/arthropod-systematics on 13.xii.2013. Abstract The described Neotropical species of Cis Latreille, 1796 belonging to the taurus group are revised here. We designate lectotypes for the following names: Malacocis bahiensis Pic, 1916, Macrocis bison Reitter, 1878, Macrocis diabolicus Reitter, 1878, Macrocis grandicornis Pic, 1917, Macrocis rufescens Pic, 1922, Macrocis testaceimembris Pic, 1916, Macrocis testaceus Pic, 1916 and Trichapus pubescens Friedenreich, 1881. We transfer T. pubescens to Cis and include it in the taurus species-group, synonymize Trichapus Friedenreich, 1881 with Cis and transfer T. glaber Friedenreich, 1881 to Porculus Lawrence, 1987. We include C. longipilis Pic, 1930 in the taurus group and propose Macrocis bison as a junior synonym of C. diabolicus (Reitter, 1878). We redescribe eight Neotropical species: C. bahiensis (Pic, 1916), C. diabolicus (Reitter, 1878), C. grandicornis (Pic, 1917), C. kawanabei Lopes-Andrade, 2002, C. pubescens (Friedenreich, 1881) new combination, C. rufescens (Pic, 1922), C. setifer (Gorham, 1883) and C. testaceimembris (Pic, 1916). We also provide information on their geographic distribution and host fungi. This work solves the major taxonomic problems with the described taurus group species and also provides detailed morphological information on them, including that of male abdominal terminalia, allowing accurate identifica- tion of these species. Key words Polyphaga, Tenebrionoidea, Ciini, male abdominal terminalia, minute tree-fungus beetles. 1. Introduction Ciidae is a diverse and cosmopolitan family of fungivo- Cis Latreille, 1796, is the most speciose ciid genus rous beetles with 43 genera and more than 650 described and comprises approximately 370 species. It is an assem- species (LAWRENCE & LOPES-ANDRADE 2008, 2010; AN- blage of 25 species-groups, which are used today as taxo- TUNES-CARVALHO et al. 2012). These small beetles (ca. nomic tools to better deal with such a diversified taxon 1 – 7 mm body length) live, feed and reproduce inside (LAWRENCE 1971; LOPES-ANDRADE 2008a). However, not basidiomes of Polyporales and Hymenochaetales fungi all species of the genus are organized in groups. The few (PAVIOR-SMITH 1960; EPPS & ARNOLD 2010; LAWRENCE proposed subgenera of Cis are currently not used by au- & LOPES-ANDRADE 2010), building tunnels and feeding thors (e.g. LAWRENCE 1971; KAWANABE 1997; LOPES-AN- on the sterile support tissue of the fungus (LAWRENCE & DRADE 2008a, 2010; LOPES-ANDRADE et al. 2009) and the LOPES-ANDRADE 2010), leaving it just to disperse. whole genus deserves a careful revision. The genus Cis ISSN 1863-7221 (print) | eISSN 1864-8312 (online) 181 Oliveira et al.: Cis taurus species-group is defined by a set of characteristics common to several tinct units. These correspond to ten described species ciid lineages; however, these characteristics can be con- pre viously included in the taurus group, two described vergences between distinct lineages or plesiomorphies ciid species not assigned to the taurus group before, plus of Ciinae. The only phylogenetic analysis of the family a wealth of taurus group specimens that are more or less (BUDER et al. 2008) suggests that the genus Cis is poly- divergent from the described species. For part of the lat- phyletic. ter group, tentative species assignments are given below The morphology-based species-group of Cis named albeit they cannot yet be reliably assigned based on the taurus group currently comprises ten species, as fol- work done herein; other specimens that are too different lows: C. bahiensis (Pic, 1916), C. bison (Reitter, 1878), from previously described species likely represent new C. cornel li Lawrence, 1971, C. diabolicus (Reitter, 1878), species of the taurus group and are not further considered C. gran di cornis (Pic, 1917), C. kawanabei Lopes-An- herein. drade, 2002, C. rufescens (Pic, 1922), C. setifer (Gorham, Redescriptions of species are based on their respec- 1883), C. taurus (Reitter, 1878) and C. testacei mem bris tive lectotypes, designated in the present work, except (Pic, 1916). Most of them were described in the genus for C. diabolicus, whose redescription was based on the Ma cro cis Reitter, 1878, later synonymized with Cis by single paralectotype that we had in hands, and C. setifer, LAW RENCE (1971). Species in the taurus group can be se- whose redescription was based on a specimen compared parated from other Cis by the following features: body ro- to the type-series and here called plesiotype (sensu EVEN- bust, strongly convex; frontoclypeal ridge usually with a HUIS 2008). We had type specimens in hands, or examined pair of lateral horns in males and two short projections in them in the past, of most of the studied species, except females; prosternum in front of coxae strongly tumid and for C. longipilis, of which no type material could be lo- carinate at the longitudinal midline; prosternal process cated. In the cases of C. bahiensis, C. bison and C. kawa­ comparatively narrower at middle and enlarged at apex; nabei we have located only a single type specimen each, protibial apex with a row of spines and outer apical angle designated here as lectotypes. We cannot consider these produced into a stout tooth. While these features allow to as holotypes because it is unknown how many specimens recognize members of the taurus group, the monophyly were examined for their original descriptions. For C. cor­ of the group cannot be demonstrated at present due to the nelli and C. diabolicus we had in hands only paratypes very limited evidence on the phylogeny of the genus Cis and paralectotypes, respectively, but it shall be noted that and other Ciidae. The taurus group is possibly the most the former was described by one of the authors of the speciose species-group of Cis in the Neotropical region, present work (see LAWRENCE 1971) and the second was as based on our own extensive collections, and many ad- compared to the lectotype by him. For C. grandicornis, ditional species seem to exist. However, few species are C. rufescens, C. pubescens n. comb. and C. testaceimem­ described and, except for C. cornelli and C. taurus, the bris we had the lectotype and one or more paralectotypes. available information in the literature does not allow ac- We have designated the lectotype of C. taurus in a previ- curate identifications. ous work (OLIVEIRA & LOPES-ANDRADE 2013) and com- The name-giving species Cis taurus has been treated plementary information on the type-series is provided in a recent paper (OLIVEIRA & LOPES-ANDRADE 2013). The here (see 3.12.). aim of the present paper is to clarify the taxonomy of We dissected lectotypes and compared sclerites of the remaining taurus group with regard to the species so male abdominal terminalia to clarify doubts on species far formally described. This is mainly reached by erect- limits of very similar morphospecies, except for C. dia­ ing standardized redescriptions of species based on the bo li cus (paralectotype dissected), C. rufescens and C. se­ study of the type material whenever available (along ti fer (a specimen compared to the type was dissected). In with designations of lectotypes), plus further material if case a morphospecies was very similar to any described present. In addition we update information on the geo- species, including the ones transferred to the taurus group graphic distribution and host fungi of the species of the here, we dissected and compared male abdominal termi- taurus group. We also newly include two species in the nalia of specimens from each recognized population. taurus group: one Cis that has so far been unassigned to The terminology used here follows LOPES-ANDRADE any species-group of the genus and a species originally & LAWRENCE (2005, 2011). Frontoclypeal horns with a described in Trichapus Friedenreich, 1881, a genus here near-circular or thickly oval cross section are described synonymized with Cis. as “cylindrical” when the transversal diameter is approxi- mately constant up to the apex (process parallel-sided), and “conical” when this diameter decreases towards the apex (process tapering); horns with a strongly flat- tened cross section and a transversal diameter that is ap- 2. Materials and methods proximately constant up to the apex are categorized as “laminar parallel-sided”. What we call tegmen herein is not homologous to what is called tegmen in other cucu- jiform taxa, which is considered to be a modification of We examined more than one thousand mounted speci- the phallobase. The phallobase in all Ciidae is reduced, mens comprising approximately 80 morphologically dis - articulated
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