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Scarabaeidae: Aphodiinae: Eupariini) INSECTA MUNDI, Vol. 19, No. 1-2, March-June, 2005 55 The New World species of Ataenius Harold, 1867. V. Revision of the A. strigatus group (Scarabaeidae: Aphodiinae: Eupariini) Zdzislawa T. Stebnicka Institute of Systematics and Evolution of Animals Polish Academy of Sciences SB´awkowska 17, 31-016 Krakow, Poland [email protected] Paul K. Lago Biology Department University of Mississippi University, MS 38677 [email protected] Abstract. The strigatus group of the New World species of Ataenius Harold is revised. Seventeen species are recognized including two species described as new: Ataenius ecruensis sp. nov. from the United States and A. oaxacaensis sp. nov. from Mexico. Fifteen previously used names are considered valid, three new synonyms are proposed: A. liogaster Bates (= A. edwardsi Chapin syn. nov. = A. hoguei Cartwright and Spangler syn. nov.), A. wenzelii Horn (= A. rudellus Fall, syn. nov.). New state records are presented for A. spretulus (Haldeman) (Washington) and A. cognatus (LeConte) (Indiana, Missouri, and Mississippi). The taxa are diagnosed, keyed and illustrated; available biological information and distribution data are given. Key words: Scarabaeidae, Aphodiinae, Ataenius strigatus group, new species, taxonomy, New World. Introduction clusters of species that share similar combinations of the external character states, often correlated This is the fifth part of a revision of the New with structures of the male genitalia, but some World species of the genus Ataenius Harold (Steb- species within a group have male genitalia similar nicka 2001, 2002, 2003b, in press) and deals with to those from other species groups. Other problems the A. strigatus group of species widely distributed are connected with a north/south vicariance dem- in the middle and southern United States, Central onstrated by several species, and with extreme and South America. As now understood, the group ecological (ecophenotypic) and seasonal variation consists of seventeen species, two of which are here of many species having larger geographic ranges. described as new. Of the seventeen species consid- Variation is expressed either externally, often in ered, eight species are hitherto known only from several character states, and/or in characters of the the United States, three species occur in the USA male genitalia. Therefore, some qualifications and and Mexico, one species is apparently endemic to limitations should be emphasized in the context of the Lesser Antilles, and five species are distributed current knowledge of the taxonomy of Ataenius. in Mexico and South America, including one an- Every student who attempts to identify a specimen thropogenic species that has invaded the Oriental using this and subsequent group revisions should and Australian regions. consider that: 1/ the specimen may be of a species The intention of intrageneric grouping is to not previously recorded from a given area; 2/ it may facilitate identification of numerous species. As be a peripheral representative of a species of north- stressed by Stebnicka and Howden (1997), and then ern or southern cohesive range; 3/ it may be from a by Stebnicka (2001), the taxonomy and biogeogra- local population with somewhat different charac- phy of the Gondwanan genus Ataenius is very ters than any examined before; 4/ it may be of a difficult. A problem with grouping as we have done species forming a transition link between two is that some species do not fit easily into any groups; 5/ it may be of a species unknown to science. particular group, and could probably be placed in In the latter case, both external and internal char- monotypic “groups.” On the other hand, there are acters should be carefully analyzed. 56 Volume 19, No. 1-2, March-June, 2005, INSECTA MUNDI Collections Studied RMC Roy Morris Collection, Lakeland, Florida, USA Approximately 1738 specimens of Ataenius RTC Robert Turnbow Collection, Enterprise, strigatus group have been selected from the mate- Alabama, USA rial of Aphodiinae hitherto identified, including all RVNH Rijksmuseum van Natuurlijke Historie, the type specimens available to us. Leiden, Netherlands Material for this study was obtained from the SMNS Staatliche Museum für Naturkunde, Stut- following institutions and private collections. The tgart, Germany abbreviations listed below are used in all text cita- SMTD Staatliches Museum für Tierkunde, Dres- tions: den, Germany TMP Transvaal Museum, Pretoria, South Afri- ANSP Academy of Natural Sciences of Philadel- ca phia, Pennsylvania, USA USNM United States National Museum of Natu- BCP Balthasar’s Collection, National Museum, ral History, Washington DC, USA Prague, Czech Republic WSUC Washington State University Collection, CFC Carlos Flechtmann Collection, Brasilia Pullman, Washington, USA (Brazil) WBWC William B. Warner Collection, Chandler, CMN Canadian Museum of Nature, Ottawa, Arizona, USA Canada ZMHB Zoologisches Museum für Naturkunde der FCC F. Chalumeau Collection, Guadeloupe, Humboldt Universität, Berlin, Germany Lesser Antilles FMLT Fundacion Miguel Lillo, Tucumán, Ar- gentina Taxonomy FSCA Florida State Collection of Arthropods, The Ataenius strigatus group Gainesville, Florida, USA FVMC Fernando Vaz-de-Mello Collection, Viçosa, Diagnostic characters. Approximate length 3.4- Brazil 6.0 mm, body (Figs. 1, 2) elongate-oblong in most HAHC Henry and Anne Howden Collection, Ot- species, moderately convex, glabrous, shiny black tawa, Canada or piceous. Epipharynx and other mouthparts are HNHM Hungarian Natural History Museum, homogenous in shape or very weakly differentiated Budapest, Hungary at the species level and do not offer any useful ISEA Institute of Systematics and Evolution of diagnostic characters within this group. Head mod- Animals PAS, Krakow, Poland erate in size, not strongly elevated medially, cly- MCZC Museum of Comparative Zoology, Har- peal margin usually broadly rounded on each side vard University, Cambridge, Massachu- of median emargination, never dentate or angu- setts, USA late; clypeal surface in some species, or occasionally MHNG Muséum d’histoire naturelle, Geneva, in one of the sexes, weakly transversely wrinkled or Switzerland rugulose, middle of head minutely to finely punc- MNHN Museum National d’histoire naturelle, tured, vertex with scattered punctures or with Paris, France regular band of closer punctures. Pronotum trans- MMU Museum of Manchester, The University, verse, surface punctate, sides and base margined, Oxford, UK margin more or less deeply grooved, lateral margin MSNUP Museo di Storia Naturale, Universita di usually fringed with short to moderate, pale setae. Pisa, Calci, Italy Scutellum triangular or suboval. Elytra parallel- MSUC Mississippi State University Collection, sided or with lateral margins slightly convex, basal Starkville, Mississippi, USA bead fine, humeral denticles small to moderate; NRS Naturhistoriska Rijksmuseet, Stockholm, elytral striae distinctly impressed and punctate, Sweden intervals convex or flat, smooth, rarely eroded PKLC Paul K. Lago Collection, Biology Depart- posteriorly, lateral intervals usually not different. ment, University of Mississippi, Missis- Ventral surface shiny; abdominal sternites gla- sippi, USA brous, finely fluted along sutures, sometimes ster- PSC Paul Skelley Collection, Gainesville, Flor- nites 4-5 with coarser fluting, surface punctures ida, USA concentrated on sides. Profemur shiny, finely to INSECTA MUNDI, Vol. 19, No. 1-2, March-June, 2005 57 Figure 1. Ataenius oaxacaensis n. sp. Figure 2. Ataenius ecruensis n. sp. coarsely punctate, never scabrous; meso- and ing a number of character states such as the exter- metafemora shiny, metafemur in most species with nal sexual differences and same basic form of the incomplete posterior marginal line, this line rarely male genitalia. The differences include the unicol- lacking; meso- and metatibiae slender, subcylindri- ored and glabrous body in the strigatus group, the cal, apex with accessory spine, slender spurs and femora with posterior marginal lines in most spe- few setae; tarsi slender, basal tarsomere of meta- cies, and the more cylindrical meso- and metatibiae tarsus longer than upper tibial spur and usually furnished with accessory spine. Male genitalia with- shorter than following tarsomeres together. in the A. strigatus group are rather well differenti- External sexual differences apparent mostly in ated and paramere shape may be used to distin- the sculpture of the head, pronotum and metaster- guish the various species satisfactorily. num, in the shape of terminal spur of protibia, and in the length of abdominal sternites 5-6. Male Key to the species genitalia (Figs. 7-23) moderately sclerotized, of Ataenius strigatus group parameres usually as long as phallobase or longer, slender, narrowed apically; internal sac (Fig. 6) in 1. Elytral intervals in apical third or fourth carinate most species with symmetrical, serrate sclerites and weakly to strongly eroded on each side (Fig. and fine spicules. 21). ..................................................................... 2 1’ Elytral intervals convex or carinate apically, not eroded on each side (Fig. 22, 23) .................... 4 Affinities. The Ataenius strigatus group is most closely allied to the indigenous West Indian A. terminalis group of species (Stebnicka 2002) shar- 58 Volume 19, No. 1-2, March-June, 2005, INSECTA MUNDI 2(1) Body alutaceous, subopaque; clypeal surface very America, Venezuela ............................................ densely, roughly punctate;
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