The Neotropical Subgenera and Species of the Pantropical Genus Anaulacus MacLeay (sensu novo) (Coleoptera: Carabidae: Masoreini): A Taxonomic Revision, with Notes about Way of Life, Evolution, and Geographical History Author(s): George E. Ball and Danny Shpeley Reviewed work(s): Source: Transactions of the American Entomological Society (1890-), Vol. 128, No. 2/3 (Jun. - Sep., 2002), pp. 265-343 Published by: American Entomological Society Stable URL: http://www.jstor.org/stable/25078782 . Accessed: 21/11/2012 15:45 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Entomological Society is collaborating with JSTOR to digitize, preserve and extend access to Transactions of the American Entomological Society (1890-). http://www.jstor.org This content downloaded by the authorized user from 192.168.82.205 on Wed, 21 Nov 2012 15:45:06 PM All use subject to JSTOR Terms and Conditions Transactions of the American Entomological Society 128(2+3): 265-343, 2002 The Neotropical Subgenera and Species of the Pantropical Genus Anaulacus MacLeay (sensu novo) (Cole?ptera: Carabidae: Masoreini): a Taxonomic Revision, with Notes about Way of Life, Evolution, and Geographical History George E. Ball and Danny Shpeley Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, CANADA ABSTRACT are Classified as genera of the tribe Masoreini the pantropical Anaulacus MacLeay, 1825, sensu novo, and three Eastern Hemisphere (Old World) genus-group taxa: Masoreus Dejean, 1821; Leuropus Andrewes, 1947; and as are Atlantomasoreus Mateu, 1984. Classified subgenera of Anaulacus the taxa: following previously described genus-group Anaulacus (sensu stricto), new new status; Aephnidius W. S. MacLeay, 1825, combination; Caphora Schmidt G?bel, 1846, new combination; Microus Chaudoir, 1876, new combination; new Macracanthus Chaudoir, 1846 (= Ophryognathus Chaudoir, 1876, junior new synonym), new combination; and Odontomasoreus Darlington, 1968, combination. a Primarily taxonomic review of the Neotropical elements of Anaulacus a a MacLeay, this paper includes classification, key to subgenera and species, new are descriptions of taxa, re-rankings, and synonymy. In total, 15 species treated, seven of which are described as new. are The Neotropical taxa arranged in the following sequence, with type name new localities of new species in parentheses, following of the taxon. The subgenus Apostolus includes only its type species, A. batesi Chaudoir. The eight are species of subgenus Aephnidius MacLeay arranged in three species groups: piceolus group, with only A. piceolus Chaudoir; ampliusculus group, with four species?A. ampliusculus Chaudoir, A. bonariensis Chaudoir, A. ciliatus Mutchler, and A. humeralis, new species (Santa Maria de Dota, Provincia San Jos?, Costa ? Rica); and exiguus group, with three species A. exiguus, new species (15 km. NE Puerto Maldonado, 12?33'S, 69?03,W, Departamento Madre de Dios, Peru), A. whiteheadi, new species (San Quintin, 16?24,N, 69?03,W, State of Chiapas, same M?xico); and A. ashei, new species (type locality as for A. exiguus, above). in Three fossil specimens in amber, from the island of Hispaniola, the Domini can are one Republic, assumed to be conspecific with another, the species to which they belong being included in subgenus Aephnidius, but treated as incertae sedis in relation to the exant members of that subgenus. The six species are in as of subgenus Macracanthus Chaudoir, arranged three species groups, follows: sericatus group, with three species?A. sericatus Chaudoir, A. thoracicus, new species (Porto Alegre, State of Rio Grande do Sul, Brazil), and A. erwini, new species (Cocha Shinguito, Rio Samiria, Departamento Loreto, Peru); the new mcclevei species group, including only A. mcclevei, species (17 km SW Moctezuma, State of Sonora, M?xico); and the A. tuberculatus group, including ? two species A. pittieri Gahan, and A. tuberculatus Chaudoir (including A. t. This content downloaded by the authorized user from 192.168.82.205 on Wed, 21 Nov 2012 15:45:06 PM All use subject to JSTOR Terms and Conditions 266 GENUS ANAULACUS tuberculatus and A. t. brasilianus van Emden, new junior synonym). Generally geophilous, most of the species live at altitudes between sea level a and 500 m., in variety of situations, from tropical evergreen forest, though some areas. live in semi-tropical forests, to dry, open Adults of many of the are species fly at night. Most of the species of subgenus Macracanthus associ some ated in way with ants of the genus Atta. The species A. (M.) sericatus is a facultative cavernicole, also. are The distribution patterns of subgenera and species fitted to a general to in model purporting explain such patterns, terms of plate tectonics, refuge theory, and climatic change during the Tertiary and Quaternary Periods. CONTENTS Introduction / 266 Taxonomic history / 267 Material, Methods, and Terms / 269 Material / 269 Methods / 270 Terms / 273 Taxonomic Treatment / 274 Tribe Masoreini / 274 to Key adults of Western Hemisphere genera of long-spurred lebiomorphs, and to the subgenera and species of Anaulacus (sensu novo) MacLeay / 274 Genus Anaulacus MacLeay, 1825 / 276 Subgenus Apostolus, new subgenus / 279 Subgenus Aephnidius MacLeay, 1825 / 381 Subgenus Macracanthus Chaudoir, 1846 / 313 Ecological Considerations / 326 Geographical history of Anaulacus / 326 Concluding Remarks / 327 Acknowledgments / 327 References Cited / 328 INTRODUCTION Small and dun-colored, Neotropical beetles of the genus Anaulacus, at first are glance, rather non-descript. They live in out-of-the-way places, such as caves (Vald?s, in press), nests of leaf cutter ants (van Emden, 1949), and in disturbed areas in are or tropical forests, where they encountered along well trodden trails, areas. in relatively open, sandy The senior author encountered them for the first near town time the Pacific coastal of Puerto Madera, Chiapas, running on sand beneath clumps of low-growing bromeliads. Pursuing small, fast-running beetles in such a situation is best avoided unless one is clad in a suit of armor with mailed as a gloves, defense against the spined leaf tips of the overhanging plants. Our in was a a interest the species of Anaulacus by-product of proposed review taxa was of the higher of the masoreine-cyclosomine carabid complex. This review not to a brought fruition because of the early demise of third party to the study: its actual instigator, Hans Reichardt, renowned, brilliant, highly productive, Brazil ian an coleopterist, who died tragically in 1976, in automobile accident. Following we Reichardt's death, continued to accumulate specimens, learning in the process we were that unable to identify many of those sent to us for determination. We to we our decided correct that situation. As proceeded with study, we became This content downloaded by the authorized user from 192.168.82.205 on Wed, 21 Nov 2012 15:45:06 PM All use subject to JSTOR Terms and Conditions G. E. BALL AND D. SHPELEY 267 aware a of limited array of interesting, taxonomically useful features, especially in cuticular microsculpture and in structure of the ovipositor of females. a are Plans for review of all of the species of Anaulacus being developed, but we are not in position to offer a general report on the group at this time. What we have to offer, which we dedicate to the memory of Hans Reichardt, will serve those interested in carabid biodiversity in the tropics of the Neotropical Region. Taxonomic history Because the names of genus-group taxa associated with Anaulacus require a names broader geographical context than the specific of the Neotropical taxa, the two groups are treated separately. Names of the genus-group. Proposed by MacLeay (1825:22-23), the Masoreus like genus Anaulacus contained, among others, two monobasic subgenera: Anaulacus (s. str.) and Aephnidius MacLeay. The nominotypical subgenus was not indicated as such, but its recognition is implicit with inclusion of subgenera in the genus. Schmidt-G?bel (1846: 88-90), in his pioneering study of Burmese Carabidae, treated Anaulacus and Aephnidius as if they were generically distinct, but included two in the latter taxon species (A. quadrimaculatus Schmidt-G?bel, and A.fasciatus was Schmidt-G?bel) that belonged in Anaulacus (s. str.). This because the type a specimens of those two species lacked mental tooth. In the original description of Anaulacus (s. str.), however, MacLeay stated that this taxon was characterized a by "mentum trilobum" (i.e., presence of tooth). As Schmidt-G?bel evidently had seen not the specimens of the type species, named by MacLeay as Anaulacus was unaware sericipennis, Schmidt-G?bel that MacLeay was in error about this a detail, and that A. sericipennis also lacked mental tooth. Chaudoir (1846: 539-541) described the genus Macracanthus, based on M. sericatus Chaudoir, the first species of the Anaulacus complex to be described from the Neotropical Region. Lacordaire (1854: 134; 265; and 308-309) recognized Masoreus, Macracanthus, as Anaulacus and Aephnidius not only generically distinct from each