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
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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.
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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
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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 other, but placed them in three widely separated groups: Masoreus, in "Tribu XIII, L?biides"; Macracanthus, in "Tribu XXVIII, Cratoc?rides"; and Aephnidius and Anaulacus, in "Tribu XXXI, Pseudo-f?ronides". The numbering of tribes gives an indication of how different from one another these genera seemed to Lacordaire. Schaum (1860: 179) included in Masoreus Dejean, 1821, the species described previously in Anaulacus, Aephnidius, and Macracanthus, the three taxa noted above, thus declar ing implicitly the latter three names to be junior synonyms. In his recognition and first revision of the masoreine assemblage (Masoreidae), Chaudoir (1876:15) followed Schaum taxonomically (evidently with reservation, as noted by Bates [1883: 174]), but in form of the text, he treated each of these as were groups if they generically distinct. Chaudoir also described the monospe cific Neotropical genus Ophryognathus. taxa in Subsequent authors treated these various ways. The following, with on their studies based the fauna (or parts thereof) of the Eastern Hemisphere, or ranked explicitly, implicitly, Masoreus, Aephnidius and Anaulacus as genera: Bedel (1904:224-225); P?ringuey (1896:542);Winkler (1924:194);Andrewes (1930: 11-12; and 21-22); van Emden (1942:47, larvae); Jeannel (1949:861); Jedlicka (1963: 284); Darlington (1968: 76); Moore et al (1987: 781); and Machado (1992: 459). Basilewsky (1986: 254) declined to offer judgment about the generic distinctness o? Anaulacus and Aephnidius, but asserted that the South African species included in Anaulacus by P?ringuey (1908: 289; 1926: 611) actually belonged to Aephnidius. Lorenz (1998: 427) separated Masoreus from Aephnidius and Anaulacus, but in as a cluded the latter pair subgenera in single genus, for which he used the name Aephnidius.
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In the Neotropical Region, from where Anaulacus (s. str.) is not known, Bates as (1883:174) listed Aephnidius, Anaulacus and Macracanthus junior synonyms of even two names were Masoreus, though he believed that the first published in 1825, three years before publication of Masoreus (Dejean, 1828: 536), and thus would have priority. Evidently, like Chaudoir, Bates believed that these taxa eventually as so would be recognized generically distinct, and it seemed hardly necessary to pursue this nomenclatural matter further. In fact, however, the name Masoreus had so been validly published by Dejean in 1821 (p. 15; cf. Silfverberg, 1983:116), there a was no nomenclatural problem with treating Masoreus as senior synonym. Darlington (1934: 113) included Aephnidius and Macracanthus in the same genus, Masoreus, as did Blackwelder (1944: 52). Both of these authors recognized as Ophryognathus Chaudoir, 1876 generically distinct. Reichardt (1977:430) recog as nized Aephnidius and Macracanthus generically distinct, but included in Macracanthus, as a subgenus, Ophryognathus. Erwin and Sims (1984: 394) treated as as on Aephnidius and Macracanthus generically distinct, but their paper focused the West Indies, which is beyond the known geographical range of Ophryognathus, they did not have occasion tomention that taxon. Vald?s (in press) in his treatment of masoreine immature forms, recognized Macracanthus, Anaulacus, and Masoreus as one generically distinct from another. Van Emden (1949: 889) treated only Ophryognathus, synonymizing with it the generic name Leptotomus Gahan (1894: 8-9). Gahan, whose knowledge of carabid classification seems to have been quite limited, included Leptotomus in the was bradybaenine Harpalini. This assignment accepted by both Csiki (1932:1187) and Blackwelder (1944: 48), neither of whom/evidently, had seen Gahan's type material.
Names of the species group. Discussion of this topic is confined to the masoreine, and putatively masoreine, species known from the Neotropical Re gion. Chaudoir (1846: 541) described Macracanthus sericatus, the first masoreine known from the Neotropical Region. Chevrolat (1863:189) described from Cuban material Macracanthus brevicillus. In 1876, Chaudoir described the following taxa: inMasoreus (Aephnidius), M. batest, p. 20, M. ampliusculus and M. bonariensis, p. 21, and M. piceolus, p. 22; inMasoreus (Macracanthus), M. unicolor, p. 23; in the genus Ophryognathus, described as new, Chaudoir included on p. 26 the single species O. tuberculatus. Schaufuss (1879: 552) described Masoreus ridiculus from St. Thomas, new Virgin Islands. Gahan (1894:9) included in his genus, Leptotomus, the species L. pittieri. Mutchler (1934:130) described from West Indian material Masoreus (Aephnidius) ciliatus. Van Emden (1949: 889), on the basis of generic synonymy, transferred L. pittieri Gahan to the genus Ophryognathus. Erwin and Sims (1984:430) transferred M. ridiculus Schaufuss to the tachyine genus Polyderis Motschulsky. Summary. First, the genus Anaulacus, with its satellites noted above, has not a been the subject of careful general review since Chaudoir's monograph, pub lished in 1876. Second, disagreements about grouping and ranking of these taxa course have been few and not very serious. Third, with two exceptions, the of development of knowledge of Neotropical masoreines has been straightforward, a new van featuring sequence of descriptions of species. The first exception was Emden's discovery that Leptotomus pittieri and Ophryognathus tuberculatus were names. congeneric, resulting in the synonymy of these two generic The second was exception the discovery by Erwin and Sims that Masoreus ridiculus was in a reality, tachyine, and their reassignment of this taxon to the genus Polyderis. A conclusion that might be drawn from this review is that the Neotropical was Anaulacus assemblage of masoreines in good form taxonomically, and that more was a little to be done. But such conclusion would be premature, as ismade evident by the information summarized in Table 1, with details provided in the following pages.
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MATERIAL, METHODS, AND TERMS Material on This study is based examination of approximately 700 specimens of Masoreini, representing all known genera and most known species. For the we seen Neotropical species of Anaulacus, have 477 specimens. Some of the material was available in the Strickland Museum, Department of Biological Sciences, University of Alberta (UASM). Additional material was or borrowed from, deposited in, the following institutions and private collections, noted in the text by the associated codens. Names of owners or curators are included, in parentheses. ? AMNH Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York, U.S.A. 10024 (L. H. Herman) ? BMNH Department of Entomology, British Museum (Natural History), Cromwell Road, London, England SW7 5BD (M. J. D. Brendell, S. J. Hine) ? CASC Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California, U.S.A. 94118 (D. H. Kavanaugh) ? CMNC Canadian Museum of Nature, Entomology, P.O. Box 3443, Station D, Ottawa, Ontario, Canada KIP 6P4 (R. S. Anderson, F. G?nier) ? CMNH Section of Entomology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania, U.S.A. 15213 (R. L. Davidson) ? CNCI Canadian National Collection, Agriculture and Agri-Food Canada, K. W. Neatby Building, Ottawa, Ontario, Canada K1A 0C6 (Y. Bousquet) ? CUIC Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York, U.S.A. 14850 (J. K. Liebherr, K. W. Will) ? DEIC Deutsches Entomologisches Institut, D-16202 Eberswalde, Germany (H. J.Muller) ? FMNH Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, U.S.A. (M. Thayer, A. Newton, Jr.) ? FSCA Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture, P.O. Box 147100, Gainesville, Florida, U.S.A. 32614 (R. E. Woodruff, M. C. Thomas) ? HPSC Henry P. Stockwell Collection, same address as USNM, below ? IEMM Instituto de Ecolog?a, A. O, Departamento de Ecolog?a y Comportamiento Animal, Apartado Postal 63, Xalapa 91000, Veracruz, M?xico (E. D. Montes de Oca) ? von IHVC Instituto Alexander Humboldt, Villa de Leyva, Colombia (F. Fernandez). ? INBIO Instituto Nacional de Bioversidad, Apto. 22-3100. Santo Domingo de Heredia, 3100, Heredia, Costa Rica (A. Solis) ? JEWC J. E. Wappes, 179 Fall Creek, Bulverde Texas, U.S.A., 78163 ? MCZC Department of Entomology, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. 02138 (P. D. Perkins, B. D. Farrell) ? MNHP Entomologie, Mus?um National d'Histoire Naturelle, 45 Rue Buff on, Paris, 75005, France (H. Perrin, T. Deuve) ? MZSP Museu de Zoolog?a da Universidade de S?o Paulo, Biblioteca, 7172, 01.051 S?o Paulo, Brazil (C. Costa, U. R. Martins). ? NHMB Entomology Department, Naturhistorisches Museum, Agustinergasse 2, 4001 Basel, Switzerland (M. Brancucci) ? OSUC Department of Entomology, Ohio State University, 1735 Neil Av enue, Columbus, Ohio, U.S.A. 43210 (C. A. Triplehorn) ? QBUM Museu Nacional, Quinta da Boa Vista, S?o Cristovao, 20.942 Rio De Janeiro, Brazil (M. A. Monn?)
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? RHTC Robert H. Turnbow, Directorate of Engineering and Housing, Build ing 1404, Fort Rucker, Alabama, U.S.A. 36362-5137 ? RMNH National Museum of Natural History, P. O. Box 9517, 2311 RA., Leiden, The Netherlands (J. van Toi) ? SEAN Sevicio Entomol?gico Aut?nomo, Museo Entomol?gico, SEA, A.P. 527, Le?n, Nicaragua (J.-M. Maes) ? SEMC Snow Entomological Museum, The University of Kansas, Lawrence, Kansas, U.S.A. 66045-2119 (J. S. Ashe, R. E. Brooks) ? SPCC Stewart B. Peck Collection, Department of Biology, Carleton Univer sity, Ottawa, Ontario K1S 5B6, Canada ? UCD The Bohart Museum of Entomology, University of California, Davis, California 95616, U.S.A. (L. S. Kimsey) ? UNAM Colecci?n Entomolog?a, Instituto Biolog?a, Universidad Nacional Aut?noma de M?xico, Apartado Postal 70133, 04510 M?xico D. F. (M. and C. Santiago Zaragosa) ? UNCB Museo de Historia Natural, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apto. 7495, Santa F? de Bogot?, Co lombia (E. Camero) ? USNM Department of Entomology, United States National Museum of Natural History, Smithsonian Institution, Washington, D. C, U.S.A. 20560 (T. L. Erwin, W. Steiner) ? voor ZMAN Instituut Taxonomische Zoologie, Zoologisch Museum, van Universiteit Amsterdam, Plantage Middenlaan 64,1018 DH Amsterdam, The Netherlands (J. P. Duffels)
Methods
Taxonomic concepts, principles, criteria for ranking, and general working methods were the same as those described previously (Ball 1975,1978; Allen and Ball 1980). Measurements.? Measurements were made with an ocular micrometer in a
Wild M5 stereoscopic microscope, at 12X, 25X, and 50X. Measurements of external are: body parts and abbreviations used for them in the text - Length of head (HL) linear distance from base of left mandible to posterior - margin of left compound eye. Width of head (HW) maximum distance across head, - including eyes. Length of pronotum (PL) linear distance from anterior to posterior - margin, measured along the midline. Maximum width of pronotum (PWM) greatest - linear transverse distance. Width at base (posterior margin) of pronotum (PWB) linear transverse distance from posteriolateral angle to posteriolateral angle. - Length of elytra (EL) linear distance from basal ridge to apex of longer elytron (if the pair of elytra is asymmetrical), measured along the suture. Standardized Body as an sum Length (SBL), used index of overall size, is the of HL, PL, and EL (Table or or 2). Values for various ratios (more less diagnostic for species groups species) were computed, using the measurements above (Tables 3-5). Because of the we general paucity of material and thus small sample sizes, regard the numerical data as illustrative rather than definitive.
To express quantitatively proportions of the median lobe of the male genitalia, were three measurements made, using left lateral aspect of Figs. 10 and 18: Length - a of median lobe (MLL) measured in straight line from basal to apical margin. - a Length ofpreapical portion (PAL) measured in straight line from apical margin of area - periostial to apical margin. Length of ostial membrane (OML) measured in a straight line from basal to apical margin of the membrane. These measurements were combined in three ratios (Table 6). were Preparation of material.? Dissections made by using standard techniques. Genitalia and other small structures were preserved in glycerine in microvials,
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were pinned beneath the specimens from which they removed. Larger structures were and those that were gold-coated for study with the SEM glued to cards were pinned beneath the specimens from which they removed. were a Photographs of isolated structures taken with Cambridge S-250 Scan a ning Electron Microscope and JEOL JSM 6301 FXV field emission SEM. Line were a camera a drawings of selected body parts prepared by using lucida on Wild W5 stereoscopic microscope. Plates were prepared by using Adobe Photoshop 4.0.1.
Descriptions.? To reduce repetition, character states of lower ranking taxa are recorded in the descriptions of higher-ranking taxa not repeated in the taxa. descriptions of the included lower-ranking Thus, the complete description of a a can species must be assembled from its taxonomic placement. Such description be obtained by reading the descriptions and diagnoses of the sequence of higher ranking taxa in which the lower-ranking taxon is placed. a Habitat and activity.? For each species, section thus entitled summarizes the label data available, with "habitat" indicating primarily forest type, altitude, and any special features about way of life. "Activity" indicates months of the year were during which the specimens collected. Phylogenetic relationships.? A general analysis was not attempted, nor are we phylogenetic relationships postulated for all taxa. For many species, felt an assessment sufficiently confident to offer of their relationships, because of on overall similarity plus features considered apotypic based generalized outgroup comparison. are Specific ranking.? Species clusters of morphologically similar adults, distinguished from other clusters by discontinuity in one or more structural features. We expect that the discontinuity between morphologically similar clus more or ters will be reflected less clearly in features of other life stages. Further, we we as are expect that the clusters recognize distinct reproductively isolated from each other. Expressed in the fashion of Hennig (1966: 19-20), the tokogenetic we relations that previously connected these clusters have been interrupted. Thus, accept the (now) classical "biological" species definition, as formulated and over defended the years by Mayr (1942). See also Darlington (1980: 125-126). are on Supraspecific ranking.? Clusters of species of Anaulacus evident, based features postulated to be synapotypic. To designate these putative monophyletic are: clusters, the categories used species group and subgenus. We have adopted a a broad generic concept to make this categorical level useful to wide variety of so biologists rather than restricting it, that its value is limited to carabid specialists (Lindroth, 1969: XVII). on Label data.? For type material, the information each label is reproduced as exactly as is possible using ordinary type. Information on each label is enclosed in quotation marks; as well, a semicolon marks the end of a label. A slash mark (/) indicates the end of each line of text. Enclosed in square brackets is information or about color of label paper (other than white) printing (other than black), form of the label (other than rectangular), and coden for collection in which type material is housed. we For other material, report in full the locality data, date of collection and collectors. Label data about altitude and dates of collection are reported in summary fashion under "Habitat". Terms for structural features.? Most of the terms used to designate details of structures are in or are found textbooks of general entomology, used by coleopter ists, generally. Other words, used to designate particular structures or parts are not in us thereof, general use, though they have been used by in previous publications. We provide information about these words here, as well as names that have been changed for certain structural features.
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Microsculpture.? A "sculpticell" is the space on the surface of the cuticle enclosed by adjacent microlines of the integumental system of Microsculpture (Allen and Ball 1980:485-486). Microsculpture of the elytra varies from mesh pattern isodiametric, to transverse with sculpticells flat (Figs. 2C, G), to slightly convex to to longitudinal, with sculpticells distinctly keeled (Figs. 2A 2E, 2B, 2F) in rows markedly longitudinal, sculpticells form of long parallel (Figs. 2J, 2L, 2N, 20). Length of a sculpticell refers to its longer dimension, with width referring to the shorter dimension. are Chaetotaxy.? This term refers to the fixed setae, which the long, evidently on tactile, commonly encountered setae carabids: dorsal labral (6); clypeal (1 pair); on supraorbital (2 pairs); stipital (1 to several each stipes); submental; mental; glossal; palpigeral; pronotal; elytral parascutellar, discal, and umbilicate (or lateral); coxal, trochanteral, femoral, and tarsomeral; abdominal sternal ambula near tory (sterna IV, V, VI); and abdominal sternal terminal (sternum VII, posterior margin). use Body parts.? The term "segment" is restricted for to those body parts that reflect embryonic somites; thus, somite-like portions of the abdomen are referred to as segments. Abdominal segments are designated by Roman numerals corre sponding to their respective somites. The first complete sternum is III, and the last one normally exposed is VII. For numbering the genital somites, we follow Bus (1976). are Portions of appendages designated by the suffix "-mere", the prefix on depending the appendage in question: antenno-, palpo-, tarso-, etc. Mandibles.? Shpeley and Ball (2001: 9-21) characterized the mandibles of the lebiine sub tribe Pericalina, and illustrated the major features with SEM figures. We use here the same system (Figs. 3A-E). Labium.? The masoreine labium is standard for Carabidae. For the combined we use glossae and paraglossae, the standard term ligula. The central sclerotized, apically setigerous structure is the glossal sclerite (Fig. 4A-C). on are as Elytra.? Shallow longitudinal grooves the dorsal surface designated as we striae, rather than interneurs (Erwin 1974:3-5). Although appreciate Erwin's arguments (and accepted them in previous publications [Shpeley and Ball, 2001: we 21]), bow here to the weight of general usage. Male tarsal vestiture.? Only one type of adhesive vestiture on the ventral surface of the fore tarsi is exhibited by masoreine males: biseriate squamo-setae see (for illustrations, Shpeley and Ball, 2001: 230, Figs. 12A-C). ? Male genitalia: The surface of the median lobe treated by convention as dorsal is really the ventral surface, and vice versa (Deuve 1993:88). We have chosen to were as: remain with the conventional usage. Median lobes classified anopic, with the ostium dorso-medial; left pleuropic-anopic, with the ostium laterad, but more toward the dorsal surface; and left pleuropic-catopic, with ostium more toward the ventral surface.
The median lobe (Figs. 10A-T, and 18A-L), illustrated in left lateral and ventral aspects, with base toward the bottom of the page, exhibits interspecific differences in are seen as are form. These differences readily overall patterns ('Gestalt') but not so with of easily described, except notation differences in size and shape of the distal area. To two are provide the basis for verbal description, principal regions distinguished, the shaft (s) and basal bulb (bb). The latter is the swollen area set at an to angle the ventrally curved shaft, surrounding the basal opening (bo). areas are a Three of the shaft recognized: more distal periostial area (po), subtend the ostial ing membrane (om), which surrounds the ostium (oo), marking the place of sac a more area egress of the internal during copulation; proximal middle (ma) a extended from the base of the ostium to the basal bulb; and preapex or preapical area (pa) extended distally from the apex of the ostial membrane to the apex.
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area Proportions of the preapical and ostial membrane (or essentially, periostial are area) indicated by the ratios PAL/ MLL, OML/ MLL, and PAL/OML (Table a 6). The mean in Table 6B indicates central tendency for variation in each ratio, a and provides rough basis for ready comparison among the supraspecific taxa of Anaulacus. ? Ovipositor. For naming the sclerites of the adephagan ovipositor, we have elected to follow the system used by Liebherr and Will (1998; see figures), rather than persisting with the older system used by Shpeley and Ball (2001: 21). = = Comparing the systems, laterotergite valvifer; gonocoxite 1 stylomere 1; and = gonocoxite 2 stylomere 2. are are For the gonocoxites, the surfaces that ventral in the infolded position lateral when the ovipositor, is extended; thus such surfaces are designated as lateral, and the other surfaces are designated accordingly. Gonocoxite 2 is much or or simplified, evidently by loss: without basal lobe ensiform setae, furrow peg setae (Shpeley and Ball, 2001:33, Fig. 13C, fp); and nematiform setae absent or very short (Fig. HD, ns). However, the ventral membrane, which seems to be derived from the basal membrane of the subapical setose organ, ranges in size from small to extending half the length of gonocoxite 2 (Figs. 12A-F, vm).
Geographical terms An important landmark that extends north-south in South America is the are Andean mountain system. Areas to the west of the Andes designated trans Andean; those to the east as eis-Andean. The term Middle America refers to M?xico plus the republics of Central America, collectively. Mesoamerica, for some authors, is the equivalent of Middle America. For anthropologists, it has a more restricted meaning, and for this reason, Mesoamerica is not used here. are Geographical parts of Middle America recognized: M?xico is the northern most and largest part; Central America includes the small republics of Guatemala, Belize, Honduras, El Salvador, Nicaragua, Cost Rica and Panam?. West Indies includes the Bahamas, Greater Antilles, Virgin Islands, Cayman Islands, and Lesser Antilles.
Biogeographical terms we use Following Frank and McCoy (1990), "indigenous" (synonyms autoch thonous, native) for a taxon that achieved its current taxonomic status in the area where it is living. If an indigeneous taxon occurs nowhere else, it is referred to as "precinctive" (from Latin, meaning to gird, or encircle). a A taxon that achieved its taxonomic status elsewhere than in given area where it occurs now, is "adventive" in that area. An adventive taxon is either a "introduced" ifmoved to given area by man; or it is "immigrant" if it was not introduced. are In Zoogeographie discussion, the following terms used: Nuclear Middle America, which includes northern Nicaragua, the Central American republics to the north thereof (Honduras, El Salvador, Guatemala and Belize), and M?xico; and Lower Central America, which includes the southern part of Nicaragua, Costa Rica, a a and Panam?, region which, until the Pliocene Epoch, consisted of series of isolated volcanic islands. Nuclear Middle America includes two parts: Nuclear Central America, from Honduras and El Salvador north to the Isthmus of Tehuantepec; and M?xico, north of that isthmus, for which we have no other designation. Phylogenetic terms or use In place of sister group sister taxon, we "adelphotaxon" (Ax 1987:36), for reasons given by that author. In place of "plesiomorphic" and "apomorphic", we use on a "plesiotypic" and "apotypic", the basis that the latter pair have more general connotation than the former pair (Tuomikoski 1967).
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TAXONOMIC TREATMENT Tribe Masoreini
Mazor?ides [vernacular; misspelling] Chaudoir, 1871: 111. Masoreidae Chaudoir, 1876: 29. Masoreinae Bates, 1883: 174 Masoreini Erwin and Sims, 1984: 394.? Ball and Bousquet, 2001: 109. Masoreina Reichardt, 1977: 430. Anaulacini Blackwelder, 1944: 52.
Type genus.? Masoreus Dejean, 1821 on Notes about synonymy.?The focus of the above list is usage of suprageneric name names for masoreines in relation to the Neotropical fauna. The first generic was to be used for designating masoreines Masoreus Dejean, 1821. So, the correct name stem for a suprageneric for this group (however itmay be ranked) must be Masore-.
Csiki (1932:1287), followed by Blackwelder (1944:52) proposed Anaulacini as name a replacement name, evidently because he believed that the correct stem had to be that of the first-described genus in the group. Having dated the Dejean name, incorrectly, as 1828, it seemed that Masoreus had been proposed three years after was the name Anaulacus MacLeay, 1825. In fact, the date of publication of Masoreus was no 1821, and thus it prior to publication of Anaulacus. However, such is of name one consequence, because the valid suprageneric is the first to be used correctly (International Code of Zoological Nomenclature, 1999: 65 [Article 64]). a a as Recognition, etc.?The tribe Masoreini is member of complex referred to to the "long-spurred lebiomorphs", in allusion the hypertrophied middle and hind tibial spurs that adults of these taxa exhibit. Being monogeneric in the Neotropical Region, the distinguishing features of the tribe, its geographical as distribution, and other aspects are the same those of its genus, Anaulacus. See the on key, and the treatment of Anaulacus, for details. A broader perspective the a Masoreini is provided by Ball and Bousquet (2001:109), in discussion of the tribe we Cyclosomini. Here we note only that follow Basilewsky (1984:527) in exclusion an of the somoplatine genera from the Masoreini, action he did not follow subsequently (Basilewsky, 1986).
Key to Adults of Western Hemisphere genera of long-spurred lebiomorphs, and to the subgenera and species of Anaulacus (sensu novo) MacLeay
1 Labial mentum toothed. Tibial spurs of middle and hind legs fimbriate, margins serrate. Mandible basally with dorsolateral and ventrolateral or margins in line, expl?nate, with lateroventral margin distinctly laterad laterodorsal ' margin.2 1 Labial mentum edentate. Tibial spurs with lateral margins smooth, not serrate. Mandible (Fig. 3E) basally with laterodorsal margin extended laterad lateroventral margin. Tribe Masoreini, Anaulacus MacLeay .3 narrow 2(1) Head posteriad eyes constricted in form of neck. Tarsal claws prominently serrate. Mandible basally expl?nate, lateroventral margin extended distinctly laterad laterodorsal margin. .Tribe Lebiini, Nemotarsus LeConte 2' Head posteriad eyes not constricted in form of narrow neck. Tarsal or claws smooth, with very short serrulations. Mandible not expl?nate, laterodorsal and lateroventral margins in line. .Tribe Cyclosomini, Tetr?gonoderus Dejean
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3(1') Elytral microsculpture with mesh pattern markedly elongate more as rows (sculpticells than 6 times long wide), sculpticells in and sharply keeled (cf. Figs. 2N-0)... subgenus Macracanthus Chaudoir .4 or 3' Elytral microsculpture with mesh pattern nearly isodiametric, as transverse, or slightly elongate (sculpticells 2 to 3 times long wide) .9
4(3) Mandibles with dorsolateral margin beaded, slightly up-turned. Elytral or not hum?rus angulate subangulate, rounded. Pronotum (cf. Figs. 17A and 19A-B) with anterior angles sharp. 5 4' Mandibles with dorsolateral margins rounded into lateral surface. Elytral hum?rus rounded, not angulate. Pronotum with anterior more 7 angles rounded. 5(4) Elytral hum?rus subangulate; basal ridge sinuate (cf. Fig. 5A). - Geographical range northwestern M?xico. new xxx .A. mcclevei, species, p. 5' Elytral hum?rus angulate; basal ridge narrow, straight, not sinuate - (Fig. 5B). Geographical range Belize, Middle America, to Argentina, South America. 6 or rows on 6(5') Elytron (cf. Fig. 19B) with two three of tubercles disc. . A. tuberculatus Chaudoir, p. xxx 6' Elytron (cf. Fig. 19A) without tubercles, smooth. .A. pittieri Gahan, p. xxx on 7(4') Pronotal microsculpture disc with mesh pattern nearly isodiametric, rather coarse.A. sericatus Chaudoir, p. xxx on 7' Pronotal microsculpture disc with mesh pattern distinctly transverse, shallow, or microlines not evident at 50X magnification. 8 areas 8(7') Pronotum bicolored, with disc piceous, lateral rufous; disc with microlines evident. Elytron bicolored, with hum?rus rufous, rest of new xxx surface piceous.A. thoracicus, species, p. 8' Pronotum and elytra concolorous, piceous. Pronotum smooth, microlines not evident at 50X magnification. new xxx .A. erwini, species, p. mm. 9(3') Size larger, SBL more than 5.5 Pronotum (cf. Fig. 1) with lateral more margins markedly rounded posteriorly, and posterior margin . . . new narrower Apostolus, subgenus. .A. batesi Chaudoir, p. xxx 9' Size smaller, SBL less than 4.0 mm. Pronotum (Figs. 8A-C) with lateral . . . margins less rounded posteriorly, posterior margin broader Subgenus Aephnidius MacLeay.10 10(9') Elytron with dorsal surface distinctly shiny, microsculpture mesh pattern in intervals 1-5 isodiametric to transverse, in intervals 6-8 diagonal. Pronotum rufotestaceous, distinctly paler than head and elytral disc. Size smaller, SBL less than 3.0 mm.11 10' Elytron with dorsal surface dull or with sericeous luster, mesh pattern narrow. slightly elongate, sculpticells Pronotum (at least disc) concolorous with head and elytral disc. Size larger, SBL more than 3.0 mm.13
11(10) Elytral microsculpture mesh pattern (Figs. 2D, H) of intervals 1-5 new xxx uniformly transverse.A. ashei, species, p. 11' Elytral mesh pattern (Figs. 2C, G) of intervals 1-2, 1-3, 1-4, or 1-5 isodiametric.12
12(11') Geographical range Middle America (Chiapas and Belize). Males with as median lobe in Figs. 10Q-R. .A. whiteheadi, new species, p. xxx
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as 12' Geographical range South America. Males with median lobe in Figs. 10O-P. A. exiguus, new species, p. xxx a 13(10') Umbilical punctures at lateral apical angle of elytron in "V" ( Figs 9E new xxx F).A. humeralis, species, p. an 13' Umbilical punctures at lateral apical angle of elytron in outward curve (Figs. 9A-D).14 14(13') Elytron with mesh pattern slightly but distinctly elongate, surface iridescent, sericeous.A. ampliusculus Chaudoir, p. xxx 14' Elytron with mesh pattern nearly isodiametric, surface dull. 15 on 15(14') Pronotum bicolored, rufo-piceous anteriorly, and lateral and xxx posterior slopes.A. bonariensis Chaudoir, p. to 15' Pronotum uniformly rufopiceous piceous. 16 sac 16(15') Antennae and legs testaceous. Internal of male genitalia with numerous xxx long spines basally.A. piceolus Chaudoir, p. 16' Antennomeres 3-11 darker than antennomeres 1-2. Legs bicolored, sac apices of tibiae darker. Internal of male genitalia without long xxx spines basally.A. ciliatus Mutchler, p.
Genus ANAULACUS MacLeay (sensu novo)
Anaulacus MacLeay, 1825: 22 Type species.?Anaulacus sericipennis MacLeay, 1825 (by subgeneric monotypy). names are Synonymy.? The of the Neotropical subgenera treated below junior are synonyms of Anaulacus (s. lot.), but since they being used, it seems unnecessary to record them in a formal synonymy. Recognition.? See the key, above. Description.? With features noted in the key, and character states below. Habitus. As in Figs. 1, 7A-B, 15,16, and 19A-B (Neotropical species, only). For see data about size (Standardized Body Length), and proportions, Tables 2A-6B. Color. Somber, body piceous to rufous, dorsal surface concolorous (except some or elytra of species) pronotum rufotestaceous, paler than head and elytral or or disc; elytra concolorous bicolored, with humeri apices paler than rest of surface; appendages piceous to testaceous. Microsculpture (Figs. 2A-0) and luster. Head capsule: Clypeus with mesh or pattern isodiametric to transverse, longitudinal, sculpticells short, slightly convex or to flat, microlines either distinct, surface subopaque, evanescent, surface shining. Frons with mesh pattern isodiametric medially, longitudinal laterally, sculpticells slightly convex, surface subopaque, or mesh pattern trans verse, sculpticells flat, and microlines finer, surface subiridescent. Vertex with mesh pattern isodiametric to longitudinal posteriorly, sculpticells slightly convex and or transverse microlines deeper, surface subopaque; generally, longitudinal posteriorly, with sculpticells flat and microlines finer, surface subiridescent. Ventral surface with mesh pattern transverse, sculpticells long, narrow, surface or iridescent. Labrum: Mesh pattern isodiametric slightly transverse, sculpticells large, convex, microlines clearly indicated, surface subopaque. Mandibles: Dorsal or surface shining, with without microsculpture; if present, mesh pattern longitu dinal, microlines shallow, surface shining, subiridescent. Pronotum: Disc with mesh pattern isodiametric, sculpticells slightly convex, microlines deep, surface subopaque, to transverse, sculpticells flat, microlines shallow, subiridescent. Lateral declivity with mesh pattern transverse, sculpticells flatter, surface shining to subiridescent. Scutellum: Mesh pattern isodiametric, sculpticells slightly convex, surface subopaque, to transverse, sculpticells flat, surface shining. Elytra: Mesh pattern isodiametric, sculpticells convex, surface opaque; or mesh pattern or or transverse, sculpticells flat, surface iridescent; longitudinal, slightly mark
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to edly so, sculpticells slightly convex, surface subiridescent markedly iridescent. Ventral surface of thorax and abdominal sterna with mesh pattern transverse on on or on (oblique proepisterna, longitudinal proepipleura), surface iridescent, prosternum medially, surface shining. Macrosculpture. Body surfaces essentially smooth, excluding elytral striation and tuberculation (for details, see below, under "Elytra"). row one Chaetotaxy. Labrum preapically with of six setae; clypeus with pair of setae; vertex with two pairs of supraorbital setae. Mouthparts: maxillary stipes er and palpif each with single lateral seta; labial submentum with pair of long setae; mentum with pair of short paramedial setae; glossal sclerite with four apical setae (Fig. 4B, gs); labial palpomere 1 (Fig. 4B, lpl) with single seta, labial palpomere 2 (Fig. 4B, lp2) with two setae preapically. Pronotum with two pairs of lateral setae, on on or posteriolateral setigerous punctures lateral bead, mediolateral puncture or close to bead. Prosternai intercoxal process with (all Western Hemisphere taxa) on without pair of setae posterior (apical) margin. Elytron with parascutellar seta, or with without pair of discal setae, and with 11-13 setae in lateral (umbilical) one one series in three groups: one medial seta, anterior, and posterior group, latter two with five or six setae, each. Legs (number of setae on each of front, middle and hind leg): coxae (1-4-2); trochanters (1-1-1); femora, dorsal (ca. 5-ca. 8 ca. 5); femora, ventral-posterior surface 0-0-2; ventral-anterior surface 0-ca. 3-ca. 3); setation of tarsomeres described below. Abdominal sterna V and VI each with pair sternum of paramedian ambulatory setae, VII with single pair of terminal setae (in both sexes) near posterior margin. near Vestiture. Antennae: antennomere 1 with several setae anteriorly base, and several preapically; antennomere 2 with single preapical seta; antennomeres 3-10 with ring of setae preapically; antennomere 4 with apical 2/3 setose; antennomeres 5-11 densely setose, seemingly pubescent. Male front tarsus: tarsomeres 1-3 each with two rows of squamo-setae. Middle tibia of males and females preapically with setal brush (Figs. 5E-F, sb), brush more extensive in males (Fig. 5E) than in females (Fig. 5F). Head: Dorsal surface slightly, evenly near convex; frontal impressions very shallow, punctiform fronto-clypeal suture. Eyes in size and convexity about average for geophilous Carabidae. Antennae: Length average, in backfolded position, antennomere 11 extended posteriad posterior margin of pronotum (Fig. 1); or shorter, with antennomere 11 not extended to posterior margin of pronotum. Antennomere 1 thicker than 2-11, subequal to length of eye; antennomere 2 short, shorter than 3, 3-10 subequal, 11 slightly longer. Antennomeres 5-10 elongate, each distinctly longer than wide more or (Fig. 1), less cylindrical, to quadrate and somewhat flattened (Figs. 19A B). or Mouthparts. Labrum transverse; anterior margin straight slightly concave to narrowly and deeply notched medially. Mandibles (Figs. 3A-E), dorsally with or narrow lateral margins broadly rounded (Fig. 3A, B), margined with raised area ridge; basal (b) ca.1/4 length of terebra (t); basal brush (bb) short; laterodorsal margin (Fig. 3E, 1dm) extended laterad ventrolateral margin; scrobe (s) not visible in dorsal aspect. Left mandible: dorsal aspect (Fig. 3A) occlusal margin with incisor (i) prominent; terebral ridge (tr) straight; terebral tooth (tt) small, blunt; retinaculum not evident dorsally; premolar tooth absent; molar tooth (m) broad; in ventral aspect (Fig. 3C) posterior occlusal groove (pog) prominent; retinacular ridge (rr) short; molar ridge complete, with occlusal and basal extensions (oemr, bemr, respectively); ventral groove (vg) moderately long. Right mandible (Figs. 3B, D) similar to left in overall size and proportions; incisor tooth (i) broader, less prominent; terebral ridge (tr) concave; anterior and posterior retinacular teeth (art, prt, respectively) evident, also premolar and molar teeth (pm, m, respectively); in ventral aspect, anterior and posterior occlusal grooves (aog, pog, respectively)
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evident, also retinacular ridge (rr) and premolar and molar ridges); ventral groove (vg) same as for left mandible. Maxillae (Ball, 1983:527, Fig. 9A) about average for Carabidae: lacinia with prominent tooth, without apical brush; galea with galeomeres about equal in length. Labium (Figs. 4A-C). Mentum (4A, m) edentate, lateral lobes (11) broad, distally, asymmetrically broadly rounded; epilobes broad, with slight dentiform projection (del) each side. Ligula with glossal sclerite (Fig. 4B, gsc) broadly keeled proximally, apical margin truncate, broad; paraglossae one (pg) membranous lobes, widely separated from another, lobes extended distinctly distad apical margin of glossal sclerite; palpus (lpl- 3) with palpomere 1 very short, 2 and 3 subequal, 3 fusiform, though apex narrowly subtruncate. Prothorax. Pronotum (Figs. 1, 8A-C, 17A-B, and 19A-B) transverse, distinctly wider than long, expl?nate, lateral margins extended laterad pleural sclerites; transverse, anterior margin moderately (Fig. 1) tomarkedly Figs. 19A-B) concave, convex more lateral margins curved, not sinuate; posterior margin slightly to markedly so and slightly lobate; anterior and posterior margins beaded laterally, more beaded or not medially; lateral margins narrowly to widely beaded through or out length, bead widened not in vicinity of posteriolateral setigerous punctures. convex Disc smooth or with pair of paramedian tubercles in posterior 1/3, slightly me medially, sloped ventrad laterally; median longitudinal impression shallow or dially, not extended to either anterior posterior margins; posteriolateral impres sions at most shallow, indistinct, in most taxa not evident. Proepipleuron joined aver to propleural sclerites at obtuse angle; propleural sclerites and prosternum age for Carabidae. Pterothorax. Average for harpaline Carabidae, without distinctive features. ca. Metepisternum with lateral margin 1.25 X longer than width at base. Elytra. (Figs. 1,5A-D, 7A-B, 9A-H,16, and 19A-B). Expl?nate, extended laterad pterothorax and abdominal sclerites. Form oblong, base distinctly broader than or pronotal base; hum?rus rounded (Fig. 1) angulate (Fig. 5,19A-B); apical margin or obliquely truncate, slightly sinuate not. Basal ridge (Figs. 5A-B, br) sinuate and or narrow or broad (Fig. 5A) straight and (Fig. 5B). Striae 1-8 shallowly impressed courses unimpressed, with their marked by sub-surface trabeculae; parascutellar stria inmost individuals unimpressed, evident in few individuals. Intervals flat or tuberculate (Fig. 19A). Posteriolateral umbilical setigerous punctures (Figs. 9A-H, upl-4) variously arranged. Hind wings. (Fig. 6A). Fully developed; venation relatively complete; oblongum cell (o) short, stalked; wedge cell not evident. Legs. Proportions about average for geophilous Carabidae, with front tibia distinctly widened distally; middle and hind tibiae terete to distinctly compressed and widened, width hind tibia apically less than half maximum width of hind more femur, to than half width of hind femur. Spines of posterior rows (Figs. 5E F) of middle and hind tibiae longer and thicker than average. Spurs long, inner to tarsomere 1. spur equal length of Tarsomeres of middle and hind legs slender, + length of tarsomere 1 equal to length of tarsomeres 2 3. Front tarsomeres of females shorter and broader, of males, tarsomeres 1-3 markedly broad, with apical margins emarginate; tarsomere 4 shorter and narrower; tarsomere 5 same as for middle and hind legs. Tarsomeres 1-4 each with pair of laterodorsal spines at apical margin, tarsomere 5 with pair of fine setae, proximad apex. Tarsomeres 1 4 with ventrolateral margins each with row of setae, tarsomere 5 with as many as on three finer setae each ventrolateral margin. Tarsomere 1with row of medio ventral spines. Claws serrate, serrations short, subject to wear, and barely evident inmany specimens. Abdominal sterna. Average for lebiomorph Carabidae. Male genitalia. Median lobe (Figs. 10A-T, and 18A-L) various in form, anopic sac or to left pleuropic-catopic; internal with without spines. For details, see
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subgeneric and species treatments, below. Female genitalia: Ovipositor. (Figs. 6B, 11A-J, and 12A-J). Gonocoxites 1 and 2 or (Fig. 12J, gel, gc2, respectively) asetose, gonocoxite 2 with two short nematiform setae (Fig. HD, ns). Gonocoxite 2 without basal lobe, shaft in form terete (ventral surface as wide as or narrower than lateral surface; cf. Fig. HA), or markedly compressed (Fig. 11F), tapered to bluntly rounded apex; ventral surface com pletely sclerotized except narrowly membranous in pit of subapical setose organ or more (Figs. 11B, D, vm), ventral surface extensively membranous (Figs. 11F, HI, 12B, 12E, 12H). Internal genitalia (Fig. 6B). Bursa copulatrix (be) short, broad; spermatheca (sp) rather thick throughout length, curved, not twisted, inserted common directly into bursa copulatrix, not into oviduct (co), spermathecal gland (spg) broad, spermathecal gland duct (spgd) narrow, sinuous. our we Morphological note.? During investigation of structural features, a coxa discovered seta on the posterior rim of the front (noted in the description seems a above, in the setal formula for legs) that to be constant and unique character state for adults of the tribe Masoreini.
Subgenera on we Based especially similarity in mandibular configuration, in Anaulacus, as include the following subgenera: the Neotropical precinctives, Apostolus, new subgenus and Macracanthus Chaudoir (including Ophryognathus Chaudoir); the Eastern Hemisphere precinctives, Anaulacus (s. str.), Microus Chaudoir, new new new status, Caphora Schmidt-G?bel, status, and Odontomasoreus Darlington, status; and the pantropical Aephnidius MacLeay. The South African species, a currently in Aephnidius (Basilewsky, 1986:253-256) comprise subgenus presently a undescribed, which will be treated in subsequent publication. Geographical distribution.? The range of Anaulacus (s. lot.) is pantropical, across extending from eastern Australia westward the Oriental Region to the Afrotropical Region to the Neotropical Region (Figs. 13,14, and 20). In the Eastern Hemisphere the range of Anaulacus extends northward to the southern reaches of the Palaearctic Region in the Japanese archipelago (Habu, 1967: 57). In the Neotropical Region, Anaulacus ranges northward in western M?xico, to within about 100 km of the Mexican- U.S. border.
Chorological affinities.? In the Eastern Hemisphere, the geographical range of or Anaulacus is overlapped partially completely by all of the other known masoreine genera. For the subgenera and species groups in the Neotropical Region, range is so as to overlap extensive render unnecessary any discussion of chorological affinities at these taxonomic levels.
Phylogenetic relationships.? Although we are not able at this time to offer a we closely reasoned statement about this topic, believe that the masoreine genera as + recognized presently may be related follows: Atlantomasoreus ((Masoreus Leuropus) Anaulacus).
new Subgenus APOSTOLUS, subgenus
Type species.? Aephnidius batest Chaudoir, 1876 (here designated). name: name a Subgeneric The subgeneric is Latin noun, meaning messenger, in to allusion the female characteristics of the single included species giving evidence a (hence delivering message) about the origin of the ventroapical membrane of 2 as gonocoxite of the ovipositor. Just the apostolic St. Paul's evocative, inspira a tional epistles delivered message of hope to the Gentile followers of the Christian so us religion, does the primitive ovipositor of A. batest give hope of discovering, eventually, the evolutionary patterns in Anaulacus.
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more Recognition.? A combination of relatively large size (SBL than 5.5 mm.) and reticulate mesh pattern of microsculpture distinguishes adults of this mono taxon members of Anaulacus. Additional specific from the other Neotropical features distinguishing adults of this subgenus from those of Macracanthus and more narrower Ophryognathus by the constricted pronotal proportions: base (Fig. 1; cf. Figs. 7A-B, 8A-C, 15, and 18A-B; Table 3B), and width and length less are divergent (thus values higher for ratio PL/PWM, Table 4B). Males distinctive in the hooked apical portion of the left pleuropic-catopic median lobe (Figs. 10A - B), with relatively long preapical portion and relatively long periostial part a (Table 6). Females are distinguished by features of ovipositor, gonocoxite 2: very are small ventroapical membrane and nematiform setae (Figs. 11A-D), which very short.
Description and other information.? Because this subgenus is monospecific, its are same character states, geographical range and phylogenetic relationships the as those of its single species, A. batest. See below, for details.
Anaulacus (Apostolus) batesi Chaudoir Figs. 1,10A-B, 11A-D, and 13
au Masoreus (Aephnidius) batesi Chaudoir, 1876: 20. TYPE MATERIAL: two thentic Chaudoir specimens, in Oberth?r-Chaudoir Collection (MNHP), to an associated with following label originally pinned the bottom of insect box: "Batesi/ Chaudoir/ Amazone/ Ega Bates" [handwritten]. LECTO TYPE male (here selected), labelled: "Ex Musaeo/ Chaudoir [red print]. same as PARALECTOTYPE (here selected) labelled lectotype.
Notes about type material.? Associated with the types is a female from the Bates was collection, also collected at "Ega". It probably obtained by Oberth?r with purchase of the Bates collection, subsequent to publication of the description of A. batesi.
Type locality.? Tefe (= Ega), State of Amazonas, Brazil. Description.? With recognition features noted above, and character states of or as genus Anaulacus, restricted amplified follows. Habitus. As in Fig. 1. For data see about size (Standardized Body Length), and proportions, Tables 2A-6B. Color. or Somber, body piceous; elytra concolorous bicolored, with apices paler than rest of surface; appendages testaceous. Microsculpture (Figs. 2A-0) and luster. Head capsule: Frons with mesh pattern isodiametric medially, longitudinal laterally, sculpticells slightly convex, surface subopaque. Vertex with mesh pattern isodia convex metric to longitudinal posteriorly, sculpticells slightly and microlines or deeper, surface subopaque. Labrum: Mesh pattern isodiametric slightly trans verse, sculpticells large, convex, microlines clearly indicated, surface subopaque. Mandibles: Dorsal surface shining, mesh pattern longitudinal, microlines shal low. Pronotum: Disc with mesh pattern transverse, sculpticells flat, surface shining. Lateral declivity with mesh pattern transverse, sculpticells flatter, surface subiridescent. Scutellum: Mesh pattern isodiametric, slightly convex, surface subopaque. Elytra: Mesh pattern slightly elongate, sculpticells narrow, convex, in surface opaque. Head. Eyes size and convexity about average for geophilous in Carabidae. Antennae. Length average, backfolded position, antennomere 11 extended posteriad posterior margin of pronotum (Fig. 1). Antennomeres 5-10 elongate, each distinctly longer than wide (Fig. 1). Mouthparts. Labrum with or concave. mar anterior margin straight shallowly Mandibles with dorsolateral gins rounded, not ridged. Prothorax. Pronotum (cf. Fig. 1) anterior margin moder ately (Fig. 1) concave, lateral margins curved, not sinuate; posterior margin slightly lobate; anterior and posterior margins beaded laterally, not beaded
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medially. Disc smooth, not tuberculate. elytra (Fig. 1). Hum?rus rounded (Fig. 1); apical margin obliquely truncate, slightly sinuate. Basal ridge (cf. Figs. 5A-B, br) sinuate and broad. Striae shallowly impressed; parascutellar stria shallow, inter or vals flat, not tuberculate. Discal setigerous punctures present (2 exx.) absent (13 exx.). Posteriolateral umbilical setigerous punctures aligned as in Figs. 9C-D. Legs. Middle and hind tibiae terete; width of hind tibia apically less than half maximum width of hind femur. Male genitalia. Median lobe (Figs. 10A-B) left pleuropic catopic; shaft (Fig. 10A) curved evenly dorsally and ventrally, with shaft-basal bulb juncture ventrally rather abrupt; in ventral aspect (Fig. 10B), shaft markedly area constricted prebasally; periostial (po) narrowed distally in form of smooth nar juncture with preapex (pa), latter in left lateral aspect sloped gradually to area rowly obtuse apex; preapical in ventral aspect flattened, broad, projected to sac right as rather blunt hook. Internal without spines. Ovipositor. Gonocoxite 2 (Fig. HA) with pair of short nematiform setae (Fig. HD, ns); terete in form, in lateral aspect (Fig. HA) parallel-sided for most of length; in ventral aspect (Fig. 11B) widened from base distally; ventral surface narrower than lateral surface; apex broadly rounded, with distinct dorso-apical ridge. an ever Habitat and activity.? This species is inhabitant of lowland tropical over an - m. green forest, altitudinal range of approximately 0 500 Collections of adults at mercury vapor light indicate nocturnal flight activity. Known adult over activity extends the period May to August and October to November. Geographical distribution.? (Fig. 13). The range of this species extends from the Amazon Basin of central Brazil, eastern Bolivia and Peru northward in Middle America to southern Panam?. on Phylogenetic relationships.? Based structure of ovipositor, gonocoxite 2, this species (and subgenus Apostolus) is postulated to be the adelphotaxon of the remaining species of Anaulacus (s. tat.). Other primitive features are the nematiform setae in the ventroapical membrane of gonocoxite 2, and in few specimens, discal setigerous punctures in elytral interval 3. we Material examined.? In addition to the types, have seen 15 specimens of this species, from the following localities. BOLIVIA. Santa Cruz. Female, Province Sara, J. Steinbach (FMNH). Ichilo Province, Buena Vista, 400 m, 3.X.1994, R. Ward (CMNH): 2 males; female, mercury vapor light. BRAZIL. Matto Grosso. Female, 1886, P. Germain (MNHP). Tapirape Ind. Vill., confluence of R. Tapirape and R. Araguaia, B. Malkin (FMNH): female, 11-20.X.1960; female, 20-30.XI.1960. PANAMA. Canal Zone. Barro Colorado Island, Shannon Trail 9.5, leaf litter, E.O. Willis (MCZC): male, 1.VIII.1961; female, 16.XI.1962. 2 males, female, Barro Colorado Island, T.L. Erwin (USNM). PERU. Loreto. Male, Iquitos, leaf litter in forest, 90 m, 8.V.1992, J.Danoff-Berg (SEMC). Madre de Dios. Male, 15 km ne Pto. Maldonado, pitfall trap, 200 m, 7.VII.1989, J.Ashe, R. Leschen (SEMC). Madre de Dios Dpto., 15 km ne Pto. Maldonado, Reserva Cuzco Amaz?nica, 12?33'S 69?03'W, m pitfall trap, 200 (SEMC): female, 4.VII.1989, J.S. Ashe, R. Leschen D. Silva; male, 28.VI.1989, J.S. Ashe, R.A. Leschen (SEMC).
Subgenus AEPHNIDIUS MacLeay
Aephnidius MacLeay, 1825: 23
Type species.? Aephnidius adelioides MacLeay, 1825: 23 Recognition.? See key to subgenera, above. Gonocoxite 2 of the Ovipositor. area more or (Figs. 11E-J, and 12A-F) is asetose, and has the ventroapical less extensively desclerotized, membranous. Male median lobes (Figs. 10C-T) vary collectively from left-pleuropic-anopic to left pleuropic, with the preapical part relatively short to long and the periostial part short to intermediate (Table 6). The sac or a internal is with without basal collar of spines.
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Description.? With recognition features noted above, and character states of or as genus Anaulacus, restricted amplified follows. Habitus. As in Figs. 7A-B. For data about size (Standardized Body Length), and see proportions, Tables 2A-6B. Color. Somber, body piceous to rufous, dorsal surface concolorous (except elytra of some species) or pronotum rufotestaceous, paler than head and elytral disc; elytra concolorous or bicolored, with humeri paler than rest of surface; appendages piceous to testaceous. Microsculpture (Figs. 2A-I) and luster. Head capsule: Clypeus with mesh convex pattern isodiametric to transverse, sculpticells short, slightly to flat, microlines distinct, surface subopaque. Frons with mesh pattern isodiametric medially, longitudinal laterally, sculpticells slightly convex, surface subopaque. to Vertex with mesh pattern isodiametric longitudinal posteriorly, sculpticells convex slightly and microlines deeper, surface subopaque. Labrum: Mesh pattern or isodiametric slightly transverse, sculpticells large, convex, microlines clearly indicated, surface subopaque. Mandibles: Dorsal surface shining, with mesh pattern longitudinal, microlines shallow. Pronotum: disc with mesh pattern isodiametric to transverse, sculpticells slightly convex, microlines deep, surface subopaque. Lateral declivity with mesh pattern transverse, sculpticells flatter, surface shining to subiridescent. Scutellum: Mesh pattern isodiametric, sculpticells or slightly convex, surface subopaque; mesh pattern transverse, sculpticells slightly convex, surface shining. Elytra: Mesh pattern isodiametric, sculpticells convex, surface opaque; or mesh pattern transverse, sculpticells flat, surface iridescent. Head. Antennae various: length average, inbackfolded position, antennomere or 11 extended posteriad posterior margin of pronotum (Fig. 1); shorter, antennomere 11 extended only to posterior margin of pronotum. Antennomeres more or 5-10 elongate, each distinctly longer than wide (cf. Fig. 1), less cylindrical. Mouthparts. Labrum with anterior margin straight. Mandibles (Figs. 3A-E). see Dorsolateral margins rounded, not ridged. For details, description of genus. Prothorax. Pronotum (Figs. 7A-B; 8A-C) with disc smooth, not tuberculate. Elytra (Figs. 1, 5A-C-D, 7A-B, and 9A-H). Hum?rus rounded; apical margin obliquely truncate, slightly sinuate. Basal ridge sinuate and broad (Fig. 5A). Striae shallowly impressed; parascutellar stria shallow. Intervals flat. Posteriolateral umbilical setigerous punctures (Figs. 9A-H, 1-4) variously arranged. Legs. Middle and hind tibiae terete, width of hind tibia apically less than half maximum width of hind femur.
Male genitalia. Median lobe (Figs. 10C-T) anopic to left pleuropic-catopic, various in form; preapical area in ventral aspect convex (not flattened), narrow to broad, apex evenly rounded, not projected to right. Internal sac with or without spines. For details, see species group and species treatments, below. Ovipositor. (Figs. 11E-J, and 12A-F). Gonocoxite 2 without nematiform setae. Gonocoxite 2: shaft in form terete (ventral surface as wide as or narrower than or lateral surface; cf. Fig. HA), markedly compressed (Fig. 11F), tapered to bluntly rounded apex; ventral surface extensively membranous (Figs. 11F, 111). Habitat.? See species treatments, below. Geographical distribution.? This subgenus occupies almost all of the known in range of the genus Anaulacus both the Eastern and Western Hemispheres. In the Western Hemisphere (Figs. 13-14), the range of subgenus Aephnidius extends: in eis-Andean South America, from northeastern Argentina northward to northern Colombia and Venezuela; inMiddle America, from Panama to the tropic of Cancer in eastern M?xico; and to the Greater Antilles and southern Bahamas, in the West Indies.
Phylogenetic relationships.? Not specified.
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Species groups
are The eight extant Neotropical species included in this subgenus arranged in three groups: the monospecific piceolus species group; the ampliusculus species group, with four species; and the exiguus species group, with three species.
The piceolus species group Recognition.? The principal features for recognition of this group are details a of the reproductive organs. Males exhibit collar of fine spines or spinules at the sac base of the internal (Fig. 10E). In proportions of the median lobe (Table 6B), the area preapical portion is relatively long, whereas the ostial is relatively short. are Females distinctive in the markedly compressed form of gonocoxite 2 (Fig. 11F).
Anaulacus (Aephnidius) piceolus Chaudoir Figs. 8A, 9A-B, 10C-E, 11E-G, and 13
Masoreus (Aephnidius) piceolus Chaudoir, 1876:22. TYPE MATERIAL: five of six specimens, in Oberth?r-Chaudoir Collection (MNHP), four unlabelled, to an associated with following label originally pinned the bottom of insect box: "piceolus/ Chaudoir/ Amazone/ Ega Bates" [handwritten]. LECTO TYPE male (here selected): unlabelled, second specimen in the series. Four PARALECTOTYPES, two males, unlabelled; female, labelled "Cayenne/ 1866"; female, labelled "Yucatan/ Pilate".?Bates, 1883: 175.
Notes about type material.? The type material includes representatives of two species, distinguished by differences in the Male genitalia: We have arbitrarily elected to associate the specific epithet "piceolus" with the species characterized sac. by spines in the male internal In the original description, Chaudoir recorded six specimens, apparently in same as the sequence specimens noted above: first three (males), unlabelled; then, one the "Cayenne" specimen, followed by from the "Yucatan" and then, followed an we was on by unlabelled male, which believe also collected by Pilate the Yucatan Peninsula of M?xico. Although four of the six specimens are not labelled, we are feel confident that all those noted in the original description. The first a unlabelled male is not member of A. piceolus, as shown by absence of spines from sac. the internal It belongs to the species Aephnidius ciliatus Mutchler. Accordingly, it is excluded from the type series of A. piceolus, with the second male being selected as lectotype. Type locality.? Tefe (= Ega), State of Amazonas, Brazil. Recognition.? See the key and distinguishing features for the piceolus species are group. Inmost respects, adults of A. piceolus much like those of the ampliusculus species group in size and form. size nor in Neither the ratios presented Tables 3A-5A distinguish between adults of A. piceolus and A. ciliatus. Small size (Table 2A) distinguishes this species A. A. from ampliusculus and bonariensis. Ratios distinguishing A. piceolus from A. ampliusculus are: PL/PWM (Table 4A) and HW/PWM (Table 5A). Ratios distin guishing A. piceolus from A. bonariensis are: PWB/PWM (Table 3A), PL/PWM (Table 4A), and HW/PWM (Table 5A). In values for the ratio PWB/PWM (Table 3A), A. piceolus overlaps only slightly with A. humeralis. or Description.? With character states of subgenus Aephnidius restricted as as amplified follows. Habitus. Approximately in Fig 7A. For data about size see (Standardized Body Length), and proportions, Tables 2A-6B. Color. Somber, body piceous; elytra concolorous; appendages piceous. Microsculpture (Figs. 2A I) and luster. Head capsule: Clypeus with mesh pattern isodiametric, sculpticells slightly convex, microlines distinct, surface subopaque. Frons with mesh pattern
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isodiametric medially, longitudinal laterally, sculpticells slightly convex, surface subopaque. Vertex with mesh pattern isodiametric to longitudinal posteriorly, convex sculpticells slightly and microlines deeper, surface subopaque. Labrum: Mesh pattern slightly transverse, sculpticells convex, microlines clearly indicated, surface subopaque. Pronotum: Disc with mesh pattern isodiametric, sculpticells slightly convex, microlines deep, surface subopaque. Scutellum: Mesh pattern isodiametric, sculpticells convex, surface subopaque. Elytra: Mesh pattern isodia metric, sculpticells convex, surface opaque. Mouthparts. Labrum with anterior or concave. as margin straight slightly Prothorax. Pronotum in Fig. 8A. Elytra. as Posteriolateral umbilical setigerous punctures arranged in Figs. 9A-B. Male genitalia. Median lobe (Figs. 10C-E) left pleuropic-anopic; basal bend gradual, area narrower area basal slightly than medial area; medial in left lateral aspect into (Fig. IOC) curved evenly periostial area, latter narrowed evenly into preapical to area, with narrowing continued preapically acute apex, especially marked in sac lateral aspect; preapical portion moderately long. Internal with basal collar of slender long spines (Fig. 10E, s). Ovipositor. Gonocoxite 2 with shaft markedly compressed (Fig. 11F). Habitat and This on activity.? species lives the ground, in dry tropical forest, and in as disturbed locations, such mango and banana plantations, at altitudes sea m. between level and 500 Collections of adults at UV and MV lights indicate seems nocturnal flight activity. Known adult activity to be year-round, specimens been in having collected every month except April and December. Geographical distribution.? (Fig. 13). The range of A. piceolus extends from eastern Bolivia northward in Middle America almost to the tropic of Cancer, in eastern M?xico.
Phylogenetic relationships.? Not specified. Material In to we seen examined.? addition the type material, have 73 speci mens of A. piceolus, as follows. BOLIVIA. Santa Cruz. Female, 5 km ese Warnes, Hotel Rio Selva, UV trap, 20.X.2000, M.C. Thomas (FSCA). COLOMBIA. Bolivar. Zambrano, Hacienda Monterrey, 9?45'N 79?45'W, 10 m, F. Fernandez (UNCB): male, berlese, 23.VL1993; female, pitfall trap, 8.VII.1993. Magdalena. Female, PNN Tayrona Nequanje, 11?17'41"N 74?06'15"W, 155 m, IX.1996, F. Escobar (UNCB). COSTA RICA. Guanacaste. Female, 18.7 km nw Bagaces, Hwy. 1, Rio Carana, 91 m, 4.VI.1973 (USNM). Est. Palo Verde, P.N. Palo Verde, 10 m, V. Chavarria (INBIO): 2 males, V.1992; male, female, 4-7.V.1991. Female, Est. Palo Verde, Ref. Nac. Fauna Silva R.L. Rodriquez, 10 m, V.1991, D. Acevedo (INBIO). San Jose. La Caja, Bahia San Jose, F. Nevermann (USNM): male, III.1929; female, 11.1932. GUATEMALA. Guatemala. Female, Guatemala City, 26.V.1964, R.E. Woodruff u-v (FSCA). HONDURAS. Cortes. Female, La Lima, light, 3. VI.1964, F.S. A.B. R.E. Blanton, Broce, Woodruff (FSCA). M?XICO. Chiapas. 7 males, 6 females, Puerto Madero, dry tropical forest, ca. sea level, 2.VIII.1972, G.E. Ball (UASM). Oaxaca. 2 males, Chila, 31.V.1987 (UNAM): E. Barrera; F. Arias. Male, n Rte. 190,22.3 km Zanatepec, 145 m, 28. VIII.1967, G.E. Ball, T.L. Erwin, R.E. Leech (UASM). Female, Rte. 190, bet. Niltepec and Zanatapec, Bell & Van Horn (RTBC). E.A. Schwarz Tamaulipas. Male, Tampico, (USNM). Veracruz. Male, Apazapan, 19?19'45"N 96?44'25"W, mango banana plantation, pitfall trap, 330 m, 11.2000, E. Montes de Oca, Q. Santiago (IEMM). Male, ex Veracruz, ex orchids, Laredo, Texas (USNM). NICARAGUA. Leon. Leon: male, 12.11.1989, J.M. Maes (SEAN); male, B. Garcete 4 u-v III, 1991, (CMNH); males, 5 females, light, 10.11.1993, H. Pogatshnik u-v (CMNH); male, light, 25.111.1993, Maes, Lecocq, Cantamessa (CMNH). Fe Finca at male, Hernandez, u-v, 8.XI.1989, J.M. Maes (SEAN). Managua. 2 females, de Laguna Xiloa, blacklight, 10.1.1992, E. vanden Berghe (CMNH). PANAMA. Canal Zone. 3 males, 3 females, Juan Mina, 30.VIII.1923, R.C. Shannon (USNM). Madden 9?13'N at Male, female, Dam, 79?37'W, under stones, light, 30.VI.1974,
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Erwin, Whitehead (USNM). Code. 5 maies, 4 females, 13 km sw Penonome at Rio Colee Auxiliar, 26.V.1983, W.E. Steiner (USNM). Chiriqui. Female, 3 km n David, 31.V.1983, W.E. Steiner (USNM). 3 males, female, Tol?, Champion (BMNH). Los Santos. Female, 6.3 km n Pocri, 7?41'N 80?10'W, 45 m, 21.VI.1973, T.L. & L.J. Erwin w mv (USNM). Panam?. Male, 2.5 km Ipeti, and bl, 11.V.1996, R. Turnbow s (RHTC). VENEZUELA. Gu?rico. Male, 12 km Calabozo Biol. Sta., u-v light, 6 12.11.1969, P. & P. Spangler (USNM).
The ampliusculus species group
Recognition and comparisons.? A combination of intermediate size (SBL be tween 3.0 and 5.0 mm.), head, pronotum and elytral disc concolorous, and reticulate microsculpture of the elytra distinguish the adults of this species group from others of the subgenus Aephnidius. Males are distinguished by absence of spines from the internal sac of the genitalia. Other features are details of the male as median lobe (Table 6, and Figs. 10F-N), follows: shaft with preapical portion short, and ostial area of intermediate length. Description.?With recognition features of subgenus Aephnidius, and character or as states of genus Anaulacus, restricted amplified follows. Habitus. As in Figs. 7A-B. For data about size (Standardized Body Length), and proportions, see Tables 2A-6B. Color. Somber, body piceous, with head, pronotum and elytral disc same as or concolorous. Pronotum laterally disc, paler; proepipleura paler than or dorsal surface, elytra dorsally concolorous bicolored, with humeri paler than or same as rest of surface; epipleura paler, dark rufous, color dorsal surface. Appendages piceous to testaceous. Microsculpture (Figs. 2A-I) and luster. Head capsule: Clypeus with mesh pattern slightly elongate, sculpticells slightly convex to keeled, microlines distinct, surface subopaque. Labrum: Mesh pattern isodia metric or slightly transverse. Pronotum: Disc with mesh pattern isodiametric, sculpticells slightly convex, microlines deep, surface subopaque. Lateral declivity with mesh pattern transverse, sculpticells flatter, surface shining. Scutellum: Mesh pattern isodiametric, sculpticells slightly convex, surface subopaque. Elytra: or convex or Mesh pattern isodiametric slightly elongate, sculpticells keeled, surface opaque. Head. Antennae: length average, in backfolded position, antennomere 11 extended posteriad posterior margin of pronotum. Antennomeres more or 5-10 elongate, each distinctly longer than wide (cf. Fig. 1), less cylindrical. Mouthparts. Labrum with anterior margin straight. Prothorax. Pronotum as in Figs. 7A-B and 8A-C. elytra (5A, 7A-B, and 9C-F). Posteriolateral umbilical setigerous punctures (Figs. 9C-F, upl-4) variously arranged. Male genitalia. Median lobe (Figs. 10F-N) anopic to left pleuropic-anopic, various in form. Internal sac without spines. For details, see species group and species treatments, below. Ovipositor. (Figs. 11H-J, and 12A-C). Gonocoxite 2: shaft in form terete (ventral surface as wide as or narrower than lateral surface; cf. Fig. HA). Habitat.? See species accounts, below. Geographical distribution.? (Fig. 14). The range of the ampliusculus species group is co-extensive with the range of the Neotropical species of subgenus Aephnidius: through much of eis-Andean South America, the Greater Antilles and southern Bahamas, but not in Middle America, where members are not known north of Costa Rica.
Phylogenetic relationships.? Not specified.
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Anaulacus (Aephnidius) ampliusculus Chaudoir Figs. 2B, 2F, 7A, lOF-H, and 14
Masoreus ampliusculus Chaudoir, 1876: 21. TYPE MATERIAL: one authentic Chaudoir specimen, the holotype, in Oberth?r-Chaudoir Collection (MNHP), associated with following label originally pinned to the bottom of an insect box: "ampliusculus/ Chaudoir/ Para/ Bates" [handwritten]. HOLOTYPE male, labelled: "Ex Musaeo/ Chaudoir" [red print].
on Note about type material.? The original description of this species is based a no single specimen, collected at Para, by Bates (Chaudoir, 1876: 22). We have doubt that the male noted above is the one that Chaudoir had before him when he
described A. ampliusculus. Type locality.? Para (= Belem), State of Para, Brazil. Recognition.? In addition to the key character of elytral microsculpture, among the members of the ampliusculus species group, those of A. ampliusculus exhibit relatively large size (Table 2A)? cf. size of the holotype of A. bonariensis. Values for the ratio PWB/PWM (Table 3A) distinguish the latter from A. ampliusculus. Values for the ratio HW/PWM (Table 5A) distinguish adults of A. ampliusculus from the other three species of the ampliusculus species group. Males, are additionally, distinguished from other members of the ampliusculus species group by form of the median lobe: preapical portion moderately long (Fig. 10F) and relatively broad in ventral aspect (Fig. 10G; cf. Figs. 10I-N). Description.? With recognition features noted above, and character states of or as the ampliusculus species group, restricted amplified follows. Habitus. As in see Fig. 7A. For data about size (Standardized Body Length), and proportions, Tables 2A-6A. Color. Dorsal surface of head, pronotum and elytra uniformly or piceous, pronotum dark rufopiceous. Epipleura of prothorax and elytra rufous. Antennae, palpi and legs rufotestaceous. Microsculpture. Elytron (Figs. 2B, F) with mesh pattern slightly elongate, sculpticells keeled. Luster. Elytral surface seri ceous. as Pronotum. Form in Fig. 7A. Male genitalia. Median lobe (Figs. 10F-H) area anopic; prebasal curvature long, basal slightly narrower than medial area area (Fig. 10G); medial in left lateral aspect (Fig. 10F) curved evenly into periostial area, latter constricted rather markedly into narrowed moderately long preapical area; apex in lateral aspect (Figs. 10F, H) acute, in ventral aspect (Fig. 10G) narrowly rounded; preapical portion nearly straight, moderately long, broad in sac ventral aspect (Fig. 10G). Internal without spines. Ovipositor. Preapical one membrane about fifth length of gonocoxite 2. an on Habitat and activity.? This species is inhabitant the ground in lowland (probably evergreen), and probably montane, tropical forest, and in disturbed as over an sea sites, such mango plantations, altitudinal range of approximately m. level to 1500 Collections of adults at light (white and UV) indicate nocturnal flight activity. Known adult activity is in January and February, and May through August. Geographical distribution.? (Fig. 14). The range of A. ampliusculus extends from northern South America (near the Atlantic coast, from northern Brazil to Venezu ela), to Lower Middle America, in Panam? and Costa Rica. Phylogenetic relationships.? Not specified. Material examined.? In addition to the type specimens, we have seen 29 specimens of this species, as follows. COSTA RICA. Puntarenas. Male, La Nat. Amistad Park, 8?57'N 82?50'W, 1500 m, 19.11.1991, N.D. Penny (CASC). PANAMA Canal Zone. Barro Colorado Island: female, Amour Zetek 6,09?09'20"N 79?51'20"W, raking litter, 19.11.1975, T.L. Erwin, J.F. Lawrence (USNM); male, female, Miller 7.8,21.11.1975, T.L. Erwin, J.F. Lawrence (USNM); male, 2 females,
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10-17.V.1964, W.D. & S.S. Duckworth (USNM); maie, 2 females, at light, V.1941, J. Zetek (USNM); maie, berlese, 9.VIII.1949, J. Zetek (USNM); male, 6-8.II.1968, J.F. Lawrence (MCZC); male, 2-9.V.1929, P.J. Darlington Jr. (MCZC); male, 14.V.1980, H. Wolda (UCD); 3 females, 2.VI.1977, H. Wolda (USNM); female, 4.VI.1977, H. Wolda (USNM); 3 females, 5.VI.1977, H. Wolda (USNM); male, 11.VII.1977, H. Wolda (USNM); female, 21.VII.1938, E.C. Williams Jr. (FMNH). Female, Ft. Clayton, u-v light, R. Hutton (USNM). Male, Para?so, 27.1.1911, E.A. Schwarz (USNM). Female, Tabernilla, 14.V.1907, A. Busck (USNM). Panam?. Female, Las Cumbres, u-v light, H. Wolda (OSUC). Female, Soberan?a Nac. Pq., 15-24.1999, J.E.Wappes s n (JEWC). VENEZUELA. Gu?rico. Male, Guatopo NP, border, 24 km Altagracia, leaf litter under mango, 300 m, 13.VI.1987, S. & J. Peck (CMNC).
Anaulacus (Aephnidius) bonariensis Chaudoir Figs. 7B, 8B, 12A-C, and 14
Masoreus bonariensis Chaudoir, 1876: 21. TYPE MATERIAL: one authentic Chaudoir specimen, the holotype, in Oberth?r-Chaudoir Collection (MNHP), associated with following label originally pinned to the bottom of an insect box: "bonariensis/ Chaudoir/ Buenos Ayres/ Germain" HO LOTYPE female, labelled: "Ex Musaeo/ Chaudoir" [red print].
on Note about type material.? The original description of this species is based a single specimen, collected at Buenos Aires, by "M. Germain" (Chaudoir, 1876: 22). We have no doubt that the female noted above is the one that Chaudoir had before him when he described A. bonariensis.
Type locality.? Buenos Aires, Province of Buenos Aires, Argentina. Recognition.? In addition to the key character of elytral microsculpture, among the members of the ampliusculus species group, the holotype of A. bonariensis a exhibits relatively large size (Table 2A) and broad pronotal base, which is reflected in the relatively high values for the ratio PWB/PWM (Table 3A). Also, this specimen exhibits a relatively low value for the ratio HW/PWM (Table 5A), an indicating overall broad pronotum. Description.? With recognition features noted above, and character states of the ampliusculus species group, restricted or amplified as follows. Habitus. As in Fig 7B. For data about size (Standardized Body Length), and proportions, see Tables 2A-5A. Color. Dorsal surface dark overall, but varied: head piceous; pronotal disc rufopiceous, laterally paler, rufous; elytra generally rufopiceous, but humeri narrowly somewhat paler. Prothoracic and elytral epipleura rufous. Antennae and palpi rufotestaceous; legs rufous, with femora slightly darker than tibiae. Microsculpture. Mesh pattern of elytral disc isodiametric, sculpticells at convex least slightly (not examined in detail). Luster. Elytra with dorsal surface rather dull. Pronotum (Figs. 7B, 8B). Broad, laterally margins only slightly incurved as as mem posteriorly, base almost wide maximum width. Ovipositor. Preapical one brane about half length of gonocoxite 2 (Fig. 12A). Habitat.? Unknown.
Geographical distribution.? (Fig. 14). This species, at the southern extremity of the known range of Anaulacus in the Neotropical Region, is known only from the type locality, in eastern Argentina. are Chorological affinities.? Possibly this species and A. ciliatus sympatric. Phylogenetic relationships.? Not postulated. see Material examined.? Holotype only. For details, "Type material", above.
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Anaulacus (Aephnidius) ciliatus Mutchler 14 Figs. 2A, E, I, 5A, 6A, 8C, 9C-D, 10I-L, 11H-J, and
Masoreus ciliatus Mutchler, 1934:130-131. TYPE MATERIAL: four specimens, as follows: HOLOTYPE (not seen by us), collected at Ensenada, Puerto as Rico, by F. E. Lutz (AMNH). PARATYPES, three males, labelled follows: "Puerto Rico/ West Indies"; "Wolcott/ legit"; "U.S.N.M."; "PARATYPE" (USNM). "Soledad Cuba/ Cienfuegos/ X.26 1926/ Darlington"; "flood blue pool/ edge of/ woods" [handwritten]; "Fig" [handwritten; paper]; "19581/ M.C.Z./ Paratype"; "Masoreus/ (Aephnidius)/ ciliatus/ Mutchler" A. "PARATYPE" (MCZC). "Cayamas/ 16 Cuba"; "E. Schwarz/ Collector"; Mutchler" [red paper] "U.S.N.M."; "Masoreus/ (Aephnidius)/ ciliatus/ (USNM).
Note about type material.? According to Mutchler (1934: 131) the holotype under in ("Type") was collected "on the side hill back of Canary Cottage, stones, a field containing numerous cacti". ciliatus is Puerto Rico. Type locality.? The type locality of Anaulacus Ensenada, to the members of Recognition.? In addition the key character of color, among size the ampliusculus species group, those of A. ciliatus exhibit relatively small in data in (Table 2A), but nothing distinctive proportions (cf. Tables 3A-5A). are most like those A. differences in Specimens of A. ciliatus of piceolus. Slight in series are arrangement of the posterior setigerous punctures the lateral elytral distinctive (Figs. 9C-D; cf. Figs. 9A-B). Additionally, the median lobe of the male in the swollen genitalia is distinctive (Fig 71) within the ampliusculus species group and 10M basomedial portion, and in the long preapical portion (cf. Figs. 10G-H, N). states Description.? With recognition features noted above, and character of or as the ampliusculus species group, restricted amplified follows. Habitus. Similar to that of A. ampliusculus (cf. Fig. 7A). For data about size (Standardized Body see Dorsal surface Length), and proportions, Tables 2A-6A. Color. of head, pronotum and elytra uniformly piceous, or elytra rufopiceous. Prothoracic and elytral antennomeres 1-3 epipleura rufous to piceous. Antennae with testaceous, antennomeres 4-11 testaceous to rufopiceous. Palpi generally rufous, but maxil 3 testaceous lary palpomere 4 and labial palpomere apically. Legs with femora piceous, tibiae and tarsi rufotestaceous. Microsculpture. Elytra with mesh pattern isodiametric to slightly longitudinal (Figs. 2A, E), sculpticells slightly tomarkedly convex, but not keeled (cf. Fig. 2F). Luster. Elytron with dorsal surface uniformly dull, opaque, or irregularly sericeous, reflections coal-like. Pronotum (Fig. 8C). Lateral margins, including posteriolateral angles, distinctly rounded. Elytra. Posterior group of umbilical setigerous punctures with up2 distinctly mediad upl (Figs. 9C-D). Male genitalia. Median lobe (Figs. 10I-L) anopic; basal bend marked; area sharp constriction between medial and basal areas; medial (Figs. 10I-J) distinctly swollen proximally, narrowed distally and curved evenly into periostial to area, latter narrowed evenly into preapical area, with narrowing continued sac narrowly rounded apex; preapical portion moderately long. Internal without Ventral membrane about one spines (Figs. 10K-L). Ovipositor. (Figs. 11H-J). quarter length of gonocoxite 2. are Variation.? Specimens from the West Indies generally darker, the are more sculpticells of the elytra elongate and convex, and the elytral surface has a sericeous, coal-like luster, resembling in the last respect the wide-ranging In Eastern Hemisphere species A. (Aephnidius) adelioides. contrast, specimens of A. ciliatus from the American mainland are generally paler, with elytral sculpticells isodiametric and flatter, and surface dull, without coal-like luster. This variation
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we provides the basis for subspecies recognition, but elect not to take such action, at this time. an on Habitat.? This species is inhabitant the ground of rather dry lowland over an sea m. tropical forest, altitudinal range of approximately level to 500 Collections of adults at light (white, UV, and MV) indicate nocturnal flight activity. Known adult activity seems to be year-round, specimens having been collected in every month except March and October. Geographical distribution.? (Fig. 14). The range of A. ciliatus extends through eastern eis-Andean South America from northern Argentina to Venezuela. North occurs ward, this species only in Panam?, in Lower Middle America, and is widespread in the West Indian Greater Antilles. Chorological affinities.? The range of A. ciliatus overlaps the ranges, more or less, of A. batesi, three other species of subgenus Aephnidius (A. piceolus, A. ampliusculus, and A. exiguus). Phylogenetic relationships.? Not postulated. Material examined.?We have seen 129 specimens of A. ciliatus, as follows. ARGENTINA. Santa Fe. Male, Est. La Noria, Rio San Javier, 21.XII.1911, G.E. Bryant (BMNH). BRAZIL. Bahia. Female (CMNH). Goias. Male, Mt. Corumba, 21-23.11.1954, G. Gans, F. Pereira (MZSP). Matto Grosso. 2 males, Chapada (CMNH). Para. Santarem: 3 females (AMNH); 8males, 14 females (CMNH); male, female, 27-28.XII.1967 (MZSP). 2 males, 3 females, Faz. Taperinha near Santarem, 29.XII.1967-9.I.1968 (MZSP). 2 males, female, Taperinha (CMNH). COLOMBIA. Female, Vichada PNN, Tupparo Cero Tomas, 5?21'N, 67?51'W, 140 m, malaise trap, 20-29.XI.2000, W. Villalba, M. 1056 (IHVC). PANAMA. Canal Zone. 2 arc females, Ancon, light globe, IV.1991, Kraft (USNM). Male, Tabernilla, 4. VI.1907, A. Busck (USNM). Panam?. Female, dist. Chepo, Altos de Maje, at lights, 17. V.1975, u-v Stockwell, Engleman (HPSC). Female, Las Cumbres, light, 28.V.1975 (OSUC). PARAGUAY. Central. San Lorenzo, u-v light trap, J.A. Kochalka (CMNH): 9 males, 12 females, 18-19.XI.1986; 2males, 2 females, 7.XI.1986; male, 23-24.XI.1986; female, 30.XII.1987. SURINAM. Surinam. Male, Zanderij, savanna, 4-7.IX.1964, D.C. Geijskes (RMNH) VENEZUELA. Bolivar. Cuidad Bolivar, 5.VI.1898, E.A. s u-v Klages (CUIC). Guarico. 12 km Calabozo, Est. Biol. Los Llanos, light, P. & P. Spangler (USNM): male, 6-12.II.1969; female, 6.II.1969; male, 6-12.II.1969. WEST INDIES. Bahama Islands. Male, 2 females, Exuma, Georgetown, Hummingbird Cay, 11.VI.1968, B.K. Dozier (FSCA). 3 males, 8 females, Mayaguana I., 21 28.VIII.1963, CM. Murvosh (FSCA). Cayman Islands. Female, 17.IV-26.VII.1938, C.B. Lewis, J.H. Thompson (BMNH). 2 males, Grand Cayman, Hell, 19?23'N 81?24'W, 19.11.1993, W.E. Steiner, J.M. Swearington (USNM). Grand Cayman, sw point, ex suction trap, 11-12.XI.1970, J. Davies (BMNH). Cuba. Oriente. Male, Maisi, 17.VII.1936, Darlington (MCZC). Dominican Republic: Monte Cristi. 2 females, 8.2 km n Villa Elisa, mv and bl, 1.VI.1994, R. Turnbow (RHTC). 3 males, 15 females, 5 km nne Botoncillo, 19?46'N71?24'W, arid thorn scrub, 50 m, 29 30.XI.1992, R. Davidson, M. Klingler, S. Thompson, J.Rawlins (CMNH). Pedernales. 2 females, Cabo Rojo, swimming pool surface, coastal thorn scrub, 10 m, 28.XI 2.XII.1991, L. Masner, S. Peck (CMNC). Puerto Rico. 2 males, Wolcott (MCZC) (USNM). Female, Smythem from Wolcott (BMNH). 2 males, La Parguera, 28.VII.1969, H..& A. Howden (UASM). Turks and Caicos Islands. 4 males, 2 u-v females, Providenciales, Grace Bay Hills, sandy scrub forest, light, 27.1.1998, W.E. Steiner, J.M. Swearington (USNM).
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new Anaulacus (Aephnidius) humeralis, species Figs. 9E-F, lOM-N, and 14
Type material? HOLOTYPE, male, labelled: "Sta.Maria, C. R./ (Sn. Mat.)/ X. '39, Bierig"; "M winged"; "ADP/ 05932" [MCZC]. PARATYPE, male, labelled same as holotype, except: "V winged"; "ADP/ 05933" [MCZC]. Type locality. The type locality of this species is Santa Mar?a de Dota, Province of San Jos?, Costa Rica. Specific epithet.? The word "humeralis" is Latin, meaning pertaining to the as a noun shoulder, and used here in apposition, in allusion to the pale-colored humeri of adults of this species. Recognition.? In addition to the key character of arrangement of the apical series of lateral setigerous punctures, among the members of the ampliusculus are species group, those of A. humeralis relatively small (Table 2A). Values for the are a narrow ratio PWB/PWM (Table 3A) low, indicating relatively pronotal base. The species is characterized externally also by the pale elytral humeri. Description.? With recognition features noted above, and character states of or as the ampliusculus species group, restricted amplified follows. Habitus. Similar to that of A. ampliusculus (cf. Fig. 7A). For data about size (Standardized Body see Length), and proportions, Tables 2A-6A. Color. Dorsal surface of head and pronotum uniformly piceous; elytron principally piceous, but hum?rus with pale vitta extended laterally from lateral margin approximately to stria 3, and posteri one orly about third length of elytron. Antennae, palpi, prothoracic and elytral epipleura paler, rufo to rufotestaceous. Microsculpture. Elytron with sculpticells convex (cf. Fig. 7E), slightly elongate. Luster. Elytra with dorsal surface dull. Pronotum: Form similar to pronotum of A. piceolus (cf. Fig. 8A), but more con stricted posteriorly. Elytra: Posterior group of umbilical setigerous punctures with seta up2 distinctly laterad seta upl (Figs. 9E-F). Male genitalia. Median lobe (Figs. area narrower 10M-N) anopic; basal bend gradual, basal slightly than medial area; area medial evenly continuous with preapex in left lateral aspect, (Fig. 10M) in more ventral aspect (Fig. ION), markedly constricted; preapical portion in left lateral aspect (Fig. 10M) parallel sided, curved ventrad rather sharply, rather short. Internal sac without spines. a Habitat.? Unknown, but probably ground inhabitant, in lowland tropical forest.
Geographical distribution.? (Fig. 14). This species is known only from the type locality in south central Costa Rica. Chorological affinities.? The range of A. humeralis is overlapped by the ranges of A. piceolus and A. sericatus, and possibly by the range of A. pittieri. Phylogenetic relationships.? Not postulated. Material examined.? We have seen only the type material of A. humeralis. For see details, that topic, above.
The exiguus species group
Recognition.? A combination of the following features distinguishes adults of the exiguus species group from those of the other Neotropical species groups of subgenus Aephnidius: small size (SBL less than 3.0 mm); pronotum distinctly paler than head and elytra; elytral microsculpture with mesh pattern isodiametric to diagonal, and sculpticells flat (cf. Figs. 2C-G), surface of disk shiny, subiridescent laterally, and umbilical puncture 3 closer to 4 than to 2 (Figs. 9G-H). Males lack spines from the internal sac. Other distinctive features are details of the male as median lobe (Figs. 10O-T), follows: shaft with preapical portion and ostial area relatively short (Table 6B). Description.? With diagnostic features of subgenus Aephnidius, and character
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Fig. 1. Photograph, illustrating habitus, dorsal aspect, o? Anaulacus (Apostolus) batesi Chaudoir. SBL = 6.44 mm.
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m
Fig. 2. SEM photographs of microsculpture of portions of elytra: dorsal aspect, A-D, I-M, and O; lateral aspect, E-H, and N. A, E, Anaulacus (Aephnidius) ciliatus Mutchler; B, F, A. (Aephnidius) ampliusculus Chaudoir; C, G, A. new new (Aephnidius) exiguus, species; D, H, A. (Aephnidius) ashei, species; I, A. (Aephnidius) ciliatus, in vicinity of basal ridge; J- O, A. (Macracanthus) near tuberculatus Chaudoir (M, lateroapical margin). Scale bars: A-C, E-H, K & = = = N-O 10 urn; D & I 50 urn; and J-K & M 100 um.
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Fig. 3. SEM photographs of mandibles of Anaulacus (Aephnidius) adelioides MacLeay. A, C, left mandible, dorsal and ventral aspects, respectively; B, D, right mandible, dorsal and ventral aspects, respectively; E, left mandible = urn. posteriolateral aspect. Scale bars 100 Legend: aog, anterior occlusal groove; art, anterior retinacular tooth; b, basal area; bb, basal brush; bemr, basal extension of occlusal groove; i, incisor tooth; 1dm, laterodorsal margin; m, molar tooth; mr, molar ridge; oemr, occlusal extension of molar ridge; pm, premolar tooth; pog, posterior occlusal groove; prt, posterior retinacular tooth; rr, retinacular ridge; s, scrobe; t, terebra; tr, terebral ridge; tt, terebral = tooth; vg, ventral groove. Scale bars 200 urn.
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Fig. 4. SEM photographs of labium of Anaulacus (Aephnidius) adelioides MacLeay. A, mentum and prementum, ventral aspect; B, prementum, ventral aspect; C, = = um. prementum, dorsal aspect. Scale bars: A 200 um; B, C 40 Legend: del, dentiform projection of epilobe; gs, glossal seta; gsc, glossal sclerite; 11, lateral lobe; lpl, labial palpomere 1; lp2, labial palpomere 2; lp3, labial palpomere 3; = urn = urn. m, mentum; pg, paraglossa. Scale bars: A 200 and B-C 40
or as states of genus Anaulacus, restricted amplified follows. Habitus. Not illus trated, but form similar to A. ampliusculus (Fig. 7A). For data about size (Standard see ized Body Length), and proportions, Tables 2B-6B. Color. Somber, head capsule, except clypeus, piceous; clypeus, prothorax, and elytral epipleura rufous to rufotestaceous; elytra with dorsal surface rufopiceous; appendages, including labrum and mandibles, testaceous. Microsculpture (Figs. 2C-G) and luster. Head convex to capsule: Clypeus with mesh pattern isodiametric, sculpticells slightly flat, microlines distinct, surface subopaque. Labrum: Mesh pattern isodiametric, sculpticells large, convex, microlines clearly indicated, surface subopaque. Scutel lum: Mesh pattern transverse. Elytra: Mesh pattern isodiametric to transverse in intervals 1-5, diagonal in intervals 6-8, sculpticells flat, surface shining to subiridescent. Head. Antennae: rather short, in backfolded position, antennomere 11 extended to posterior margin of pronotum. Antennomeres 5-10 elongate, each more or only slightly longer than wide, less cylindrical. Mouthparts. Labrum with anterior margin straight. Elytra. Posteriolateral umbilical setigerous punctures as arranged in Figs. 9G-H. Male genitalia. Median lobe (Figs. 10 O-T) left pleuropic area narrow catopic, various in form; preapical in ventral aspect flattened, to broad. Internal sac without spines. For details, see species treatments, below. Ovipositor. (Figs. 11E-J, and 12A-F). Gonocoxite 2: shaft in form terete (ventral surface as wide as or narrower than lateral surface (cf. Fig. 11A); ventral membrane one half length of gonocoxite 2.
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Fig. 5. SEM photographs of elytra and legs of Anaulacus (s. tat.) species: A-B, A. elytral base, respectively, of: A, A. (Aephnidius) ciliatus Mutchler and B, (Macracanthus) tuberculatus Chaudoir. C-D, posterior portions of elytra of A. new (Aephnidius) exiguus, species, showing arrangement of umbilical setiger ous setae: C, posteriolateral portion of left elytron, dorsolateral aspect; D, left elytron, dorsal aspect. E-F, left middle tibia, distal portion, showing setal brush = of A. (Macracanthus) sericatus Chaudoir: E, male and F, female. Scale bars 200 urn. Legend: br, basal ridge of elytron; mt, middle tibia; psp, parascutellar setigerous puncture; sb, setal brush of middle tibia; tsl, tarsomere 1; tsp, tibial spine; upl- 3 umbilical setigerous punctures 1-3, respectively.
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of Anaulacus Fig. 6. Line drawings of hind wing and female genitalia species. A, left hind wing, dorsal aspect, of A. (Aephnidius) ciliatus Mutchler. B, female Gahan. Scale bars = 1mm. genitalia, ventral aspect, of A. (Macracanthus) pittieri anterior veins 1 +2 and 3+4 Legend: AA1+2 and 3+4, anal respectively; be, common 1 and bursa copulatrix; co, oviduct; gcxl, 2, gonocoxites 2, respec tively; MP3, 4, posterior median veins 3 and 4, respectively; o, oblongum cell; RA4, anterior radial vein 4; RP2, and 3+4, posterior radial veins 2 and 3+4, sternum respectively; sIX, abdominal IX; sp, spermatheca; spg, spermathecal gland; spgd, spermathecal gland duct.
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7. Fig. Photographs, illustrating habitus, dorsal aspect, of Anaulacus (Aephnidius): = A, A. ampliusculus Chaudoir, SBL 4.14 mm.; B, A. bonariensis Chaudoir, = mm. Holotype, SBL 3.84
Habitat.? See this topic for the species treated below. Geographical distribution.? (Fig. 14). The range of the exiguus species group extends in cis-Andean South America from southern Paraguay northward to southernmost Chiapas, in M?xico, assuming that the large gap between southern eastern some M?xico and Peru is occupied by yet to be discovered populations of or described species, by presently unknown species. Phylogenetic relationships.? Not postulated. Included species.? This group includes three species, all described here for the first time: A. exiguus, A. whiteheadi, and A. ashei.
Anaulacus (Aephnidius) exiguus, new species Figs. 2C-G, 5C-D, 9G-H, 10O-P, and 14
Type material.?HOLOTYPE, male, labelled: "PERU: Tambopata Prov./ Madre de Dios Dpto./ 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nico/ "7 12?33'S, 69?03'W/ 200m, swamp trail"; July 1989, J. S. Ashe,/ R. A. Leschen #444/ ex., flower fall, sifted/ berlese" [SEMC]. ALLOTYPE, female, labelled: "PERU: Tambopata Prov./ Madre de Dios Dpto./ 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nico/ 12?33'S, 69?03'W/ 200m, Z2U21"; "10 July 1989, J. S. Ashe,/ R. A. Leschen #468/ ex., flowerfall/ slush /litter" [SEMC]. 29 PARATYPES, sex and label data follows. 2 males, female: "PERU: Tambopata Prov./15 km NE Pto. Maldonado/ 7 July 1989,200 m/ J.Ashe, R.Leschen, #438/ ex: liana fruit fall" [SEMC]. Female: "PERU: Tambopata Prov./15 km NE Pto. Maldonado/ 5 July
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mm
Fig. 8. Line drawings of pronotum, dorsal aspect, of species of Anaulacus (Aephnidius): A, A. piceolus Chaudoir; B, A. bonariensis Chaudoir; and C, A. ciliatus Mutchler. Scale bars = 1.0 mm.
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Fig. 9. Line drawings of elytral apices, dorsal aspect, o? Anaulacus (Aephnidius) species. A, C, E, G, dorsal aspect; B, D, F, H, dorsolateral aspect. A-B, A. piceolus new Chaudoir; C-D, A. ciliatus Mutchler; E-F, A. humeralis, species; and G-H, A. new = mm. exiguus, species. Scale bar 1.0 Legend: upl-4, umbilical setigerous punctures 1-4, respectively.
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Fig. 10. Line drawings of male median lobes of: Anaulacus (Aephnidius) species: A, C, F, I, K, M, O, Q, S, left lateral aspect; E, H, L, right lateral aspect; B, D, G, J,N, P, R, T, ventral aspect. A-B, A. batesi Chaudoir; C-E, A. piceolus Chaudoir; F-H, A. ampliusculus Chaudoir; I-L, A. ciliatus Mutchler; M-N, A. humeralis, new new new species; O-P, A. exiguus, species; Q-R, A. whiteheadi, species; and S new = mm. = mm. T, A. ashei species. Scale bars: A-B 1 and C-T 0.5 Legend: bb, basal bulb; bo, basal orifice; m, medial part; oo, ostium; pa, preapical part; pb, sac. prebasal bend; po, periostial part; s, basal spine patch of internal
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Fig. 11. SEM photographs of left gonocoxite 2 of species of Anaulacus (s. lat.): A, E, H, lateral aspect; B, D, F, I, ventral aspect; C, G, J,median aspect. A-D, A. (Apostolus) batest Chaudoir; E-G, A. (Aephnidius) piceolus Chaudoir; and H-J, A. = = (Aephnidius) ciliatus Mutchler. Scale bars: A-C and E-J 100 urn and D 10 urn. Legend: ns, nematiform seta; vm, ventral membrane.
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Fig. 12. SEM photographs of ovipositor sclerites of species o? Anaulacus (s. lat.). Left gonocoxite 2, only: A, D, G, lateral aspect; B, E, H, ventral aspect; C, F, I, median aspect. Left gonocoxites 1 and 2: J. A-C, A. (Aephnidius) bonariensis Chaudoir; D-F, A. (Aephnidius) adelioides MacLeay; and G-J, A. (Mact-acanthus) = urn. sericatus Chaudoir. Scale bars 100 Legend: gel, 2, gonocoxites 1 and 2, respectively; vm, ventral membrane.
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Fig. 13. Map of southern part of United States of America, Middle America, and South America showing positions of known localities for the species of Anaulacus (Apostolus) batest Chaudoir and A. (Aephnidius) piceolus Chaudoir.
1989, 200 ml J.Ashe, R.Leschen, #395/ ex.:flower fall" [SEMC]. Female: "PERU: Tambopata Prov./ Madre de Dios Dpto./ 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nico/ 12?33'S, 69?03'W/ 200m, swamp trail"; "13 July 1989, J. S. Ashe,/ R. A. Leschen #501/ ex., fungusy log" [SEMC]. Male, 2 females: "PERULoreto Prov./Iquitos, 90 m, 8 May/ 1992, J. Danoff-Berg/ ex:sifter leaf litter in/ forest" [SEMC]. Female: "PERU: Madre de Dios/ Pakitza, Zone 02/ fine Tr. 09Sep89 T.L.Erwin/ 12?07'S 70?58'W"; "Under leaves on/ loam/sand/ Aguajal/107"; "BIOLAT/Cole? 000007975" [USNM]. Male: "COLOMBIA: Amazonas/ Leticia, 15-VI-65/ P. R. Craig/ and J. Robb [CASC]. Male: "Braz.MatoGrosso/ Barra do Tapi-/ rape XII-23-62/ B.Malkin [CASC]. Male: "BRASIDM. Grosso./ Dec.1-11,1960./ B. Malkin leg."; "Tapirape Indian/ Village
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Fig. 14. Map of southern part of United States of America, Middle America, and South America showing positions of known localities for the species of the ampliusculus and exiguus species groups o? Anaulacus (Aephnidius) (MacLeay). at confluence/ of "at/ [FMNH]. Female, same R.Tapirapeand/" R.Araguia"; light" as previous except Nov. 11-20, 1960" [FMNH]. Male: "BRASIL:/ M. Grosso./ at XI:15:60/ B. Malkin leg."; "Tapirape Indian/ Village confluence/ of R.Tapirape and/ R.Araguia"; "at/ light" [FMNH]. Female: "BRAZIL, M.G. / Jacare,P.N.Xingu/ XI-1965,at light/ M. Alvarenga" [USNM]. Female: "Jacar?/ Pq.Nac.Xingu, MT/ XI.1961, Alvarenga/ & Bokermann col." [USPB]. Male, female: "BOLIVIA: COCHABAMBA/ 67.5 km E Villa Tunari, Est. Biol./ Valle Sajita, Univ. San Simon/ 300m, 17?06'19"S 64?46'57"W/ 7.II.1999, R.Anderson, lowland/ rain same as forest litter, 99-027A" [CMNC]. Female, previous, except "99-027C" [CMNC]. 2 males, 2 females: "ARGENTINA: Chaco/ 100km NW Resistencia/
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Fig. 15. Photograph, illustrating forebody, dorsal aspect, of amber-fossilized specimen of Anaulacus (Aephnidius) species incertae sedis, from "Puerto Plata", Dominican Republic, Hispaniola.
P.N. Chaco, 14.XII.1990/ S.&J. Peck, rotted logs &/ sticks with fungus, forest/ 90 113" [CMNC]. ]. 2 males: "ARGENTINA: Chaco/ 100km NW Resisten-/ cia, P.N. Chaco, 14.XII./ 1990, S.&J. Peck, thorn-/ forest leaflitter, 90-115" [CMNC]. ]. 2 females: "ARGENTINA: Chaco/ 100km NW Resistencia/ P.N. Chaco, 13.XII.1990/ S.&J. Peck, forest tree-/ base litter, 90-109" [CMNC]. Female: "ARGENTINA: Chaco/ Chaco NP, 100km NW/ Resistencia 15.XII.1990/ S.&J. Peck subtropical thorn/ for. Rotted logs. 90-116" [CMNC]. Type locality.? The type locality of this species isMaldonado Reserve, 12?33'S m 69?03'W, 200 altitude, 15 km NE Puerto Maldonado, Departamento Madre de Dios, Peru,. Specific epithet.? The word "exiguus" is Latin, meaning small, and used here as a noun in apposition, in allusion to the small size of adults of this species. Recognition.? Elytral microsculpture (Figs. 2C, G) with mesh pattern in intervals 1-2 to 1-5 isodiametric, distinguishes adults o? A. exiguus from those of A. ashei, specimens of which exhibit completely transverse mesh pattern in elytral intervals 1-5. Within this group, the male median lobe (Figs. 10O-P; cf. Figs. 10Q T) is distinctive in ventral aspect with very broad preapical part.
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Fig. 16. Photograph illustrating habitus, dorsal aspect, of Anaulacus (Macracanthus) sericatus Chaudoir. SBL = 4.20 mm.
Description.? With recognition features noted above, and character states of or as the exiguus species group, restricted amplified follows: For data about size see (Standardized Body Length), and proportions, Tables 2A-6A. Microsculpture (Figs. 2C, G). See "Recognition", above. Male genitalia. Median lobe (Figs. 10O-P) not area left pleuropic-catopic; basal bend evident dorsally (Fig. 10O), basal broad, basal bulb area in (Fig. 10P) prominent; medial left lateral aspect (Fig. 10O) curved evenly into periostial area, latter narrowed evenly into preapical area, with narrowing continued preapically to rather broadly rounded apex, especially marked in ventral aspect (Fig. 10P); preapical part rather short. Internal sac without spines. Ovipositor. Ventral membrane about half length of gonocoxite 2. Variation.?Mesh pattern of elytral microsculpture varies in intervals 3-5 from to isodiametric transverse, with isodiametric being the pattern in intervals 3 and 4 most for individuals, but transverse being predominant in interval 5. a Habitat.? This species is ground-living inhabitant of tropical evergreen m. forest and subtropical thorn forest, between 90 and 300 Collections of adults at light indicate nocturnal flight activity. Known adult activity extends possibly
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Anaulacus Fig. 17. Line drawings of pronotum, dorsal aspect, of species of new (Macracanthus): A, A. sericatus Chaudoir and B, A. mcclevei, species. Scale bars = 1.0 mm.
are throughout the year, though records for only the months of May, July, September, November and December. in Geographical distribution.? (Fig. 14). This species is known from localities the upper Amazon Basin of Brazil, Peru, and Bolivia, and in the Rio Paran? Basin of Paraguay and Argentina. Chorological affinities.? The ranges of A. exiguus and of its relative, A. ashei, A. overlap broadly in the upper reaches of the Amazon Basin. However, exiguus is geographically isolated from the range of its other relative, A. whiteheadi (Fig. 14). are Phylogenetic relationships.? Although adults of A. exiguus and A. whiteheadi similar in their varied elytral microsculpture mesh pattern in intervals 1-5, the isodiametric pattern is predominant in A. exiguus. On this basis A. exiguus is more ancestor + postulated to be the primitive of the two, and the of A. whiteheadi A. ashei. seen Material examined.? We have only type specimens of A. exiguus. For see details, "Type material", above.
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Fig. 18. Line drawings of male median lobes of species of Anaulacus (Macracanthus): A, C, E, G, I, K, left lateral aspect; B, D, F, H, J, L, ventral aspect. new A-B, A. sericatus Chaudoir; C-D, A. thoracicus, species; E-F, A. erwini, new new species; G-H, A. mcclevei, species; I-J, A. pittieri Gahan; and K-L, A. = tuberculatus Chaudoir. Scale bars: A-D and G-L 1.0 mm., and E-F = 0.5 mm. Legend: bb, basal bulb; bo, basal orifice; m, medial part; mb, medial bend; oo, ostium; pa, preapical part; pb, prebasal bend; po, periostial part.
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Fig. 19. Photographs illustrating habitus, dorsal aspect, of Anaulacus = (Macracanthus) tuberculatus species group: A, A. pittieri Gahan, SBL 4.84 mm. = and B, A. tuberculatus Chaudoir, SBL 4.48 mm.
new Anaulacus (Aephnidius) whiteheadi, species Figs. 10Q-R, and 14
Type material? HOLOTYPE, male, labelled: "M?XICO, Chiapas/ San Quintin7007 91o20'-16o24'/rain forest/ II. 5-20. 1966"; "George E. Ball/ D. R. Whitehead/ Collectors" [USNM]. PARATYPE, male, labelled: "BELISE: Orange Walk Dist./ Rio Bravo Conserv. Area/ ( Archaeological camp)/ 17?50'04"N,89?03'04"W/ 2-V-1996, C Carlton #089/ ex: berlese [SEMC]. Type locality.? The type locality of this species is the town of San Quintin, State at m of Chiapas, M?xico, 16?24'N latitude, 91?20'W longitude, and 213 altitude. The holotype is mislabelled in that longitude is recorded before latitude. a Specific epithet.? The word "whiteheadi" is Latinized genitive eponym, based on surname now the of Donald R. Whitehead, deceased, formerly scientist in the Systematic Entomology Laboratory, United States Department of Agriculture, and former associate and long time friend of the senior author, who accompanied on a to him year-long expedition M?xico, during which the holotype of this species was collected. a Recognition.? Adults of A. whiteheadi exhibit microsculpture mesh pattern of as the elytra which, in A. exiguus, is partially isodiametric and partially transverse to diagonal. This feature distinguishes A. whiteheadi from A. ashei. Additionally, are within the exiguus species group, males distinguished by the preapical part of the median lobe (Figs. 10Q-R; cf. Figs. 10O-P and 10S-T): in left lateral aspect, broader, but in ventral aspect narrower.
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Fig. 20. Map of southern part of United States of America, Middle America, and South America, showing positions of known localities for the species of Anaulacus (Mact-acanthus).
Description.? With recognition features noted above, and character states of or as the exiguus species group, restricted amplified follows: For data about size see (Standardized Body Length), and proportions, Tables 2A-6A. Male genitalia. Median lobe (Figs. 10Q-R) left pleuropic-catopic; basal bend rather abrupt, basal area narrower rather large, slightly than medial area; basal bulb moderately area prominent (Fig. 10R); medial in left lateral aspect (Fig. 10Q) curved evenly area in into periostial area, latter hardly narrowed to broad preapical (narrower ventral aspect (Fig. 10R), apex, broadly rounded; preapical portion short. Internal sac without spines. Variation.? Mesh pattern of elytral microsculpture varies in interval 3 from isodiametric to transverse. were on Habitat and activity.? Both known specimens collected the ground in we assume lowland tropical evergreen forest, and that such is the habitat of this
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species. Collections were made in February and May. Geographical distribution.? (Fig. 14). This species is known only from south eastern M?xico and nearby Belize. Chorological affinities.? The known range of this Middle American species is widely isolated from the ranges of its South American, putatively close, relatives. on Phylogenetic relationships.? Based the putatively synapotypic transverse elytral microsculpture mesh pattern of intervals 3-5, A. whiteheadi and A. ashei are postulated to be adelphotaxa. Material examined.? We have seen only the type material of A. whiteheadi. For details, see that topic, above.
new Anaulacus (Aephnidius) ashei, species Figs. 2D, H, 10S-T, and 14
Type material? HOLOTYPE, male, labelled: "PERU: Tambopata Prov. / Madre de Dios Dpto./ 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nico/ 12?33'S, 69?03'W/ 200m, camp"; "19 July 1989, J. S. Ashe,/R. A. Leschen#148/ ex., compost pile" [SEMC]. ALLOTYPE, female, labelled: "PERU: Tambopata Prov./ Madre de Dios Dpto./ 15kmNE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nica/ 12?33'S, 69?03'W/ 200m, swamp trail"; "13 June 1989, J. S. Ashe,/R. A. Leschen sex #501/ ex., fungus" [SEMC]. 10 PARATYPES, and label data follows. Male, same as same as holotype [SEMC]. Female, allotype [SEMC]. Female, "PERU: Tambopata Prov. / Madre de Dios Dpto. / 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nica/ 12?33'S, 69?03'W/ 200m"; "22 June 1989, J. S. Ashe,/R. A. " Leschen #214/ ex., compost pile [SEMC]. Male, "PERU: Tambopata Prov./ 15 km NE Pto. Maldonado/ 19 June 1989, 200 m/ J.Ashe, R.Leschen #148/ ex., compost pile" [SEMC]. Male, female, "PERU: Tambopata Prov./ 15 km NE Pto. Maldonado/ 22 June 1989, 200 m/ J.Ashe, R.Leschen #218/ ex., compost pile" 2 [SEMC]. females, "PERU: Tambopata Prov./ 15 km NE Pto. Maldonado/ 13 July 1989,200 m/ J.Ashe, R.Leschen #501 / ex., fungusy log" [SEMC]. 2 females, "Para, Brazil/ Ace. No. 2966" [CMNH]. Type locality.? The type locality of this species isMaldonado Reserve, 12?33'S m 69?03'W, 200 altitude, 15 km NE Puerto Maldonado, Departamento Madre de Dios, Peru. a Specific epithet.? The word "ashei" is Latinized genitive eponym, based on surname the of James S. Ashe, Snow Entomological Museum, University of Kansas, friend and former associate of the authors, and one of the collectors of the type series of A. ashei. are Recognition.? Adults of A. ashei distinguished from other two members of the exiguus species group by the transverse mesh pattern of elytral intervals 1-5. are Additionally, within the exiguus species group, males distinguished by the very short preapical part of the median lobe (Figs. 10S-T; cf. Figs. 10 O-R). Description.? With recognition features noted above, and character states of or as the exiguus species group, restricted amplified follows. For data about size see (Standardized Body Length), and proportions, Tables 2A-6A. Microsculpture (Figs. 2D, H). See "Recognition", above. Male genitalia. Median lobe (Figs. 10S-T) left area pleuropic-catopic; basal bend rather abrupt, basal rather large, slightly narrower than medial area; basal bulb moderately prominent (Fig. 10T); medial area in left lateral aspect (Fig. 10S) curved evenly into periostial area, latter hardly area narrowed to broad preapical (narrower in ventral aspect (Fig. 10R), apex, sac broadly rounded; preapical portion very short. Internal without spines. a Habitat and activity.? This species is ground-living inhabitant of lowland at tropical evergreen forest, known presently only from localities approximately m 200 altitude, with adults active in June and July.
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Geographical distribution.? (Fig. 14). This species is known only from two localities in South America: one in Brazil, near the mouth of the Amazon River; and one, from the upper reaches of the Amazon Basin, in southeastern Peru. Chorological affinities.? The range of A. ashei probably is overlapped by the a range of the close relative, A. exiguus, and by the ranges of number of species of Anaulacus, in other groups and subgenera. are Phylogenetic relationships.? This species and A. whiteheadi postulated to be adelphotaxa, based on the more derived elytral microsculpture that they exhibit, compared to A. exiguus. Material examined.? We have seen only type specimens of A. ashei. For details, see "Type material", above.
Anaulacus (Aephnidius) species incertae sedis Fig. 15
Included here are three specimens, preserved in amber, from the Dominican on are as Republic, the island of Hispaniola. They follows: female, Cordillera Septentrional, vicinity of the Palo Alto de las Auryamas amber mine; Susan sex Hendrickson (UASM); not determined, La Toca Mine (No. C7-86C George O. Poinar, Jr. coll.); and female (Fig. 15), from "Puerto Plata", without identifying was a number (UASM). The specimen purchased in jewelry store, in New York on City. Puerto Plata is the Atlantic coast of the Dominican Republic, and is the source one of origin of of the roads to the nearby Cordillera Septentrional, where are located many amber mines, including the one at La Toca, and at Palo Alto. The a Puerto Plata specimen probably is also from deposit in the Cordillera Septentrional. According to Poinar (1992: 37) the age of the amber from the La Toca mine is Oligocene-Miocene (30-45 mybp) and for the Palo Alto mine, it is Early Miocene as (20-25 mybp). However, noted by Poinar & Mastalerz (2000: 180), Dominican a amber is secondarily deposited in sedimentary rocks, which makes definite age we on determination difficult. So, prefer not to place much weight the putative age difference between the two sites.
The La Toca specimen is smaller (length or elytra, 2.69 mm.) than the Palo Alto specimen (length of elytra, 2.98 mm.). One may be tempted to argue that, since these specimens may have lived at markedly different times, and in view of the size difference, they could be specifically distinct from one another. But, consid to one ering their general similarity another, and that the putative temporal we are difference may be incorrect, disinclined to place much weight on the size difference. (The La Plata specimen is the same size as the La Toca specimen, but without knowing its exact source, it cannot be considered in relation to the matter at extant issue). So, for purposes of comparison with Western Hemisphere speci mens we treat as and species o? Anaulacus, the fossils conspecific with one another. as Important details for classification, such male genitalia and sclerites of the are not evident. ovipositor The slightly elongate mesh pattern of elytral microsculpture excludes these specimens from subgenus Macracanthus, and from the exiguus species group of subgenus Aephnidius. Relatively small size and as pronotal form exclude them from the subgenus Apostolus this taxon is currently understood. However, Apostolus ismonobasic, so its limits are circumscribed very are most or tightly. The fossils like members of either the piceolus ampliusculus in group of subgenus Aephnidius, size, general form (Fig. 15; cf. Fig. 7A), and elytral microsculpture. Because of difficulty in measuring two of the specimens, we confine remarks about size to relative elytral measurement, which, for the three specimens, ranges to mm a mean mm. from 2.69 2.98 with of 2.79 In comparison, for a series of six specimens of A. ciliatus from the Dominican Republic, the only species of the
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ampliusculus group inhabiting the West Indies, elytral length ranges from 1.84 to amean mm. are 2.52 mm, with of 2.33 Thus, the fossils consistently larger than the seem measured specimens of A. ciliatus, which, in size, to be typical of the series from which they were drawn. As shown by data in Tables 2A and 2B, adults of A. piceolus (easily confused with A. ciliatus; wide-ranging in the Neotropical main are same land, but not occurring in the West Indies) about the size as are those of A. ciliatus, and consequently, are smaller than the fossils. The mean value of elytral length for the sample of A. ampliusculus, with largest was individuals included in the ampliusculus species group, 2.55 mm., the largest mm. are specimen with elytral length of 2.68 Thus, the fossils consistently and at A. least marginally larger than the specimens of ampliusculus. This evidence of distinctness of the fossils is not particularly convincing, but are it shows that they consistently larger, and indicates that the species they represent may be different from the known extant species of the ampliusculus group, and from A. piceolus, the only species of the piceolus group. we In the absence of convincing evidence to the contrary, exclude the fossils from membership in any of the extant species of Western Hemisphere Aephnidius. as a new This could lead to describing them members of species. However, in the are one absence of evidence that the fossils really conspecific with another, and considering that size is the only feature available at present to distinguish them, we are a we disinclined to propose formal taxon for them. Accordingly, treat these specimens as noted above.
Subgenus MACRACANTHUS Chaudoir, NEW COMBINATION
Macracanthus Chaudoir, 1846: 539.? Reichardt, 1977: 430.? Erwin and Sims, 1984: 395. Masoreus (Macracanthus); Schaum, 1860: 179.? Chaudoir, 1876: 23.? Bates, 1883: 174. van Ophryognathus Chaudoir, 1876: 26.? Emden, 1949: 889.? Reichardt, 1977: 431. NEW SYNONYMY.
Leptotomus Gahan, 1894: 115.? van Emden, 1949: 889. NEW SYNONYMY.
name Type species.?For the Macracanthus, M. sericatus Chaudoir, 1846 is type species; for Ophryognathus, O. tuberculatus Chaudoir, 1876; and for Leptotomus, L. are pittieri Gahan, 1894. All three type species by monotypy. Notes about synonymy and classification.? As understood by Chaudoir (1876) van were and Emden (1949), Macracanthus and Ophryognathus clearly distinguish was able from one another. The marked gap recognized by those authors partially filled by inclusion of L. pittieri in Ophryognathus. The diagnostic combination of differences that remained was bridged with discovery of A. mcclevei, new species. In many respects, this species isMacracanthus (s. sfr.)-like. But specimens exhibit the color and mandibular lateral marginal ridge of Ophryognathus, and the form of two the hum?rus is about midway between that of the subgenera. Under these circumstances, one could include A. mcclevei in its own monospe seems us to use cific subgenus. But it to preferable the virtual continuum of a structural features to combine in single subgenus Macracanthus and Ophryognathus, as same ranking them species groups, and treating in the way, because of its own intermediacy, A. mcclevei, assigning it to its monobasic species group. Thus, we recognize three species groups named, respectively sericatus (=Macracanthus s. str.), tuberculatus, (^Ophryognathus) and mcclevei. to Recognition.? See key the Neotropical subgenera and species o? Anaulacus, above. The pronotum is broad at base (Figs. 16,17A-B, and 19A-B), with length and maximum width moderately divergent (values for ratio PL/PWM less than 0.6;
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Table 4B). The internal sac of the male genitalia is without spines, and the median lobe is left pleuropic, or left pleuropic-anopic (Figs. 18A-L). Gonocoxite 2 of the Ovipositor. (Figs. 6B, and 12G-J) is asetose, relatively slender, and with ventroapical membrane approximately one fifth length of gonocoxite 2. Description.? With recognition features noted above, and character states of genus Anaulacus, restricted or amplified as follows. Habitus. As in Figs. 16, and 19A-B. For data about size (Standardized Body see Length), and proportions, Tables 2A-6B. to Color. Somber, body piceous rufous; elytra concolorous; appendages pi ceous to testaceous.
Microsculpture (Figs. 2A-0) and luster. Head capsule: Clypeus with mesh pattern isodiametric, sculpticells slightly convex, microlines distinct, surface or subopaque, mesh pattern transverse, microlines evanescent, surface shining. Frons with mesh pattern isodiametric medially, longitudinal laterally, sculpticells or slightly convex, surface subopaque, mesh pattern transverse, sculpticells flat, and microlines finer, subiridescent. Vertex with mesh pattern isodiametric to convex sur longitudinal posteriorly, sculpticells slightly and microlines deeper, face subopaque. Labrum. Mesh pattern isodiametric, sculpticells large, convex, microlines clearly indicated, or mesh pattern transverse, sculpticells flat. Man or dibles. Dorsal surface shining, mesh pattern longitudinal, microlines shallow, microlines not evident, surface smooth. Pronotum. Disc with mesh pattern trans verse to irregular, sculpticells flat, microlines shallow, surface subopaque to subiridescent. Lateral declivity with mesh pattern transverse, sculpticells flatter, surface shining to subiridescent. Scutellum. Mesh pattern isodiametric to trans verse, sculpticells slightly convex to flat. Elytra. Mesh pattern (Figs. 2J-L) of disc markedly longitudinal, sculpticells slightly convex, in parallel rows, surface markedly iridescent. Head. Antennae: shorter, with antennomere 11, back-folded, extended only or less than distance to posterior margin of pronotum. Antennomeres 5-10 quadrate or (Fig. 16) slightly wider than long (Figs. 19A-B), and somewhat compressed. or Mouthparts. Labrum transverse; anterior margin straight slightly to mark edly concave, or with narrow deep medial notch. Mandibles with dorsolateral margins rounded or ridged. Prothorax. Pronotum as in Figs. 16, 17A-B, and 19A-B. Disc smooth, not tuberculate. Lateral margin distinctly beaded, bead widened around posteriolateral setigerous puncture, or bead very narrow, rather indistinct, not widened posteri or orly. Anteriolateral angles slightly rounded (Figs. 16,17A-B) sharp, subacute (Figs. 19A-B). or Elytra. Hum?rus rounded (cf. Fig. 5A) angulate, apical margin obliquely truncate, slightly sinuate. Basal ridge (cf. Fig. 5A, br) sinuate and broad, or narrow or and straight (Fig. 5B). Striae shallowly impressed, striae unimpressed, their courses marked by sub-surface trabeculae. Intervals flat or tuberculate. as Posteriolateral umbilical setigerous punctures arranged in Figs. 9C-D. Legs. Middle and hind tibiae terete, width hind tibia apically less than half or maximum width of hind femur, middle and hind tibiae distinctly compressed more and widened, width hind tibia distinctly than half width of hind femur. or Male genitalia. Median lobe (Figs. 18A-L) left pleuropic left pleuropic area anopic; various in form; preapical in ventral aspect convex (not flattened) narrow, apical margin evenly rounded, not projected to right. Internal sac without spines. For details, see species group and species treatments, below. Ovipositor. As in Figs. 6B, and 12A-J. Habitat.? The species of Macracanthus occupy principally lowland to lower one montane tropical forest, with species occurring in dry open areas. At least one are species occupies caves, probably facultatively, and several associated with
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some ants of the genus Atta. Adults of at least species are night-active, and have been collected in every month of the year. For details, see the following species treatments. Geographical distribution.? (Fig. 20). The range of subgenus Macracanthus extends in eis-Andean South America from eastern Argentina to northern Colom bia and Venezuela, in the West Indies in the islands of the Greater Antilles, and in Middle America from Panam? to northwestern M?xico.
Phylogenetic relationships.? On the basis of larval features, Vald?s (in press) was more to to postulated that Macracanthus (s. str.) closely related Masoreus than on a Anaulacus (s. str.). But this postulate is hinged shared derived emargination we more of the nasale, which contend is likely to evolve independently in separate lineages than the group of defining features for Anaulacus (s. lat.), which places Anaulacus (s. str.) and Macracanthus (s. str.) in a group apart from Masoreus. In our so more opinion, then, the suite of larval features presented ably by Vald?s is likely to be of value in postulating relationships within Anaulacus than between this genus and the other genera of the Masoreini. But, what then are the relationships of Macracanthus, within Anaulacus (s. lat)? This group shares with Aephnidius and a presently undescribed South African a seems as a subgenus, setiferous prosternai intercoxal process. This promising synapotypy. Other features, such as short antennae (shared with Old World Microus and Anaulacus (s. str.)) and markedly derived pattern of elytral some seem Microsculpture (shared with species of each of those subgenera) less promising as indicators of phylogenetic relationship. The question remains open for the present. are Included species.? The six species o?Macracanthus arrayed in three species groups. See below, for details.
The sericatus species group
are: Recognition.? Characteristic of this species group body color predomi nantly piceous; antennomeres 5-10 approximately quadrate, antenna, backf olded, extended to posterior margin of pronotum; mandibles with dorsal surface shining, mesh pattern longitudinal, microlines shallow, dorsolateral margins rounded, not ridged; pronotum with relatively high values for ratio PL/PWM and HW/PWM (Tables 4B and 5B, respectively), pronotal disc smooth, without paramedian tubercles; elytral hum?rus rounded (Fig. 15), basal ridge relatively broad and sinuate (cf. Fig. 5A); and middle and hind tibiae terete, with width of hind tibia apically less than half maximum width of hind femur. The median lobe of the male genitalia is left pleuropic-anopic. The shaft is slender, not appreciably widened distad the preapical bend. In proportions, (Table 6B and Figs. 18A-F; cf. Figs. 18G L), the preapical part of the shaft is short tomoderately long, and the periostial part is relatively short. Description.? With recognition features noted above, and character states of or as subgenus Macracanthus, restricted amplified follows. Habitus. As in Fig. 16. see For data about size (Standardized Body Length), and proportions, Tables 2A 6B. Color. Somber, body predominantly piceous; pronotum and elytra concolorous or bicolored. Microsculpture and luster. Head capsule: Clypeus with mesh pattern isodiametric, sculpticells slightly convex, surface subopaque. Frons with mesh pattern transverse, sculpticells flat, and microlines finer, surface subiridescent. convex Vertex with mesh pattern longitudinal posteriorly, sculpticells slightly and microlines deeper, surface shining. Pronotum: Disc with mesh pattern trans verse, sculpticells flat, microlines shallow, surface shining to iridescent. Scutel convex. lum: Mesh pattern isodiametric, sculpticells slightly Mouthparts. Labrum with anterior margin shallowly concave, not narrowly notched medially. Man
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dibles dorsolaterally rounded, not ridged. Prothorax. Pronotum as in Figs. 16 and 17A. Disc smooth, not tuberculate. Lateral margin distinctly beaded, bead wid ened around posteriolateral setigerous puncture. Anteriolateral angles slightly rounded. Elytra. As in Fig. 16. Striae very shallow, but evident. Hum?rus (cf. Fig. 5A) rounded. Basal ridge (cf. Fig. 5A, br) sinuate and broad. Striae shallowly impressed. Intervals flat, interval 3 not tuberculate. Legs. Middle and hind tibiae terete, width hind tibia apically less than half maximum width of hind femur. Male genitalia. Median lobe (Figs. 18A-F) left pleuropic or left pleuropic-anopic; shaft with preapical portion short to moderately long. Ovipositor. Gonocoxites 1 and 2 as in Figs. 12G-J. Geographical distribution.? (Fig. 20). The range of the sericatus species group extends throughout eis-Andean South America from northern Argentina, north ward. In Middle America the range of this group extends northward on the Caribbean versant almost to the Tropic of Cancer, in eastern M?xico, and to the islands of the Greater Antilles, in the West Indies. Phylogenetic relationships.? We postulate that the sericatus species group is the + on adelphotaxon of the mcclevei tuberculatus species groups, based the absence from the former of the synapotypic features that characterize the latter pair. Included species.? Three species, treated below, are included in the sericatus species group.
Anaulacus (Macracanthus) sericatus Chaudoir Figs. 5E-F, 12G-J, 16,17A, 18A-B, and 20
Macracanthus sericatus Chaudoir, 1846: 541. TYPE MATERIAL: one female labelled "TYPE" [red paper], putatively the holotype, in Oberth?r-Chaudoir Collection (MNHP), associated with following label originally pinned to the bottom of an insect box: "sericatus/ Chaudoir/ Br?sil/ coll. Falder". Masoreus (Macracanthus) sericatus; Chaudoir, 1876: 24. Masoreus brevicillus Chevrolat, 1863: 189. TYPE MATERIAL: one specimen, HOLOTYPE, labelled: "Masoreus/ brevicillus Chr. an. 1863, p. 189/ Cuba D. Poey" (Hope Museum, Oxford U). NEW SYNONYMY ? Masoreus (Macracanthus) brevicillus; Chaudoir, 1876: 24. Peck, 1974: 24.? Peck et al, 1998: 157, 162. Vald?s, in press. Masoreus (Macracanthus) unicolor Chaudoir, 1876: 23. TYPE MATERIAL: HO LOTYPE, male (without head), in Oberth?r-Chaudoir Collection (MNHP), associated with following label originally pinned to the bottom of an insect ? box: "unicolor/ Chaudoir/ Amazone/ Ega Bates". Bates, 1883: 175. NEW SYNONYMY.
Notes about material and as type synonymy.? The specimen treated holotype of M. sericatus in the Oberth?r-Chaudoir Collection, is without a Head. Next to it is a on a head, mounted point; associated with it is a printed label: "Macracanthus sericatus" in what we take to be Chaudoir's hand. A second female associated with
this putative holotype is labelled "Ste Catharina", presumably Santa Catarina state. one In the original description, Chaudoir notes only specimen, collected at Rio de Janeiro. Thus, the Santa Catarina specimen cannot be the type, nor can it be included in the type series. on a The original description of Masoreus brevicillus is based single specimen, collected by Poey in Cuba. Two females in the Oberth?r-Chaudoir Collection are a (MNHP) associated with label stating "brevicillus Guerin M" and "Cuba". The structural details of these specimens simply confirm that Chaudoir's and taxon Chevrolat's concept of this coincide. They, and additional West Indian material, indicate specific identity with the type material o?Macracanthus sericatus.
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a Associated with the putative holotype of M. unicolor are male and female labelled "Ega", and "Ex Musaeo H. W. Bates 1892". Additionally, the male bears a label "this insect/ crepitates", and the female is labelled "unicolor/ Chaud". These specimens must have been acquired by Oberth?r following his acquisition as of the Chaudoir collection; thus they cannot be regarded type material, though they were collected by Bates, and from the locality where the type of M. unicolor was obtained.
Type locality.? Rio de Janeiro, State of Rio de Janeiro, Brazil. Recognition.? See key to subgenera and species of Neotropical Anaulacus (s. lat.). Additionally, within the sericatus species group, males of A. sericatus are distinguished by details of the median lobe (Figs. 18A-B; cf. Figs. 18C-F): preapical part in left lateral aspect curved slightly dorsally, and in ventral aspect apex broadly rounded. Description.? With recognition features noted above, and character states of or as the sericatus species group, restricted amplified follows. Habitus. As in Fig. 16. see For data about size (Standardized Body Length), and proportions, Tables 2A as 6A. Prothorax. Pronotum in Fig. 16. Male genitalia. Median lobe (Figs. 18A-B) left area narrower pleuropic; basal bend gradual, basal slightly than medial area; basal area more bulb (Fig. 18B) moderately prominent; medial curved ventrally than in most other Anaulacus species, in left lateral aspect (Fig. 18A) curved evenly into periostial part, latter narrowed evenly into preapical part; preapical part moder ately long, curved, in left lateral aspect; in ventral aspect (Fig. 18B) straight, as moderately broad, apex broadly rounded. Ovipositor. Gonocoxite 2 in Figs. 12G-J. Immature stages and life history.?Information about these topics is provided by name Vald?s (in press), under the Macracanthus brevicillus Chevrolat. a Habitat.? This species is ground-living inhabitant of lowland evergreen and over an lower montane tropical forests, altitudinal range of approximately sea m. a level to 1000 It is facultative cavernicole, reported in West Indian studies name under the Macracanthus brevicillus Chevrolat by: Peck (1974: 24, and 1981: 76); Peck et al (1998:157); and Vald?s (in press). It lives also in association with ants of the genus Atta. If crepitation (explosive release of defensive secretions), as noted a common by Bates, is feature of this species, perhaps the explanation of this habit is in the association of the beetles with ants, and the consequent need for added protection from attacks of the latter organisms. Collections of adults at light sources indicate nocturnal flight activity. Geographical distribution.? (Fig. 20). The geographical range of A. sericatus is nearly co-extensive with the range of subgenus Macracanthus, including much of subtropical and tropical eis-Andean South America, most of the Middle American tropics, and the Greater Antilles, in the West Indies. Chorological affinities.? The geographical range of this species overlaps the ranges of its close relatives, A. thoracicus and A. erwini. The latter species and A. are near sericatus sympatric Tambopata, Peru. on Phylogenetic relationships.?Based the less derived mesh pattern and coarser microlines of the pronotal disc, A. sericatus is postulated to be the adelphotaxon of A. thoracicus + A. erwini.
Material examined.?In addition to the type material noted above, we have seen as 101 specimens of A. sericatus, follows. BELIZE. Orange Walk Dist. Rio Bravo nr. Conser. Area, P. Kovarik (FMNH): female, Well Trail, well, flight intercept trap ca. ex in dry creek, 10-16.IV.1996; male, 2 females, La Milpa ruins, Atta debris pile, 18.IV.1996. Toledo Dist. 2males, female, 8 km nw Punta Gorda, (Borland's Cave), zone in loose powdery soil and bat guano, dark of limestone cave, 29 .VI. 1981, W.E. near u-v Steiner (USNM). Female, Blue Creek Village, rain forest, valley creek, light, 10.VI.1981, W.E. Steiner (USNM). BRAZIL. Distrito Federal. Female, Estacao
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Florestal, Cabeca do Vecado, 100 m, 23-27.X.1971, E.G., I. & E.A. Munroe (CNCI). Para Male, female, Santarem (CMNH). Rio de Janeiro. Female, Rio de Janeiro, (CMNH). Rondonia. Ouro Preto d'Oeste, W. Groeneveld, A. Akker (ZMAN): male, 24.VI.1983; female, 3.III.1983. COLOMBIA. Magdelena. Sierra Nevada de near Santa Marta, Municipio Minea, Minea, natural forest, 700 m, 1997, E. Camero (UNCB). COSTA RICA. Cartago. Male, Chirripo, under loose bark, 20.IV.1925, F. Nevermann (USNM). Guanacaste. Male, Est. Santa Rosa, P.N. Sta. Rosa, 300m, 3 12.VI.1992 (INBIO). Limon. Female, Santa Clara, Ramal, Reventaz?n R., under loose bark on rotten wood, 2.VI.1925, F. Nevermann (USNM). Puntarenas. 2 females, Est. Quebrada Bonita, R.B. Carara, 100m, III.1995, R. Guzman (INBIO). Est. Quebrada Bonita, Res. Biol. Carara, 50m, J.C. Saborio (INBIO): 2 males, 3 w females, 11.1993; male, 2 females, VII.1992. San Jose. Male, La Caja, 8 km San Jose, 1934, Schmidt (DEIC). CUBA. Cienfuegos. Female, Soledad, V-VI.1939, Parsons (MCZC). Havana. Male, female, Cueva Mariana, Quivican, 4.IV.1968, F. de Zayas (MCZC). Pinar del Rio. Female, Vinales, San Vicente, 19.VIII.1963 (USNM). Santiago de Cuba. Female, Vista Alegre, at light, 16.VI.1942, C.T. ne Ramsden (USNM). Female, 10 km Caney, Arroyo Grovert, leaf and log litter, 300m, S.Peck (SPCC). ECUADOR. Napo. Female, Yasuni Sei. Sta., 00?40'36"S 76?24'02"W, 220 m, 15.IV.1998, K.W. Will (CUIC). GUATEMALA. Jutiapa. Fe male, San Jose, 16.V.1951, E.S. Ross (CASC). HONDURAS. Yoro. Female, Subirana, e 400 m, Stadelman (MNHP). M?XICO. Campeche. Female, 51.2 km Escarcega, 61 m, 14-15.IV.1966, G.E. Ball, D.R. Whitehead (UASM). Male, 2 km s Bolenchenticul, Grutas de Xtacumbilxunam, 14.VII.1975, J. Reddell, A. Grubbs (UASM). Chiapas. u-v Male, Palenque ruins, light, 122 m, 12.IV.1966, G.E. Ball, D.R. Whitehead n ex (UASM). 5 males, 2 females, Mpio. Cintalapa, 8 km Nuevo Tenochitlan, subter. Atta dump, 13.VII.1991, P.W. Kovarik, T.K. Philips (FMNH). Oaxaca. Male, 2 km w Los Corniles, Cueva de los Corniles, 30.XIL1976, D. McKenzie (JRCC). Female, 6 km s Acatlan, Cueva de las Maravillas, 29.XII.1976, D. McKenzie (JRCC). Quintana Roo. Female, Esperanza, 48.3 km se Chemex, in great cavern, 16.XI.1965, J.G. Edwards (JRCC). Male, Ruinas de Tulum, Cueva de Tulum, 18.VII.1983, J. Reddell (JRCC). 3 females, Felipe Carrillo Puerto, Cenote de Juan Coh, 4.VII.1975, J. Reddell, A. Grubbs, D. McKenzie (JRCC). Veracruz. Female, 4 km w Sontecomapan, 30.5 m, 3&10.IV.1966, G.E. Bail, D.R. Whitehead (UASM). Maie, 33 km ne Catemaco, Los Tuxtlas Biol. Sta., ravine, litter, 1.VII.1983, S. & J. Peck (UASM). Yucatan. Female, in cave, A.S. Pearse (JRCC). Male, 12 km e Catzin, Cenote de San Diego, 5.VII.1975, J. Reddell, A. Grubbs (JRCC). 2 males, female, 7 km sw Oxkutzcab, Actun Loltun, 25-26. VII.1975, J.Reddell, A. Grubbs, D. McKenzie, S. Wiley (JRCC). 3 males, Kaud Cave, 16.VI.1936, A.S. Pearse (UNAM). NICARA GUA. Chontales. Janson (BMNH). PANAMA. Canal Zone. Barro Colorado or Island: female, berlese, debris of field, under bark large stub, 21.1.1959, H. Dybas (FMNH); female, 8.VI.1928, J. Zetek (USNM); female, light traps, 20.V.1978, H. Wolda (USNM). Panam?. 4 males, 3 females, Chilibre, Chilibrillo Cave, bat guano, thinlayer,7.II.1959,H. Dybas (FMNH). Female, Rio Chilibrillobat caves, 29.111.1923, Zetek, Molino, Shannon (USNM). PERU. Madre de Dios. 3 males, female, 15 km ne ex on Puerto Maldonado, fungus log, 200m, 13.VI.1989, J. Ashe, R. Leschen (SEMC). PUERTO RICO. Male, Aguas Buenas, Aguas Buenas Cave, XI.1968, B. s Fenton (CNCI). 3males, 4 females, 35 km Cabo Rojo, Cueva Tuna, berlese guano, 14.VI.1974, S.B. Peck (CNCI). 2 females, Cueva de Carozal, 4.1.1967, S.B. Peck (CNCI). TRINIDAD. Male, Lopinot Valley, Colado Cave, 11.IX.1989, J.P.E.C n Darlington (CNCI). Simia, 8 km Arima, 20.VIII.1969, H. & A. Howden (UASM). 2 n VENEZUELA. Miranda males, Guatopo Nat. Pk., El Lucero, 28 km Altagracia, rotted log litter, 8. VI.1987, S. & J. Peck (CMNC). 2males, 4 females, Puente Bucaral, n Guatopo Nat. Pk., 30 km Altagracia, bat guano berlese, 600 m, 9.VI.1987, S. & J. Peck (CMNC).
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Anaulacus (Macracanthus) thoracicus, new species Figs. 18C-D, and 20
Type material? HOLOTYPE, male, labelled: "BRASILIEN/ Rio Grde.do Sul/ K.E. H?depohl"; "Porto Alegre/ X 1958" [ZSMC]. ALLOTYPE, female, labelled: "BRASILIEN/ Rio Grde.do Sul/ K.E. H?depohl"; "Porto Alegre/ X 1958"; "Sammlung/ Jakob Sturm" [ZSMC]. PARATYPE, female, labelled: "BRAZILGorcovado/ Guanabara,700 m/ Nov.1-7,1963/ Wygodzinsky" [AMNH]. PARATYPE, female, labelled: "Nova Teutonia, Sta/Catarina, BRAZ./ XI: [hand written] :1940/ 300-500 m [handwritten] alt./ Fritz Plaumann leg" [FMNH] Type locality.? The type locality of A. thoracicus is Porto Alegre, State of Rio Grande do Sul, Brazil. Specific epithet.? The word "thoracicus", pertaining to the thorax, in Latin, is a used here as noun in apposition, in allusion to the distinctive pronotum of adults of this species. Recognition.? See key to subgenera and species o? Anaulacus (s. lat.). Adults of A. thoracicus are markedly similar to those of A. erwini, but are larger (Table 2A), and with higher values for the ratio PL/PWM (Table 4A). Additionally, within are subgenus Macracanthus, males of A. thoracicus distinguished by details of the median lobe (Figs. 18C-D; cf. Figs. 18A-B and 18E-F), particularly the marked seen relatively sharp bend in the medial portion, in left lateral aspect. Description.? With recognition features noted above, and character states of or as the sericatus species group, restricted amplified follows. Habitus. Similar to A. sericatus (cf. Fig. 16). For data about size (Standardized Body Length) and see proportions, Tables 2A-6A. Microsculpture. Pronotal disc with mesh pattern at transverse, microlines very fine, hardly visible 50X, close together. Male genita area lia. Median lobe (Figs. 18C-D) left pleuropic; basal bend gradual, basal narrower slightly than medial area; basal bulb (Fig. 18D) moderately prominent; medial part in left lateral aspect (Fig. 18C) curved sharply toward middle (mb); distally widened into periostial part, latter narrowed rather abruptly into preapical area; preapical part relatively short, sloped ventrally, parallel-sided in lateral and ventral aspects, with apex obliquely subtruncate. a Habitat.? This species is probably ground-living inhabitant of the Eastern Atlantic Forest, living, like its relatives, at lower altitudes. Geographical distribution.? (Fig. 20). This species is known only from three localities, two nearly coastal, in southeastern Brazil. Chorological affinities.? The geographical range of A thoracicus is overlapped by the range of A. sericatus, but is widely isolated from the range of its putatively closest relative, A. erwini. on Phylogenetic relationships.? Based similarity in the putatively apotypic on feature of pronotal microsculpture, and the apparent allopatric distribution pattern, this species is the adelphotaxon of A. erwini. seen Material examined.? We have only the type material of this species. For details, see that topic, above.
Anaulacus (Macracanthus) erwini, new species Figs. 18E-F, and 20
Type material? HOLOTYPE, male, labelled: "PERU Dpto. Loreto / Rio Samiria / on Cocha Shinguito/ logs, under bark/ 28-29.VIII.91 28-91"; "T.L.Erwin PERU EXP./ Res. Pacaya-Samiria/ G.E.Ball & D.Shpeley/ collectors 1991" [USNM]. same as ALLOTYPE, female, labelled holotype [USNM]. 3 PARATYPES, sex and as label data follows. Male, same holotype [UASM]. Female, "PERU: Tambopata Prov./ 15 km NE Pto. Maldonado/ 13 July 1989,200 m/ J.Ashe, R.Leschen #507/
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ex:rotting tree" [SEMC]. Female, "PERU: Tambopata Prov./ Madre de Dios Dpto./ 15km NE Puerto"; "Maldonado Reserva/ Cuzco Amaz?nico/ 12?33'S, 69?03'W/ 200m, Plot #Z2E20-21"; "25 June 1989, J. S. Ashe,/R. A. Leschen #400/ ex., berlesate,Palmaceae/ flower fall/logs" [SEMC]. Type locality.?The type locality of A. erwini is Cocha Shinguito, Departamento a Loreto, Peru. Cocha Shinguito is large oxbow lake beside the Rio Samiria, which a is minor tributary of the Rio Amazonas, in the Upper Amazon Basin. a Specific epithet.? The word "erwini" is Latinized genitive eponym, based on surname the of Terry L. Erwin, Department of Entomology, Smithsonian Institu tion, friend and associate, who invited the authors to participate in the expedition some on during which of the specimens (including the holotype), which A. erwini is based, were collected. Recognition.? Adults of this species are similar to those of A. thoracicus, differing in smaller size (Table 2A), and in lower values for the ratio PL/PWM (Table 4A). Additionally, within subgenus Macracanthus, males of A. erwini are distinguished by details of the median lobe (Figs. 18E-F; cf. Figs. 18A-D): in ventral aspect, the medial part is expanded distally, and the preapical part is relatively narrow, with apex narrowly rounded. Description.? With recognition features noted above, and character states of or as the sericatus species group, restricted amplified follows. Habitus. Similar to size A. sericatus (cf. Fig. 16). For data about (Standardized Body Length) and see as proportions, Tables 2A-6 A. Microsculpture. For pronotum, same A. thoracicus. Male genitalia. Median lobe (Figs. 18E-F) left pleuropic-catopic; median lobe in straighter than any other Neotropical species of Anaulacus; basal bend hardly evident basal narrower (Fig. 18E), part slightly than medial part; basal bulb (Fig. 18E-F) only slightly developed; medial part in left lateral aspect (Fig. 18E) curved into evenly periostial part, latter narrowed evenly into preapical part; preapical part rather short, rather narrow, apex obliquely subtruncate in left lateral aspect (Fig. 18E), broadly rounded in ventral aspect (Fig. 18F). Internal sac without spines. Habitat.? The only known specimens of A. erwini were taken in association with fallen trees: from under loose bark, in association with fungus, in lowland riparian tropical evergreen forest. Like most Neotropical species o? Anaulacus, this species is probably ground-inhabiting. Geographical distribution.? (Fig. 20). This species is known only from Amazo nian Peru.
Chorological affinities.? The geographical range of A. erwini, overlapped by that its A. of relative, sericatus, is widely separated from the known range of its putative adelphotaxon, A. thoracicus. Specimens of the former two species were near collected virtually together, Tambopata, Peru. on Phylogenetic relationships.? Based similarity in derived pronotal on microsculpture, and the apparent allopatric distribution pattern, we believe this species is the adelphotaxon of A. thoracicus. Material examined.? We have seen only the type material of A. erwini. For details, see that topic, above.
The mcclevei species group
the features this are same as Being monospecific, of group the those of its single new species, A. mcclevei, species. See below, for details.
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Anaulacus (Macracanthus) mcclevei, new species Figs. 17B, 18G-H, and 20
Type material? HOLOTYPE, male, labelled: "MEXICO: SONORA 17/ km SW ex Moctezuma/ vii-22-80/ Atta nest S.McCleve,P.Jump" [USNM]. ALLOTYPE, sex female, labelled same as holotype [USNM]. 23 PARATYPES, and label data as follows. 2 males, 3 females, same holotype [SMC]. Male, "MEX.VII. 12.62/ Sonora, 1 mi. NW/ Navojoa"; "E.Sleeper,R.An-/ derson,A.Hardy,/ R.Somerby Collr." [CASC]. Male, female, "13.5 mi E Alamos/ Sonora, MEX./ X-29-1972/ DSChandler coir."; "in Atta/ mexicana nest/ det. C. Gaspar" [OSUC]. 4 males, 6 females, "Mexico;Sonora/ 13mSE Alamos/ Oct.301972/ K.Stephan leg." [FSCA]. Male, "Venedio/ SinaloaMex./ VI-10-1918"; "Van Dyke/ Collection" [CASC]. Female, "5 mi.N.Mazatlan/ Sin., MEXICO/ 30.VII.1964/ W.C.McGuffin" [CNC]. Female, "MEXICO: CHIHUAHUA/ Sierra Madre Mts./ La Bufa/ 7-VII-1972/ Derham Giuliani" [CASC]. Female, "MEXICO, Jalisco/ Cihuatlan, Rte./ 110, nr. sea/ level. 1.18.66/ D.R. Whitehead" [UASM]. Male, "Mexico;Nayarit/ San Bias/ July 5 1972/ K.Stephan leg." [FSCA]. Type locality.? The type locality of this species is 17 km. southwest of Moctezuma, State of Sonora, M?xico. a Specific epithet.? The word "mcclevei" is Latinized genitive eponym, based on surname the of Scott McCleve, Douglas, Arizona, amateur coleopterist and specialist on scarabaeid beetles, generous host and field companion of the senior author, and one of the collectors of the type series of A. mcclevei. Recognition.? Similar to adults of the tuberculatus species group in the overall rufous body color, antennomeres 5-10 broader than long, and mandibles with are ridge dorsolaterally, adults of A. mcclevei smaller (Table 2B) with lower values for the ratio PWB/PWM (Table 3B), and higher values for HW/PWM (Table 5B). Further, the anteriolateral angles of the pronotum are more broadly rounded than are those of the tuberculatus species group (Fig. 17B; cf. Figs. 19A-B). The elytral are hum?rus is nearly rectangular. Additionally, males of A. mcclevei distin guished by details of the median lobe (Figs. 18G-H; cf. Figs. 18A-F and 18I-L): preapical part long and slender; medial and prebasal parts similar in width, relatively narrow; and periostial part relatively short (high value for the ratios PAL/MLL and PAL/OML, and relatively low value for the ratio OML/MLL (Table 6B). Description.? With recognition features noted above, and character states of or subgenus Macracanthus, restricted amplified as follows. For data about size (Standardized Body Length), and proportions, see Tables 2A-6B. Color. Somber, body rufous; appendages testaceous. Microsculpture (Figs. 2A-0) and luster. Head capsule: Clypeus with mesh pattern isodiametric, sculpticells slightly convex, surface subopaque. Frons with mesh pattern transverse, sculpticells flat, and microlines finer, surface subiridescent. Vertex with mesh pattern longitudinal convex posteriorly, sculpticells slightly and microlines deeper, surface shining. Mandibles: Dorsal surface shining, mesh pattern longitudinal, microlines shal low. Pronotum: Disc with mesh pattern transverse, sculpticells flat, microlines shallow, surface shining to iridescent. Scutellum: Mesh pattern isodiametric, sculpticells slightly convex, surface subopaque. Mouthparts. Labrum with anterior margin shallowly concave, not narrowly notched medially. Mandibles. Dorsolat eral margins ridged. Pronotum. As in Fig. 17B. Disc smooth, not tuberculate. Lateral margin distinctly beaded, bead widened around posteriolateral setigerous puncture. Anteriolateral angles slightly rounded. Elytra. Hum?rus subangulate. Basal ridge (cf. Fig. 5A, br) sinuate and broad. Striae shallowly impressed. Intervals flat, interval 3 not tuberculate. Legs. Middle and hind tibiae terete, width hind tibia apically less than half maximum width of hind femur. Male genitalia:
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Median lobe (Figs. 18G-H) left pleuropic; basal bend gradual, basal part slightly narrower than medial area; basal bulb (Fig. 18H) moderately prominent; medial part in left lateral aspect (Fig. 18G) curved evenly into periostial part, latter narrowed evenly into preapical part; preapical part relatively very long, straight, parallel-sided in lateral and ventral aspects, apex moderately broadly rounded. Habitat.? This species lives in relatively dry areas, in association with ants of the genus Atta, where adult beetles have been found in the ants' refuse heaps. Geographical distribution.? (Fig. 20). Confined to northwestern M?xico, the range of A. mcclevei extends from Jalisco to northern Sonora, to the west of the Sierra Madre Occidental.
Chorological affinities.? This species is isolated geographically from all other its Neotropical species of Anaulacus, including those of putative adelphotaxon, the tuberculatus species group. on Phylogenetic relationships.? Based its diagnostic features, A. mcclevei is postulated to be the adelphotaxon of the tuberculatus species group. Material examined.? In addition to the type material (see above for details), we seen one have female from Alamos, Sonora, used to obtain a DNA sample (K. Ober, Department of Entomology, University of Arizona).
The tuberculatus species group
Recognition.? Characteristic of this species group are: body color rufous; antennomeres 5-10 wider than long, antenna backfolded not extended to base of pronotum; mandibles dorsally without microlines, dorsolaterally with distinct ridge; pronotum with broad base (Figs. 19A-B), length and maximum width moderately divergent (values for ratio PL/PWM less than 0.53; Table 4B), pronotal more or disc less tuberculate, elytral hum?rus rectangular (Figs. 19A-B), and basal narrow ridge and straight (Fig. 5B); and middle and hind tibiae somewhat compressed, width of hind tibia apically less than half maximum width of hind femur. The median lobe of the male genitalia is left pleuropic-anopic. Distad the is than in preapical bend, the shaft appreciably wider other species of Neotropical Anaulacus. In proportions, (Table 6A-B and Figs. 18I-L), the preapical part of the shaft is relatively short, whereas the periostial part is relatively long. Description.? With recognition features noted above, and character states of restricted or as subgenus Macracanthus, amplified follows. Habitus. As in Figs. 19A-B. For data about size (Standardized Body Length), and proportions, see Tables 2B-6B. Color. Somber, body rufous; elytra concolorous; appendages testa ceous. Microsculpture (Figs. 2N-0) and luster. Head capsule: Clypeus with mesh pattern transverse, microlines evanescent, surface shining. Frons with mesh pattern transverse, sculpticells flat, and microlines finer, surface subiridescent. Pronotum: Disc with mesh pattern irregular, sculpticells flat, microlines shallow, subiridescent. Scutellum: Mesh pattern transverse, sculpticells flat, surface shin Labrum anterior ing. Mouthparts. transverse; margin narrowly notched medially. Mandibles with lateral with narrow dorsally margins raised ridge. Prothorax. as in Pronotum Figs. 19A-B. Disc with pair of paramedian tubercles in posterior 1 / 3. Lateral rather margin narrowly, indistinctly beaded through length, bead not widened in of vicinity posteriolateral setigerous puncture. Anteriolateral angles sharp, subacute. elytra (Figs. 5B, 19A-B). Hum?rus angulate; apical margin ob not Basal narrow liquely truncate, sinuate. ridge (Figs. 5B, br) straight and (Fig. courses 5B). Striae unimpressed, their marked by sub-surface trabeculae; Intervals flat or tuberculate (Fig. 19A). Male genitalia. Median lobe (Figs. 18I-L) left shaft with area pleuropic-anopic; preapical portion very short; preapical in ventral convex aspect (not flattened), narrow, apical margin evenly rounded, not pro to area jected right; periostial long. Female genitalia. Ovipositor and internal
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genitalia as in Fig. 6B. Habitat.? The members of the tuberculatus species group are residents of lowland tropical forest, both evidently associates of ants of the genus Atta. Geographical distribution.? (Fig. 20). The range of the tuberculatus species group extends in eis-Andean South America from southeastern Brazil to approxi mately central Venezuela and in Middle America to Belize, on the Caribbean versant.
Phylogenetic relationships.? This species group is the putative adelphotaxon of A. mcclevei. are Included species.?Included in the tuberculatus species group two species: A. pittieri Gahan, and A. tuberculatus Chaudoir.
Anaulacus (Macracanthus) pittieri Gahan Figs. 6B, 18I-J, 19A, and 20
Leptotomus pittieri Gahan, 1894: 116. TYPE MATERIAL: two specimens, as follows. HOLOTYPE female, labelled: "Type" [circular, ringed with red]; "Costa Rica/ 93-69/ 711"; "Leptotomus/ pittieri/ Gahan/ Type"; "Lecto type [female symbol]/ Leptotomus/ pittieri Gahan/ By Erwin 174" (BMNH). PARATYPE male, labelled: "PARA"; "LECTOTYPE" [red paper]; "Costa Rica/ 92-48"; "Leptotomus/ pittieri/ Gahan/ Cotype" [handwritten]; "Brit ish/ Museum"; "Van Emden/ vidit 1935"; "Aephnidius/ pittieri/ Gahan/ PARALECTOTYPE/ det George E. Ball 1990" (MNHP). van Ophryognathus pittieri; Emden, 1949: 889.
Notes about type material.? Because both of the specimens noted above have handwritten "type" labels, it seems reasonable to assume that Gahan had both at hand when he described this species, though it is unclear from the original was so. description if this Because the specimen in The Natural History Museum as seems is labelled "type", evidently by Gahan, it reasonable to regard it as the one as holotype, with the in the Paris Museum, labelled "Cotype", being a paratype, previous opinions to the contrary notwithstanding. Note about synonymy.? Lorenz (1998:427) listed the name Macracanthus pittieri as a junior synonym of M. t. tuberculatus. Although M. tuberculatus and M. pittieri are we see no closely related, basis for combining them in one species. Type locality.? As determined from the original description (Gahan, 1894:116) the type locality is Buenos Aires, Puntarenas Province, Costa Rica. In to Recognition.? addition only slightly developed paramedial tubercles of the pronotum and discal elytral intervals smooth, not tuberculate, within the are tuberculatus species group, males of A. pittieri distinguished from males of O. tuberculatus by details of the male genitalia (Figs. 18I-J; cf. Figs 18K-L): in left lateral narrower aspect, prebasal bend (pb) gradual, and preapical part (pa) and slightly acute or longer, with apex very narrowly rounded. Description.? With recognition features noted above, and character states of the tuberculatus or as species group, restricted amplified follows. Habitus. As in Fig. 19A. For data about size (Standardized Body Length), and proportions, see Tables as in 2A-6A. Prothorax. Pronotum Fig. 19A. Elytra. As in Fig. 19A. Male genitalia. Median lobe (Figs. 18I-J) left pleuropic-anopic; prebasal bend in left lateral aspect curved but not area (Fig. 181, pb) abruptly angulate (cf. Fig. 18K), basal markedly narrower than medial area; basal bulb moderately prominent (Fig. 18J); medial area thick, curved evenly into periostial area; periostial portion narrowed mark edly toward preapical portion, preapical portion rather short, narrowed to acute Internal sac apex. without spines. Ovipositor and internal female genitalia. As in Fig 6B.
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Habitat.? Data are available for only one locality, which is located in lowland m. was a tropical evergreen forest, at 200 That specimen taken in refuse pile of an Atta nest.
Geographical distribution.? (Fig. 20). This species is known only from Costa Rica and Belize, in Middle America. Chorological affinities.? The geographical ranges of A. pittieri and its putative are close relative, A. tuberculatus, complementary, and non-overlapping. are Phylogenetic relationships.? This species and A. tuberculatus postulated to numerous are be adelphotaxa, based on the shared derived features that charac teristic of the tuberculatus species group. Material examined.? In addition to the type material, we have seen three specimens of A. pittieri, as follows. BELIZE. Orange Walk Dist. Female, Rio Bravo Conservation Area, ca. La Milpa ruins, ex Atta debris pile, 18.IV.1996, P. Kovarik (FMNH). COSTA RICA. Female (USNM);male (BMNH).
Anaulacus (Macracanthus) tuberculatus Chaudoir Figs. 2J-0, 5B, 18K-L, 19B, and 20
Ophryognathus tuberculatus Chaudoir, 1876: 26. TYPE MATERIAL: one authen tic Chaudoir specimen, the holotype, in Oberth?r-Chaudoir Collection (MNHP), associated with following label originally pinned to the bottom of an insect box: "tuberculatus/ Chaudoir/ Colombie/ Valencia/ Moritz/ Mus. de Berlin". The HOLOTYPE female is labelled: "Holo-/ type" [circu lar, ringed with red]; "Ex Musaeo/ Chaudoir" [red print]; "tuberculatus van ? Ch?. I type/ Emden det. 1935" [handwritten], van Emden, 1949:889. Macracanthus (Ophryognathus) pittieri; Lorenz, 1998: 427. Ophryognathus tuberculatus brasilianus van Emden, 1949: 889. TYPE MATE RIAL: nine specimens, in BMNH, MNHP, and coll. G. Nick. Eight speci mens as seen, follows. HOLOTYPE male, labelled: "Type" [circular; ringed with red]; "Mend?s b./ Rio de Janeiro/ H. Eidmann"; "b. Atta 6-dens/ aus alter Pilz-/ kammerern/ 27.9.1933 (BMNH). PARATYPE female, labelled: "Para-/ type" [circular; ringed with yellow]; "Mend?s/ Rio de Janeiro/ H. - Eidmann" [handwritten]; "f. Best 6 1935/ von Eidmann"; "brasilianus/ Paratypus" [handwritten; red paper]; "Ophryognathus/ tuberculatus/ brasilianus/ Emd./ E.B. Britton/ det 9.2.1937 (BMNH). PARATYPE fe male, labelled: "Mend?s/ e. d. Rio/ Brasilien/ 28.IX.1933"; "f. Best. 7.1933/ von Eidmann" (MNHP). Two PARATYPES, similarly labelled to preced ing, male and female, dated "27.IX.1933" and "28.IX.1933", respectively (BMNH). PARATYPE female, similarly labelled to preceding, but without date (BMNH). PARATYPE female, labelled "Paraguay Villarica 3.36 F. Schade leg." (BMNH). PARATYPE female, labelled: "Sn Paulo/ Br?sil"; "E Musaeo/ V. Mayet/ 1909 (MNHP). New synonymy.
Notes about type material.? Associated with the putative holotype of O. tuberculatus in the Oberth?r-Chaudoir Collection is a female labelled "Bahia/ van as a Lacerda", determined by Emden (in 1935) member of O. t. brasilianus. source Although Chaudoir did not record the of the material on which the name O. tuberculatus is one can infer based, from the wording of the original description that it was on a based single specimen. Further, because the description fits the character states of O. t. tuberculatus, rather than those of O. t. brasilianus. one can as a make the further inference that the specimen labelled holotype must be that, van conclusion reached previously by Emden. van as Two additional specimens, both identified by Emden O. t. brasilianus are were associated with the authentic specimens of O. tuberculatus, but both acquired
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not as by Oberth?r following the death of Chaudoir. These have been labelled are as paratypic, but they noted such in the original description. Note about synonymy.? See this topic for M. pittieri, above. a area Type localities.?For O. t. tuberculatus, only type is indicated: "Valencia", van which according to Emden (1949:889) is Venezuela. Because of taxonomically seems amore significant geographical variation in this species, it desirable to have specific locality. We choose arbitrarily Ciudad Bolivar, State of Bolivar, Venezu ela. For O. t. brasilianus, the type locality isMend?s, State of Rio de Janeiro, Brazil, as determined from the original description of this subspecies, and from the labelling associated with the holotype. Recognition.?In addition to pronounced paramedial tubercles of the pronotum and tuberculate elytral intervals 3 and 5, within the tuberculatus species group, are males of A. tuberculatus distinguished from males of A. pittieri by details of the male genitalia (Figs. 18K-L; cf. Figs 18I-J): in left lateral aspect, prebasal bend short more and sharp, and preapical part broader and shorter, with apex broadly rounded.
Description.? With recognition features noted above, and character states of or as the tuberculatus species group, restricted amplified follows. Habitus. As in Fig see 19B. For data about size (Standardized Body Length), and proportions, Tables 2A-6A. Microsculpture. Elytral microsculpture as in Figs. 2J-0. Elytra. Hum?rus and basal ridge as in Fig. 5B. Male genitalia. Median lobe (Figs. 18K-L) left pleuropic-anopic; prebasal bend in left lateral aspect (Fig. 18K; cf. Fig. 181, pb) area narrower abrupt, angulate, basal markedly than medial area; basal bulb area moderately prominent (Fig. 18L); medial thick, extended evenly into periostial area area area; periostial portion narrowed markedly toward preapical preapical rather short, narrowed slightly to broadly rounded apex. Internal sac without spines. Variation.? Van Emden (1949: 889-890) recognized two subspecies (see syn on onymy, above), based distinctions in pronotal form (especially development of the anteriolateral angles), development of anterioparamedial tubercles, and form of the elytral apices. But in the material seen by us, these features seem to vary in an inconsistent manner relative to each other. Accordingly, we elect not to recognize subspecies. a Habitat.? This species is ground-living inhabitant of lowland and lower an montane tropical evergreen forest, with altitudinal range approximately from m. sea level to 1000 Probably adults are active year-round, though they have been collected only during five of the 12 months. Geographical distribution.? (Fig. 20). The geographical range of A. tuberculatus probably extends through most of the lowlands of eis-Andean South America. Chorological affinities.? The geographical ranges of A. tuberculatus and its are putative close relative, A. pittieri, complementary, and non-overlapping. are Phylogenetic relationships.? This species and A. pittieri postulated to be on numerous are adelphotaxa, based the shared derived features that character istic of the tuberculatus species group. Material examined.? In addition to the types, noted above, we have seen 17 specimens of A tuberculatus, as follows. BOLIVIA. Sara. Female, Steinbach (CMNH). Santa Cruz. Male, Asuncion, 500 m, XI.1963 (MNHP). BRAZIL. Amazonas Female, Tefe, 8.XII.1919, H. Parish (CASC). Distrito Federal. Female, Ilha do Governador, b. Atta sexdens, 16.XI.1955, M. Alvarenga (QBUM). Pernambuco. Female, Pery-Pery (=Pirituba), 11.XII.1892, Grounelle (MNHP). Rio de Janeiro. 3 males, female, Mendes, 29.IX.1933, Eidmann (NHMB). S?o Paulo. Female, Itu, 11.1959, U. Martins (MZSP). FRENCH GUIANA. Female, Maroni River, W. Schaus (USNM). GUIANA. Male, Bartica Demerara, R.J. Crew (USNM). SURINAM.
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D.C. 6 Lelydorp, in rubbish pit of Atta sexdens nest, Geijskes (RMNH): male, 16.XII.1938; 2 males, female, 19.IX.1938. VENEZUELA. Bolivar. Male, Ciudad Bolivar, 28.VIII.1898, E.A. Klages (CUIC).
Ecological Considerations
This section summarizes available information about natural history of the All the are most of them Neotropical species of Anaulacus. of species geophiles, sea level and 500 m. Most are inhabitants of living at altitudes between tropical some in and in either man evergreen forest, though live semi-tropical forests, or in areas. Adults of of disturbed situations (mango plantations), dry, open many as At least adults of A. the species fly at night, shown by light-trapping. piceolus as seem to be diurnally active, well. are in some with Most of the species of subgenus Macracanthus associated way is a also. ants of the genus Atta. The species A. (M.) sericatus facultative cavernicole, were associated with fallen The type specimens of A. (M.) erwini trees, perhaps of these kinds are not for the other two only facultatively. Associations reported subgenera, Aephnidius and Apostolus. for life seem to Among the species of Anaulacus, requirements overlap broadly, co-occurrence in the same areas Co as indicated by of several species (Table 7). for and occurrence provides the potential competition genetic exchange among either of these in Anaulacus. related species. No evidence is available for phenomena in Thus, co-occurrence is evidence of reproductive isolation this group.
Geographical history of Anaulacus
on a Based previous analysis of Neotropical biogeographical history (Shpeley & Ball 1993: 157-169) which featured the distribution of the Neotropical taxa of we an that follows a of the Coptodera Dejean, offer evolutionary scenario summary of the taxa. and 20 illustrate the facts of geographical distribution Figures 13,14 of Anaulacus. Maximum known geographical ranges of the Neotropical species both north and and with most diversity is toward the equator, decreasing south, in few latitudinal zones 8 and Of the 15 species represented relatively (Tables 9). nine are to the The extant species of Anaulacus, the ranges of confined tropics. one north of ranges of five species extend beyond the tropics: (A. mcclevei, Fig. 20) The the tropic of Cancer, only; four, south of the tropic of Capricorn, only. range known from a is of only one species (A. bonariensis, Fig. 14), single specimen, the completely outside of the tropics (to south). Anaulacus is considered as well continen The geographical distribution of by are tal occurrence. Of the 15 species, 10 precinctive continentally, i.e., known only from either the South American continent or from the Middle American portion of North America, and most are still more localized within each continent (Table over most of 10). The ranges of five species extend parts of both continents, with In more the area being on the South American continent (Table 11). detail, many to are taxa postulated be closely related distributed vicariously (Table 12). in we that Because Anaulacus is basically Gondwanan distribution, postulate the ancestral stocks of the extant lineages were part of the original South American biota, when the South American continent separated from Africa during Mesozoic time. We are not prepared to deal with the question of how many stocks actually were Gondwanan, with one vicar in Africa and another in South America. one and Certainly, Aephnidius must have been of these. Possibly Apostolus Macracanthus also had African vicars, but we have no evidence for that suggestion. occurred in the South American stocks in the Probably substantial differentiation of the latter part of the Mesozoic Period. Because of the marked range overlap
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no subgenera, we have geographical clues about this early differentiation. an Probably fairly early in the Tertiary period, early stock of Anaulacus (possibly amember of subgenus Aephnidius) reached the Middle American part of North America, crossing a sea barrier to do so (connection between what are now the North American and South American continents was not achieved until the
early Pliocene epoch- [Stehli and Webb 1985: 11]). This early-dispersing stock reached the West Indies, as indicated by Miocene age fossils discovered there, on to we the island of Hispaniola. Being unable relate these fossils to the extant fauna, more offer nothing about their origin and relationships. Probably somewhat later, one or more now extant lineages, represented by species, entered Middle America, or either crossing narrow water gaps, over recently, emerged land that formed the intercontinental connection.
Following entry into Middle America, these Neotropical lineages o? Anaulacus, each with its geographically isolated pair of vicars, differentiated. Further differ entiation of these vicars occurred on each continent, followed by dispersal and some range interp?n?tration of the resulting adelphotaxa. We expect that intracontinental differentiation was the result of interruption of range continuity of beetle species through interruption of habitat continuity. Present intercontinen tal range continuity of five extant species probably is the result of dispersal that a occurred with development of land connection between continental South America and North America.
As indicated above and in conclusion, the distribution pattern of Anaulacus is readily interpretable using the biogeographical paradigm involving Tertiary age geology, Tertiary and Quatrnary age climatic change, and refuge theory (Prance, 1982, and references therein; Vanzolini and Heyer, 1988, and references therein; Savage, 1982).
Concluding Remarks
Inmany respects, this study has been frustrating. Material, overall, was scarce. sense as as we To make taxonomic of the group, small it is, felt compelled to a propose number of monospecific taxa of supraspecific rank, a less than optimal practice in hierarchical classification.The available characters seemed too few and a their variation patterns too discordant to make worthwhile comprehensive phylogenetic analysis, and the absence of information about way of life and shortage of details about habitats occupied made less than useful interpretation of an the characters in evolutionary context. Because of the paucity of geographical net a data, the of evidence for biogeographic analysis had very large mesh. In spite of these defects, this paper offers ameasure of progress in understand ing the Neotropical taxa included in Anaulacus. It is possible now to at least a associate specimens with readily diagnosable taxonomic units, and framework is available for interpreting additional information, as it becomes available. We this is to serve us hope framework sufficiently robust and anyone else who might be interested in seeking understanding of the taxa of Anaulacus that inhabit not only the Neotropics, but also the tropics and warm temperate areas of the Eastern Hemisphere. ACKNOWLEDGMENTS
our to are We express gratitude the following, many of whom close personal friends, and have been, for many years. We are indebted to the curators and other individuals noted above, in the "Materials" section of the paper, for loan of the specimens in their care that were our vital to this study. We acknowledge especially friend, James K. Ryan (Edmonton,
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a an Alberta), for his thoughtful purchase, in jewelry store in New York City, of a amber-preserved carabid from the Dominican Republic that proved to be specimen of Anaulacus, part of which is illustrated in Fig. 20, above. Further, with we are regard to fossil material, thankful for the help that George O. Poinar, Jr. (Oregon State University, Corvallis, Oregon) provided with alacrity and grace, and for the advice and assistance provided by Robert E. Woodruff (Department of Plant Industry, Gainesville, Florida). We thank Terry L. Erwin (USNM) for the privilege of joining his Pacaya use Samiria entomological excursion, in 1991, and for permission to study and his substantial and important Middle American masoreine material. James Stephen Ashe and Robert W. Brooks (SEMC), in the course of their South American field us some work, obtained and provided with fine masoreines. was Special assistance received from George C. McGavin and Darrell Mann (OXUM), who provided us with information about type material of Masoreus brevicillus Chevrolat. Robert L. Davidson (CMNH) and Yves Bousquet (CNCI) some provided helpful advice regarding thorny nomenclatural problems that we encountered.
Although it is possible to obtain much of the working material from museums, on loan, the study of some type specimens in situ is desirable. This is especially true of the Chaudoir types (MNHP). During his stay in Paris to carry out said study, the was senior author made welcome in the Laboratoire d'Entomologie by Thierry Deuve and Jean J.Menier. The cordiality of these colleagues is appreciated very much. are a Illustrations of high quality valuable, if not essential, part of taxonomic we papers. For their contributions, note the following close associates: Diane Hollingdale, who made the line drawings of pronota; George D. Braybrook and Edith Schwaldt (Departmental of Earth and Atmospheric Sciences), skilled opera was tors of the stereoelectron microscope which used to obtain photographs of mouthparts, Ovipositor, s, and microsculpture; and John S. Scott, who prepared the habitus photographs. a a Careful reviews of substantial portion of previous draft of the final manuscript were made by our colleagues Yves Bousquet (CNCI) and Douglas A. Craig (University of Alberta). We are grateful for their considerable efforts on our behalf. museum Funds for limited field work, study, and page charges were met with grant OGP 1399 to the senior author, from the Natural Sciences and Engineering Research Council of Canada.
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Table 1. Checklist of the Western Hemisphere Species of the Genus Anaulacus MacLeay.
new Subgenus Apostolus, subgenus A. batesi Chaudoir Subgenus Aephnidius MacLeay piceolus species group A. piceolus Chaudoir ampliusculus species group A. ampliusculus Chaudoir A. bonariensis Chaudoir A. ciliatus Mutchler new A. humeralis, species. exiguus species group new A. exiguus, species new A. whiteheadi, species new A. ashei, species new subgenus Macracanthus Chaudoir, combination sericatus species group (= Macracanthus (sensu stricto)) A. sericatus Chaudoir new A. thoracicus, species new A. erwini, species mcclevei species group new A. mcclevei, species tuberculatus species group (=Ophryognathus Chaudoir) A. pittieri Gahan A. tuberculatus Chaudoir
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Table 2A. Data about variation in SBL (mm) among the New World species of Anaulacus Macleay.
Name of Taxon N Range Mean
MALES
A. (Apostolus) batest (Chd.) - Tambopata, Peru 3 5.68 6.04 5.91 Buena Vista Bolivia 6.11-2 6.21 6.16 unlabelled (USNM) 5.66- 2 5.70 5.68 - A. (Aephnidius) piceolus (Chd). 3.14 5 3.42 3.31 - A. (Aephnidius) ampliusculus (Chd). 3 3.72 4.08 3.85 ? ? A. (Aephnidius) bonariensis (Chd). - A. (Aephnidius) ciliatus (Mutch.) 2.98 8 3.56 3.22 n. - A. (Aephnidius) humeralis, sp. 3.20 2 3.36 3.28 n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 3 2.51-2.54 2.53 Iquitos, Peru 1 2.48 Parque Nac. Chaco, Argentina 4 2.34-2.59 2.50 A. (Aephnidius) whiteheadi, n. sp. 2 2.48-2.58 2.53 A. (Aephnidius) ashei, n. sp. - Tambopata, Peru 4 2.46 2.86 2.70 - A. (Macracanthus) sericatus (Chd.) 4.523 4.88 4.75 A. (Macracanthus) thoracicus, n. sp. 4.802 A. (Macracanthus) erwini, n. sp. Pacaya-Samiria, Peru 23.36- 3.42 3.39 - A. (Macracanthus) mcclevei, n. sp. 33.36 4.28 3.96 A. (Macracanthus) pittieri (Gahan) 5.16 1 - A. (Macracanthus) tuberculatus (Chd.) 3 4.24 4.48 4.37 FEMALES
A. (Apostolus) batest (Chd.) Tambopata, Peru 1 6.44 Buena Vista, Boliva 1 6.43 - A. (Aephnidius) piceolus (Chd.) 5 3.30 3.70 3.55 - A. (Aephnidius) ampliusculus (Chd.) 3 3.84 3.96 3.91 A. (Aephnidius) bonariensis (Chd.) 1 3.84 - A. (Aephnidius) ciliatus (Mutch.) 8 3.20 3.76 3.41 ? A. (Aephnidius) humeralis, n. sp. n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 4 2.53- 2.60 2.57 Iquitos, Peru 3 2.53-2.61 2.58 Parque Nac. Chaco, Argentina 5 2.53- 2.69 2.63 ? A. (Aephnidius) whiteheadi, n. sp. A. (Aephnidius) ashei, n. sp Tambopata, Peru 6 2.58- 2.74 2.69 Brazil 2 2.62- 2.72 2.67 - A. (Macracanthus) mcclevei, n. sp. 3 4.00 4.32 4.16 - A. (Macracanthus) sericatus (Chd.) 3 4.68 5.04 4.85 A. (Macracanthus) thoracicus, n. sp. 1 4.52 A. (Macracanthus) erwini, n. sp. Tambopata, Peru 3 3.42- 3.78 3.65 - A. (Macracanthus) pittieri (Gahan) 2 4.84 5.36 5.10 - A. (Macracanthus) tuberculatus (Chd.) 3 4.48 5.16 4.76
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Table 2B. Summary by genus, subgenus and species group of data about variation in SBL (mm) among the New World species of Anaulacus Macleay.
Name of Taxon Range of Values
Genus Anaulacus Macleay Males 2.34 - 6.21 Females 2.53 - 6.44
Subgenus Apostolus, new subgenus Males 5.68 - 6.21 - Females 6.43 6.44
Subgenus Aephnidius Macleay - Males 2.34 4.08 - Females 2.53 3.96
piceolus species group - Males 3.14 3.42 - Females 3.30 3.70
ciliatus species group - Males 3.20 4.08 - Females 3.20 3.96 exiguus species group - Males 2.34 2.86 Females 2.53 - 2.74
Subgenus Macracanthus Chaudoir - Males 3.28 5.16 - Females 3.42 5.36 sericatus species group - Males 3.28 4.88 - Females 3.42 5.04
mcclevei species group Males 3.36 - 4.28 - Females 4.00 4.32 tuberculatus species group Males 4.24 - 5.16 - Females 4.48 5.36
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Table 3A. Data about variation in PWB/PWM among the New World species of Anaulacus Macleay.
Name of Taxon N Range Mean
MALES A. (Apostolus) batest (Chd.) - Tambopata, Peru 3 0.828 0.869 0.847 - Buena Vista, Bolivia 2 0.800 0.843 0.822 - Unlabelled (USNM) 2 0.845 0.867 0.856 - A. (Aephnidius) piceolus (Chd.) 5 0.828 0.879 0.866 - A. (Aephnidius) ampliusculus (Chd.) 3 0.816 0.865 0.837 ? ? A. (Aephnidius) bonariensis (Chd.) - A. (Aephnidius) ciliatus (Mutch.) 8 0.829 0.867 0.847 n. - A. (Aephnidius) humeralis, sp. 2 0.781 0.800 0.791 n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 3 0.840-0.852 0.844 Iquitos, Peru 1 0.820 - Parque Nac. Chaco, Argentina 4 0.828 0.860 0.846 - A. (Aephnidius) whiteheadi, n. sp. 2 0.774 0.803 0.788 - A. (Aephnidius) ashei, n. sp. 4 0.828 0.857 0.840 - A. (Macracanthus) sericatus (Chd.) 3 0.927 0.945 0.937 A. (Macracanthus) thoracicus, n. sp. 0.9632 A. (Macracanthus) erwini, n. sp. - Pacaya-Samiria, Peru 2 0.960 0.980 0.970 - A. (Macracanthus) mcclevei, n. sp. 3 0.920 0.950 0.935 A. (Macracanthus) pittieri (Gahan) 0.9501 - A. (Macracanthus) tuberculatus (Chd.) 3 0.943 0.961 0.949
FEMALES
A. (Apostolus) batest (Chd.) Tambopata, Peru 1 0.833 Buena Vista, Bolivia 0.857 1 - A. (Aephnidius) piceolus (Chd.) 5 0.844 0.886 0.865 - A. (Aephnidius) ampliusculus (Chd.) 3 0.825 0.846 0.839 A. (Aephnidius) bonariensis (Chd.) 0.9251 - A. (Aephnidius) ciliatus (Mutch.) 8 0.833 0.871 0.850 ? ? ? A. (Aephnidius) humeralis, n. sp. n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 4 0.808-0.830 0.818 Iquitos, Peru 3 0.817-0.825 0.821 - Parque Nac. Chaco, Argentina 5 0.848 0.885 0.863 ? ?? A. (Aephnidius) whiteheadi, n. sp. A. (Aephnidius) ashei, n. sp. - Tambopata, Peru 6 0.821 0.858 0.833 2Brazil 0.800-0.828 0.814 - A. (Macracanthus) sericatus (Chd.) 3 0.943 0.948 0.945 A. (Macracanthus) thoracicus, n. sp. 0.9581 A. (Macracanthus) erwini, n. sp. - Tambopata, Peru 3 0.943 0.964 0.956 - A. (Macracanthus) mcclevei, n. sp. 3 0.920 0.939 0.932 - A. (Macracanthus) pittieri (Gahan) 2 0.935 0.947 0.941 - A. (Macracanthus) tuberculatus (Chd.) 3 0.944 0.949 0.946
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Table 3B. Summary by genus, subgenus and species group of data about variation in PWB/PWM among the New World species o? Anaulacus Macleay.
Name of Taxon Range of Values
Genus Anaulacus Macleay - Males 0.78 0.98 - Females 0.82 0.95 new Subgenus Apostolus, subgenus - Males 0.83 0.87 - Females 0.833 0.857
Subgenus Aephnidius Macleay - Males 0.78 0.88 - Females 0.80 0.92
piceolus species group - Males 0.83 0.88 - Females 0.84 0.89
ciliatus species group - Males 0.78 0.87 - Females 0.82 0.92
exiguus species group - Males 0.82 0.86 - Females 0.80 0.88
Subgenus Macracanthus Chaudoir - Males 0.92 0.98 - Females 0.92 0.96
sericatus species group - Males 0.93 0.98 - Females 0.94 0.96
mcclevei species group - Males 0.92 0.95 - Females 0.92 0.94 tuberculatus species group - Males 0.94 0.96 - Females 0.94 0.95
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Table 4A. Data about variation in PL/PWM among the New World species of Anaulacus Macleay.
Name of Taxon N Range Mean
MALES A. (Apostolus) batest (Chd.) 3 0.578-0.594 0.587 Tambopata, Peru - Buena Vista, Bolivia 2 0.566 0.575 0.570 - Unlabelled (USNM) 2 0.600 0.620 0.610 - A. (Aephnidius) piceolus (Chd.) 5 0.606 0.655 0.627 - A. (Aephnidius) ampliusculus (Chd.) 3 0.579 0.595 0.586 ? ? ? A. (Aephnidius) bonariensis (Chd.) 8 - A. (Aephnidius) ciliatus (Mutch.) 0.588 0.633 0.608 n. - A. (Aephnidius) humeralis, sp. 2 0.594 0.600 0.597 n. A. (Aephnidius) exiguus, sp. 3 0.640 Tambopata, Peru 0.635-0.645 1 Iquitos, Peru 0.606 4 - Parque Nac. Chaco, Argentina 0.625 0.635 0.630 - n. 2 0.645 0.640 A. (Aephnidius) whiteheadi, sp. 0.636 A. (Aephnidius) ashei, n. sp. 4 - Tambopata, Peru 0.621 0.643 0.632 - A. (Macracanthus) sericatus (Chd.) 3 0.545 0.580 0.557 - A. (Macracanthus) thoracicus, n. sp. 2 0.556 0.574 0.565 A. (Macracanthus) erwini, n. sp. 2 - Pacaya- Samiria, Peru 0.540 0.550 0.545 - A. (Macracanthus) mcclevei, n. sp. 3 0.520 0.575 0.549 A. (Macracanthus) pittieri (Gahan) 0.5001 - A. (Macracanthus) tuberculatus (Chd.) 3 0.500 0.529 0.513
FEMALES A. (Apostolus) batest (Chd.) 1 Tambopata, Peru 0.591 Buena Vista, Bolivia 0.583 1 - A. (Aephnidius) piceolus (Chd.) 5 0.611 0.629 0.620 - A. (Aephnidius) ampliusculus (Chd.) 3 0.564 0.590 0.576 A. (Aephnidius) bonariensis (Chd.) 0.6001 - A. (Aephnidius) ciliatus (Mutch.) 8 0.583 0.618 0.607 n. ? ? ? A. (Aephnidius) humeralis, sp. n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 4 0.593-0.625 0.614 Iquitos, Peru 3 0.635-0.669 0.648 - Parque Nac. Chaco, Argentina 5 0.612 0.639 0.624 n. ? ? ? A. (Aephnidius) whiteheadi, sp. n. A. (Aephnidius) ashei, sp. Tambopata, Peru 6 0.606-0.652 0.630 2Brazil 0.614-0.631 0.622 - A. (Macracanthus) sericatus (Chd.) 3 0.517 0.556 0.540 A. (Macracanthus) thoracicus, n. sp. 0.5831 A. (Macracanthus) erwini, n. sp. Tambopata, Peru 3 0.530-0.547 0.537 - A. (Macracanthus) mcclevei, n. sp. 3 0.511 0.551 0.527 - A. (Macracanthus) pittieri (Gahan) 2 0.491 0.500 0.496 - A. (Macracanthus) tuberculatus (Chd.) 3 0.500 0.509 0.506
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Table 4B. Summary by genus, subgenus and species group of data about in variation PL/PWM among the New World species of Anaulacus Macleay.
Name of Taxon Range of Values
Genus Anaulacus Macleay Males 0.49 - 0.66 - Females 0.50 0.67
Subgenus Apostolus, new subgenus - Males 0.58 0.60 Females 0.59
Subgenus Aephnidius Macleay Males 0.58 - 0.64 Females 0.58 - 0.67
piceolus species group - Males 0.61 0.66 - Females 0.61 0.63
ciliatus species group - Males 0.58 0.64 - Females 0.57 0.68
exiguus species group - Males 0.61 0.64 Females 0.60 - 0.67
Subgenus Macracanthus Chaudoir - Males 0.50 0.58 - Females 0.51 0.58 sericatus species group - Males 0.54 0.58 - Females 0.52 0.58
mcclevei species group - Males 0.52 0.58 - Females 0.51 0.55 tuberculatus species group - Males 0.50 0.53 Females 0.49 - 0.51
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Table 5A. Data about variation in HW/PWM among the New World species of Anaulacus Macleay.
Name of Taxon NRange Mean
MALES A. (Apostolus) batest (Chd.) Tambopata, Peru 3 0.639-0.656 0.650 - Buena Vista, Bolivia 2 0.650 0.662 0.656 - Unlabelled (USNM) 2 0.653 0.676 0.664 - A. (Aephnidius) piceolus (Chd.) 5 0.710 0.727 0.721 - A. (Aephnidius) ampliusculus (Chd.) 3 0.683 0.703 0.690 ? ?? A. (Aephnidius) bonariensis (Chd.) - A. (Aephnidius) ciliatus (Mutch.) 8 0.714 0.767 0.750 n. - A. (Aephnidius) humeralis, sp. 2 0.719 0.733 0.726 n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 3 0.698-0.721 0.713 Iquitos, Peru 1 0.688 - Parque Nac. Chaco, Argentina 4 0.688 0.719 0.708 n. - A. (Aephnidius) whiteheadi, sp. 2 0.697 0.725 0.711 n. A. (Aephnidius) ashei, sp. Tambopata, Peru 4 0.655-0.680 0.671 - A. (Macracanthus) sericatus (Chd.) 3 0.636 0.655 0.644 A. (Macracanthus) thoracicus, n. sp. 0.6302 A. (Macracanthus) erwini, n. sp. - Pacaya-Samiria, Peru 2 0.632 0.640 0.636 - A. (Macracanthus) mcclevei, n. sp. 3 0.617 0.640 0.627 A. (Macracanthus) pittieri (Gahan) 0.6171 - A. (Macracanthus) tuberculatus (Chd.) 3 0.585 0.593 0.589
FEMALES A. (Apostolus) batest (Chd.) Tambopata, Peru 1 0.667 Buena Vista, Bolivia 0.643 1 - A. (Aephnidius) piceolus (Chd.) 5 0.706 0.743 0.725 - A. (Aephnidius) ampliusculus (Chd.) 3 0.692 0.700 0.695 A. (Aephnidius) bonariensis (Chd.) 0.6751 - A. (Aephnidius) ciliatus (Mutch.) 8 0.733 0.750 0.741 ? ? ? A. (Aephnidius) humeralis, n. sp. n. A. (Aephnidius) exiguus, sp. Tambopata, Peru 4 0.692-0.738 0.709 Iquitos, Peru 3 0.698-0.733 0.719 - Parque Nac. Chaco, Argentina 5 0.682 0.712 0.697 ? ? ? A. (Aephnidius) whiteheadi, n. sp. n. A. (Aephnidius) ashei, sp. Tambopata, Peru 6 0.652-0.679 0.665 2Brazil 0.686-0.708 0.697 - A. (Macracanthus) sericatus (Chd.) 3 0.655 0.679 0.667 A. (Macracanthus) thoracicus, n. sp. 0.6671 A. (Macracanthus) erwini, n. sp. Tambopata, Peru 3 0.607-0.642 0.625 - A. (Macracanthus) mcclevei, n. sp. 3 0.617 0.633 0.623 - A. (Macracanthus) pittieri (Gahan) 2 0.597 0.614 0.606 - A. (Macracanthus) tuberculatus (Chd.) 3 0.593 0.610 0.601
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Table 5B. Summary by genus, subgenus and species group of data about variation in HW/ PWM among the New World species of Anaulacus Macleay.
Name of Taxon Range of Values
Genus Anaulacus Macleay - Males 0.58 0.77 - Females 0.59 0.74 new Subgenus Apostolus, subgenus - Males 0.64 0.66 Females 0.67
Subgenus Aephnidius Macleay - Males 0.66 0.77 - Females 0.65 0.76
piceolus species group - Males 0.71 0.73 Females 0.71 - 0.74
ciliatus species group Males 0.68 - 0.77 - Females 0.69 0.75
exiguus species group - Males 0.66 0.72 - Females 0.65 0.74
Subgenus Macracanthus Chaudoir - Males 0.58 0.64 - Females 0.59 0.68
sericatus species group - Males 0.63 0.64 - Females 0.61 0.68
mcclevei species group - Males 0.62 0.64 Females 0.62 - 0.63 tuberculatus species group - Males 0.58 0.62 - Females 0.59 0.61
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Table 6A. Data about variation in proportions of the male median lobe of the Anaulacus genitalia of species of Macleay.
Name of Taxon PAL/MLL OML/MLL PAL/OML
0.108 0.7270.381 A. (Apostolus) batest (Chd.) 0.600 A. (Aephnidius) piceolus (Chd.) 0.136 0.227 0.227 0.400 A. (Aephnidius) ampliusculus (Chd.) 0.136 0.462 A. (Aephnidius) ciliatus (Mutch.) 0.154 0.333 n. 0.308 A. (Aephnidius) humeralis, sp. 0.100 0.325 n. 0.385 A. (Aephnidius) exiguus, sp. 0.114 0.295 n. 0.2290.454 A. (Aephnidius) whiteheadi, sp. 0.104 n. A. (Aephnidius) ashei, sp. 0.081 0.2700.300 A. (Macracanthus) sericatus (Chd.) 0.117 0.2000.583 A. (Macracanthus) thoracicus, n. sp. 0.123 0.193 0.636 A. (Macracanthus) erwini, n. sp. 0.122 0.1710.714 A. (Macracanthus) mcclevei, n. sp. 0.167 0.2170.769 A. (Macracanthus) pittieri (Gahan) 0.104 0.3770.276 A. (Macracanthus) tuberculatus (Chd.) 0.071 0.3710.192
Table 6B. Summary by genus, subgenera and species groups of proportions of the male median lobe of the genitalia of Anaulacus Macleay.
Name of Taxon PAL/MLL OML/MLL PAL/OML
- - - Genus Anaulacus Macleay 0.07 0.17 0.18 0.40 0.19 0.60 Mean of range of variation 0.12 0.29 0.48 new 0.11 0.730.38 Subg. Apostolus, subg. - - - 0.15 0.23 0.34 0.30 0.60 Subg. Aephnidius Macl. 0.08 piceolus sp. group 0.14 0.23 0.60 - - - ciliatus sp. group 0.10 0.15 0.32 0.34 0.31 0.46 - - - exiguus sp. group 0.08 0.11 0.23 0.30 0.30 0.45 - - - 0.17 0.40 0.19 0.77 Subg. Macracanthus Chd. 0.07 0.17 - - sericatus sp. group 0.12 0.17 0.20 0.58 0..71 mcclevei sp. group 0.17 0.22 0.77 - - - tuberculatus sp. group 0.07 0.10 0.37 0.40 0.19 0.28
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Table 7. Sympatric assemblages of species of Anaulacus MacLeay in the Neotropical Region
Locality Resident Species
BELIZE Orange Walk, Dist. Rio Bravo Cons. Area A. pittieri, sericatus, whiteheadi BRAZIL Amazonas, Tef? A. batest, erwini, piceolus, tuberculatus BRAZIL
Matto Grosso, Tapirip? Indian Vill. A. batest, exiguus BRAZIL Para, Santarem A. ciliatus, sericatus BRAZIL Para, Para A. ampliusculus, ashei PANAMA
Canal Zone, Barro Colorado Island A. ampliusculus, batest, sericatus PANAMA Panam?, Las Cumbres A. ampliusculus, ciliatus PERU Madre de Dios, 15 km. NE Puerto Maldonado A. ashei, batest, erwini, exiguus, sericatus
Table 8. Number of species of Anaulacus MacLeay, by 10? intervals.
Latitude Number of species
30-20?N 20-10?N 10-0?N 0-10?S 10-20?S 20-30?S 30-40?S
Table 9. Number of Neotropical species of Anaulacus MacLeay, by number of 10? intervals occupied.
Number of units Number
occupied of species
one 5 two 3 three 4 four 1 five six 1 seven 1
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Table 10. Distribution of zonal or continental precinctive Neotropical species of Anaulacus MacLeay.
CONTINENTAL SOUTH AMERICA
Taxon Area of occurrence
A. bonairiensis Southern Atlantic area A. thoracicus Southern Atlantic area A. ashei Amazon Basin
A. exiguus Western Amazon Basin A. erwini Western Amazon Basin A. tuberculatus South America CONTINENTAL NORTH AMERICA
Taxon Area of occurrence
A humeralis Lower Central America A pittieri Central America A. whiteheadi Nuclear Central America A. mcclevei Northwestern Mexico
Table 11. Distribution in Middle America of Neotropical species of Anaulacus source area. having South America as putative
Species Lower Central Nuclear Central West Indies America America
A. batesi X
A. ampliusculus X A. piceolus X X A. ciliatus X X A. sericatus X X X
Table 12. Putative places of orgin of adelphotaxa of Neotropical species of Anaulacus MacLeay. Vicar/ Vicar/ Putative place of origin Putative place of origin + A. exiguus I A. whiteheadi ashei/ Western Amazon Basin NE South America
A. whiteheadi! A. ashei/ Central America Eastern Amazon Basin
A. sericatus species group/ A. mcclevei+tuberculatus sp. grp/ South America Central America
A. mcclevil A. tuberculatus species group/ Northwestern Mexico Central America
A. pittieri/ A. tuberculatus/ Central America South America
A. sericatus/ A. thoracicus+erwini/ Central America and/or South America: Amazon Basin, Northern South America southward
A. thoracicus/ A. erwini/ Southern Atlantic area Western Amazon Basin
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