ISSN 0071-075X
Memoirs of the Number 170
Entomological Society of Canada 1996
A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON (COLEOPTERA: SCARABEIDAE, SCARABAEINAE)
Franc;ois Genier
Entomological Society of Canada Table of contents on first page Societe d'Entomologie du Canada
Date of issue: November 1996 393 Winston Avenue, Ottawa, Ontario, Canada K2A 1 Y8 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON (COLEOPTERA: SCARABAEIDAE, SCARABAEINAE)
FRAN<;OIS GENIER
Canadian Museum of Nature, Collection Division, Invertebrate Section, PO Box 3443, Station D, Ottawa, Ontario, Canada KIP6P4
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA-No. 170 V. Behan-Pelletier, Editor
THE ENTOMOLOGICAL SOCIETY OF CANADA 393 Winston Avenue Ottawa K2A IY8 1996 FIG. 1. Habitus: Ontherus pilatus sp.nov. CONTENTS
ABS TRAC T ...... 3
RESUME ...... 3
INTRODUCTION ...... 4
MATERIAL ...... 5
ME THODS ...... 6
SYS TEMATICS ...... 7
KEY TO ADULTS OF SPECIES OF ONTHERUS ...... 7
CLAVE PARA SEPARAR LAS ESPECIES DE ONTHERUS ...... 15 Genus ONTHERUS Erichson, 1847 ...... 22
Subgenus CAELONTHERUS nov...... 23
HADROS GROUP ...... 24
1. Ontherus (Caelontherus) hadrossp.nov ...... 24 2. Ontherus (Caelontherus) magnus sp.nov ...... 25
LAMINIFER GROUP ...... 26 3. Ontherus (Caelontherus) laminiferBalthasar, 1938 comb.nov ...... 27
BRE VICOLLIS GROUP ...... 28
4. Ontherus (Caelontherus) gilli sp.nov...... 28
5. Ontherus (Caelontherus) ashei sp.nov...... 30
6. Ontherus (Caelontherus) brevicollis Kirsch, 1871 comb.nov ...... 31 7. Ontherus (Caelontherus) politus sp.nov ...... 33
8. Ontherus (Caelontherus) aequatorius Bates, 1891 comb.nov ...... 34
KIRSCHII GROUP ...... 35 9. Ontherus (Caelontherus) kirschii Harold, 1867 comb.nov ...... 35
10. Ontherus (Caelontherus) sanctaemartae sp.nov ...... 37
ALEXIS GROUP...... 39 11. Ontherus (Caelontherus) alexis (Blanchard), 1845 comb.nov ...... 39
12. Ontherus (Caelontherus) tenuistriatus sp.nov ...... 41 13. Ontherus (Caelontherus) obliquus sp.nov ...... 43
14. Ontherus (Caelontherus) incisus (Kirsch), 1871 comb.nov ...... 45
15. Ontherus (Caelontherus) howdeni sp.nov ...... 47 16. Ontherus (Caelontherus) diabolicus sp.nov ...... 48
MEXICANUS GROUP ...... 50 17. Ontherus (Caelontherus) compressicornis Luederwaldt, 1931
comb.nov ...... 51
18. Ontherus (Caelontherus) pilatus sp.nov ...... 52 19. Ontherus (Caelontherus) trituberculatus Balthasar,
1938 comb.nov...... 53
20. Ontherus (Caelontherus) mexicanus Harold, 1868 comb.nov ...... 55
21. Ontherus (Caelontherus) sextuberculatus sp.nov ...... 58
22. Ontherus (Caelontherus) lunicollis sp.nov ...... 59 23. Ontherus (Caelontherus) monilistriatus sp.nov ...... 62
24. Ontherus (Caelontherus) pseudodidymus sp.nov ...... 63
Subgenus PLANONTHERUS nov...... 67 25. Ontherus (Planontherus) bridgesi Waterhouse,
1891 comb.nov ...... 67
26. Ontherus (Planontherus) rectus sp.nov ...... 69
Subgenus ONTHERUS s. str...... 70
SULCATOR GROUP ...... 71 27. Ontherus (Ontherus) pubens sp.nov ...... 71
28. Ontherus (Ontherus) sulcator (Fab ricius), 1775 comb.nov ...... 74 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
APPENDICULATUS GROUP ...... 78 29. Ontherus (Ontherus) appendiculatus (Mannerheim), 1829 comb.nov...... 78 30. Ontherus (Ontherus) ulcopygus sp.nov ...... 82 31. Ontherus ( Ontherus) podiceps Harold, 1868 comb.nov...... 85 32. Ontherus (Ontherus) azteca Harold, 1869 comb.nov...... 87 33. Ontherus (Ontherus) camheforti sp.nov...... 89 34. Ontherus (Ontherus) grandis Luederwaldt, 1931 comb.nov...... 91 35. Ontherus (Ontherus) carinifrons Luederwaldt, 1930 comb.nov...... 92 36. Ontherus (Ontherus) virescens (Lucas), 1857 comb.nov...... 94 37. Ontherus (Ontherus) irinus Balthasar, 1938 comb.nov...... 96 BREVIPENNIS GROUP...... 97 38. Ontherus (Ontherus) raptor sp.nov...... 97 39. Ontherus (Ontherus) planus sp.nov...... 99 40. Ontherus (Ontherus) lichyi Martinez, 1947 comb.nov...... 100 41. Ontherus (Ontherus) edentulus sp.nov ...... 102 42. Ontherus (Ontherus) hrevipennis Harold, 1867 comb.nov...... 104 43. Ontherus (Ontherus) rectangulidens sp.nov ...... 106 44. Ontherus (Ontherus) carinicollis Luederwaldt, 1931 stat.nov...... 108 45. Ontherus (Ontherus) zikani Luederwaldt, 1930 comb.nov ...... 110 46. Ontherus (Ontherus) lohifrons sp.nov ...... 112 47. Ontherus (Ontherus) cephalotes Harold, 1869 comb.nov ...... 115 DIGI TATUS GROUP ...... 117 48. Ontherus (Ontherus) aphodioides Burmeister, 1874 comb.nov ...... 117 49. Ontherus (Ontherus) digitatus Harold, 1868 comb.nov...... 120 50. Ontherus (Ontherus) elegans Luederwaldt, 1930 comb.nov...... 121 51. Ontherus (Ontherus) androgynus sp.nov ...... 124 52. Ontherus (Ontherus) erosioides Luederwaldt, 1930 comb.nov...... 125 53. Ontherus (Ontherus) amplector sp.nov ...... 127 54. Ontherus (Ontherus) atlantidis sp.nov ...... 129 55. Ontherus (Ontherus) dentatus Luederwaldt, 1930 comb.nov ...... 130 EROSUS GROUP ...... 131 56. Ontherus ( Ontherus) insolitus sp.nov...... 131 57. Ontherus (Ontherus) stridulator sp.nov...... 133 58. Ontherus (Ontherus) erosus Harold, 1875 comb.nov...... 134 PHYLOGENY...... 136 BIOGEOGRAPHY ...... 139 ACKNOWLEDGMENTS...... 141 REFERENCES ...... 141 CHECKLIST OF THE GENUS ONTHERUS ...... 144 ILLUSTRATIONS ...... 146 INDEX...... 168
The Memoirs are subject to the same standards and rev iew requirements as are contrib utions to The Canadian Entomologist excep t that more editorial latitude is permitted. Instructions to authors are printed in the January/Feb ruary issue of The Canadian Entomologist. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON (COLEOPTERA: SCARABAEIDAE, SCARABAEINAE)
FRAN<;OIS GENIER
Abstract Memoirs of the Entomological Society of Canada 170: 168 pp. The species of the genus Ontherus, endemic to Latin America, are revised. A key in English and Spanish for the identification of adults of the 58 recognized species is provided. Three subgenera are recognized (Caelontherus nov., Planontherus nov. and Ontherus s.str.). The following 29 species are described as new: 0. hadros, 0. magnus, 0. gilli, 0. ashei, O.politus, 0. sanctaemartae, 0. tenuistriatus, 0. obliquus, 0. howdeni, 0. diabolicus, 0. pilatus, 0. sextuberculatus, 0. lunicollis, 0. monilistriatus, 0. pseudodidymus, 0. rectus, 0. pubens, 0. ulcopygus, 0. cambeforti, 0. raptor, 0. planus, 0. edentulus, 0. rectangulidens, 0. lobiji-ons, 0. androgynus, 0. amplector, 0. atlantidis, 0. insolitus, 0. stridulator. The following new synonymies are proposed (the first specific name being valid): 0. kirschii Harold = 0. elongatus Waterhouse; 0. alexis (Blanchard) = 0. didymus Erichson= 0. glaucinus Erichson = 0. nevinsoni Waterhouse; 0. incisus (Kirsch) = 0. thoracicus Waterhouse; 0. azteca Harold = 0. villosus Luederwaldt = 0. strius Howden and Young; 0. cephalotes Harold = 0. quadrituberculatus Luederwaldt. The variety 0. zikani var. carinicollis Luederwaldt is now recognized as a valid species. Species are arranged into groups according to their phylogenetic relationship, with a brief diagnosis given for each of the 11 groups of species. The following information is given for each species, when appropriate: detailed literature review, synonymy, description or redescription, illustration of important morphological characters, data of material studied, and geographic distribution.
Genier, F. 1996. Revision des especes du genre Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Memoirs of the Entomological Society of Canada 170: 168 pp.
Resume Les especes appartenant au genre Ontherus, endemique a l' Amerique latine, sont revisees. On donne des cles en langue anglaise et espagnole pour la determination des adultes des 58 especes reconnues. On reconnait trois sous-genres, soit: Caelonthetus nov., Planontherus nov. et Ontherus s.str. Les 29 especes suivantes sont nouvelles: 0. hadros, Cf. magnus, 0. gilli, 0. ashei, 0. politus, 0. sanctaemartae, 0. tenuistriatus, 0. obliquus, 0. howdeni, 0. diabolicus, 0. pilatus, 0. sextuberculatus, 0. lunicollis, 0. monilistriatus, 0. pseudodidymus, 0. rectus, 0. pubens, 0. ulcopygus, 0. cambeforti, 0. raptor, 0. planus, 0. edentulus, 0. rectangulidens, 0. lobifrons, 0. androgynus, 0. amplector, 0. atlantidis, 0. insolitus, 0. stridulator. On propose les synonymies suivantes (le premier nom etant celui qui est valide: 0. kirschii Harold = 0. elongatus Waterhouse; 0. alexis (Blanchard) = 0. didymus Erichson = 0. glaucinus Erichson = 0. nevinsoni Waterhouse; 0. incisus (Kirsch)= 0. thoracicus Waterhouse; 0. azteca Harold= 0. villosus Luederwaldt = 0. strius Howden et Young; 0. cephalotes Harold = 0. quadrituberculatus Luederwaldt. La variete 0. zikani var.carinicollis Luederwaldt est maintenant reconnue comme buena species. Toutes les especes ant ete regroupees selon leurs affinites phylogenetiques et une breve diagnose est donnee pour chacun des 11 groupes d'especes. L'information suivante, lorsque pertinente, est donnee pour chaque espece: revue detaillee de la litterature, synonymie, description ou redescription, illustration des characteres morphologiques importants et donnees de repartitions geographique.
3 4 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
INTRODUCTION
The genus Ontherus was proposed in 184 7 by Erichson for two new species [ 0. didymus and 0. glaucinus (=O. alexis (Blanchard, 1845))] and Copris sulcator (Fabricius). The main character used by Erichson to separate his new genus from Copris was the absence of transverse carinae on the posterior tibia, combined with the reduced median width of the abdominal stemites. The genus Ontherus traditionally has been included in the tribe Dichotomiini. However, this placement, as well as the validity of the two tribes Coprini and Dichotomiini, must be reconsidered. The diagnostic character for defining the tribes, in their most recent sense, is the presence (Coprini) or absence (Dichotomiini) of a transverse carina on the median and/or posterior tibia and the number of main elytral striae, nine in Coprini and eight in Dichotomi ini (Balthasar 1963; Hanski and Cambefort 1991). Although a phylogenetic analysis at the tribal level is beyond the scope of this study, I suggest that these characters are subject to strong external selection pressure, especially the character associated with the legs, and use of these characters as the basis for the definition of tribes is questionable. Presently, I consider Ontherus to be a constituent of a lineage comprising moderate to large (usually over 10 mm) tunnellers (Hanski and Cambefort 1991), with a relatively thin and flexible tegument. Species belonging to this lineage possess the following combination of characters: first segment of antenna! club completely tomentose, anterior edge of head broadly arcuate to semicircular with or without anterior tooth/teeth, lateral edge of gena usually forming an angle less than 90°, prothorax rather loosely articulated with metathorax, pronotum usually marginate posteriorly and with lateral pronotal carina (primitive character), mesostemum lacking deep transverse pubescent sulcus anteriorly (character usually hidden by the proster num), median lobe of metastemum usually rather narrow with anterior edge straight (in primitive species) or broadly arcuate or angulate medially, middle coxae relatively closely set, middle and posterior tibiae with or without transverse carina(e) and with muscle disc attachment to membrane of posterior coxa. This lineage consists of at least the following genera: Chalcocopris Burmeister, Copris Muller, Dichotomius Hope, Holocephalus Hope, Isocopris Pereira and Martinez, Ontherus Erichson (all New World); Catharsius Hope, Copris Muller, Coptodactyla Burmeister, Litocopris Waterhouse, Metacatharsius Paulian, Thyregis Blackbum and Xinidium Harold (all Old World). The genus Heliocopris Hope, although a large tunneller and possessing some of the characters mentioned above, is removed from this lineage. I suggest that the genus Heliocopris belongs to a lineage that comprises Synapsis Bates and the genera currently included in the tribe Scarabaeini (see Hanski and Cambefort 1991). This assumption is based on the followingshared combination of characters: antenna with first segment of antenna! club lacking thick tomentose coating on proximal surface (except in Synapsis), pronotum with a thick fringe of setae at head insertion, elytron with basal portion of fifthinterval much wider than fourth and sixth, lateral edge of elytron carinate on eighth interval (feebly in some species), tarsal claws lacking axillary lamella and internally the muscle disc attachment to posterior coxa is lacking. In addition the proportionally shorter pronotum and the Old World distribution distinguish this lineage. As it now stands, the genus Ontherus consists of 58 species that are distributed throughout continental tropical and subtropical Latin America. A single species [ 0. sulcator (Fabricius)] has been recorded from an insular locality (Trinidad). The greatest diversity of species occurs in southeastern Brazil and the Andes. The present work recognizes three subgenera. The most important characters used to define the subgenera arethe configuration of the mesometasternal suture and the mesepister num. The subgenus Caelontherus nov. comprises 24 species and is restricted mostly to higher elevations in the Andes. The subgenus Planontherus nov. comprises only two species and is distributed at higher and lower elevations in Argentina, Bolivia and Peru (plus doubtful A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 5 records from Colombia and Venezuela). Finally, the subgenus Ontherus s.str. comprises 32 species and is distributed mostly at lower elevations from Mexico to Argentina. In addition to the subgenera, the species have been organized into six species groups for the subgenus Coelontherus nov. and five species groups for the subgenus Ontherus s.str. No species group has been recognized for the subgenus Planontherus nov. as it comprises only two species. The most important characters used to define groups are the organization of the sclerites of the internal sac and the shape of parameres of the aedeagus in combination with the male secondary sexual characters. The Digitatus group is the least homogenous and eventually may need to be reconsidered. This group is kept herein to avoid groups containing only one or two species. The other groups are believed to be natural and are organized in phylogenetic order within each subgenus, as much as possible; the firstlisted being the most primitive. Most Ontherus are coprophagous and/or saprophagous. Some species have evolved complex associations with ants (species belonging to the Brevipennis group) and most likely with mammals, as suggested by the highly modifiedtarsal claws in 0. virescens (see remarks under 0. virescens). The biology of a single species (0. mexicanus) has been studied and described. It is of type II, as defined by Halffter and Edmonds (1982). The nest is assembled as follows: each brood ball is constructed independently and resides in its own chamber; the entranceof the chamber is horizontal; the spherical brood ball bears two raised concentric rings at the egg pole which is oriented laterally. For more detail see Halffter and Edmonds (1982). The genus was last revised by Luederwaldt ( 1931 ), a paper that included identification keys to the Brazilian and "foreign species". Although representing a good synthesis, Luederwaldt's work has few, and poor, illustrations. It is obvious that Luederwaldt did not have access to representatives of some of the species included in the key. Consequently, the identification keys are difficult to use. Important characters were overlooked, such as the shape of the aedeagus and posterior femur in males of some species. Luederwaldt did not study types of previously described species and the correct status of some of these species is not clear.
MATERIAL
This study is based on the examination of more than 9800 adult specimens of Ontherus including type material for most of the available names, as well as some representatives of closely related genera from all tropical and subtropical regions of the world. This material was borrowed from several institutions and private collections. Most of the initialisms listed below were taken from Arnett et al. (1993) and refer to the material listed in the text. The assistance of the curators responsible for these collections, along with the assistance of individuals who sent the specimens for study, is gratefully acknowledged.
AMIC: Antonio Martinez, Rosario de Lerma, Argentina, personal collection. AMNH: American Museum of Natural History, New York, NY; L.H. Herman, Jr. AVEC: Arthur V. Evans, Los Angeles, CA, personal collection. BCRC: Brett C. Ratcliffe, Lincoln, NE, personal collection. BDGC: Bruce D. Gill, Ottawa, ON, personal collection. BMNH: The Natural History Museum, London, U.K.; M. D. Kerley. CASC: California Academy of Sciences, San Francisco, CA; D.H. Kavanaugh, R. Brett. CNCI: Canadian National Collection of Insects, Ottawa, ON; A. Smetana, J. McNamara. CMNC: Canadian Museum of Nature, Ottawa, ON. CUIC: Cornell University Insect Collection, Ithaca, NY; E.R. Hoebeke. DEIC: Deutsches Entomologisches Institut, Eberswalde, Germany; L. Zerche. FGIC: Frarn;:ois Genier, Ay lmer, PQ, personal collection. HAHC: Henry and Anne Howden, Ottawa, ON, personal collection. ICCM: Carnegie Museum of Natural History, Pittsburgh, PA; J.E. Rawlins, R.L. Davidson. 6 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
IMLA: Universidad Nacional de Tucuman, Miguel Lillo, Argentina; J.A. Haedo, A. Teran. INBC: Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; A. Solis. ISNB: Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgique; P. Dessart, M. Cludts. IZAV: Universidad Central de Venezuela, Maracay, Venezuela; J. Clavijo, L. Joly. LACM: Los Angeles County Museum of Natural History, Los Angeles, CA; C.L. Hogue, A.V Evans. LEMC: Lyman Entomological Museum and Research Laboratory, Sainte-Anne-de-Bellevue, PQ; P. M. Sanbome, G. Pelletier. MACN: Museo Argentina de Ciencias Naturales, Buenos Aires, Argentina; A.O. Backmann. MAMC: Miguel Angel Moron, Xalapa, Mexico, personal collection. MCNZ: Funda�ao Zoobotanica do Rio Grande do Sul, Porto Alegre, Brazil; G. Carvalho Ferraz, M.H. Galileo. MCZC: Museum of Comparative Zoology, Cambridge, MA; S.R. Shaw, C.A. Vogt. MLPA: Universidad Nacional de La Plata, La Plata, Argentina; R.A. Ronderos. MLUH: Martin-Luther-Universitat, Halle, Germany; M. Dom. MNHN: Museum national d'Histoire naturelle, Paris, France; Y. Cambefort. MVMA: Museum of Victoria, Natural History, Abbotsford, Australia; K. Walker. MZLU: Lund University, Lund, Sweden; R. Danielsson. MZSP: Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, Brazil; U.R. Martins. NMPC: National Museum of Natural History, Praha, Czechoslovakia; J. Jelinek. OSUC: Ohio State University Collection of Insects and Spiders, Columbus, OH; C.A. Triplehom. PAIC: Patrick Arnaud, Saintry-sur-Seine, France, personal collection. QBUM: Museu Na�ional, Rio de Janeiro, Brazil; M.A. Monne. QCAZ: Universidad Catolica del Ecuador, Quito, Ecuador; G. Onore. SEMC: Snow Entomological Museum, University of Kansas, Lawrence, KS; J.S. Ashe, R.W. Brooks. SMFD: Forschungsinstitut Senckenberg, Frankfurt, Germany; R. zur Strassen. SMTD: Staatliches Museum fiir Tierkunde, Dresden, Germany; R. Krause. TAMU: Texas A&M University, College Station, TX; H.R. Burke, E.G. Riley. UNSM: Department of Entomology, University of Nebraska State Museum, Lincoln, NE; B.C. Ratcliffe. USNM: United States National Museum of Natural History, Washington, DC; R.D. Gordon, G.F. Hevel. UZMH: Universitetets Zoologisca Museum, Helsinki, Finland; H. Silfverberg. WBWC: William B. Warner, Chandler, AZ, personal collection. WDEC: W. David Edmonds, California State Polytechnic, Pomona, CA, personal collection. ZFMK: Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany; H. Roer. ZMAS: Zoological Institut, Academy of Sciences, St. Petersburg, CIS; B.A. Korotyaev. ZMHB: Museum fiir Naturkunde der Humboldt-Universitat, Berlin, Germany; M. Uhlig, F. Hieke. ZMUC: Zoologisk Museum, University of Copenhagen, Denmark; 0. Martin. ZSMC: Zoologische Staatssammlung, Mtinchen, Germany; G. Scherer.
METHODS
Initially the material was sorted using external characters. In a detailed study of the aedeagus it became apparent that the shape of the parameres offered good and stable species and species groups level characters. The anatomy of the internal sac also proved to be useful for specific determination. The latter is so intricately folded that all attempts to evert and inflate it, to study the sclerites in a natural position, failed. It was easier to extract the internal sac fromthe tubus and detach each sclerite fromthe membrane. Three sclerites were present in all species: a double flagella, the lambda sclerite, the sigmoid sclerite, and, in some species, one or two accessory sclerites and/or a small to wide and clearly defined area covered with velum. The aedeagus was mounted on a point and attached to the specimen pin. The dissected sclerites and remaining parts of the membrane were mounted in Canada balsam on an acetate slide, which was also attached to the pin.
MEASUREMENTS. Body length: in dorsal view, from the anterior edge of the clypeus, when the head was fully retracted in a reposed position, to the posteriormost portion of the pygidium. The value is rounded to the nearest millimetre. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 7
Pronotal proportion (W/L): is the width divided by the length along the midline. Median lobe of mesosternum proportion (L/mW): equals the length along midline. This measurement is taken from the anteriormost portion of the median lobe (excluding the longitudinal carina, if present) posteriorly to the intersection of a perpendicular imaginary line at the posterior edges of the median coxal cavities. This value is divided by the minimum width (which is taken approximately medially depending on the species). Pygidium proportion (WIL): equals the maximum width, which is taken at the outer edges of the angles formed by the basal and lateral margins, divided by the length along the midline from the lower edge of the basal margin to the apical edge.
To avoid confusion the following terms and expressions are briefly defined: Accessory sclerites (Figs. 18, 19): the variously shaped and usually slightly sclerotized area of internal sac of the aedeagus (occasionally present). Axillary lamella (of tarsal claws): a small appressed leaflike sclerite situated laterally at the base of each tarsal claw. Connecting striae: the portion of the striae between two strial punctures. Edge: the two dimensional outermost portion of a three-dimensional structure. Lambda sclerite: the trifid and strongly sclerotized area of the internal sac of the aedeagus (always present). Margin: the usually elevated or otherwise modified area along the edge. Muscle disc attachment to membrane of posterior coxa: small funnel-shaped internal structure attached to the membrane connecting metasternum to posterior coxa. The exact function of this sclerotized structure is unknown to me. Sigmoid sclerite: The flat and S-shaped sclerotized area of the internal sac of aedeagus (always present).
Distribution of all species is based on label data associated with specimens examined. Full label data, with original spelling and punctuation, is given only for type material. The slash symbol is used to separate data on different labels. The condensed locality information is listed under State, Department, or Provinces for each country. An asterisk ( * ) following a locality indicates that the information defining the locality was incomplete or the locality could not be situated. Other data, such as ecological notes, habitats, trapping methods, collecting month(s) are listed under" additional data" when available, or within the complete label data for type material. Missing letters in abbreviated words have been put in square brackets when required. Wherever elevation is converted from feet to metres, the metric value is rounded to the nearest 10 m. Ty pes or specimen deposition is indicated by the collection initialism in parentheses.
SYSTEMATICS
KEY TO ADULTS OF SPECIES OF ONTHERUS
REMARKS ABOUT THE KEY. This key is to adult males and females. The most reliable and easy way to separate males from females is by examining the shape of the last abdominal sternite. In males (Figs. 2, 4) the sternite is always distinctly narrowed medially, whereas in females (Figs. 3, 5) it is parallel or slightly wider medially. Males can be distinguished from females using secondary sexual characters such as cephalic and pronotal developments and the modification of the anterior and posterior legs. These characters are not constant throughout the genus. The key has been constructed using only external characters. In some cases the extraction of the aedeagus will be necessary to confirm the identity of small and/or abraded individuals.
1. Mesometasternal suture (Figs. 62-66) straight, in some specimens broadly arcuate, never angulate medially; median lobe of metasternum usually lacking margin anteriorly. Clypeofrontal suture always tuberculate medially. Species restricted to higher elevations (subgenus Caelontherus nov.) ...... 2 8 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
1'. Mesometastemal suture (Figs.8-13, 67- 70) angulate medially (if junction appears straight then clypeofrontal suture carinate); median lobe of metastemum marginate anteriorly, similar to coxal margin, margin rarely atrophied, in some specimens anterior edge lacks margin but anterior edge raised higher than mesosternum.Species usually restricted to lower elevations ...... 25
2(1'). Anterior tibia with 3 teeth in males, 4 in females.Male secondary sexual characters always located on head and pronotum. Posterior femur not modified.Female head with pair of closely set tubercles or transverse horn (exceptionally in females of 0. compressicornis process appears conical, but then minimum width of median lobe of mesosternum subequal to half maximum width of middle femur)...... 3 2'. Anterior tibia with 4 teeth in males and females.Male secondary sexual characters restricted to shape, orientation of apical tooth of anterior tibia (Fig.59) and/or shape of posterior femur (Figs. 71, 72) or rarely to disc of pronotum if apical tooth of anterior tibia similar to that of females. Female head with conical process or with clypeofrontal carina widely interrupted medially ...... 16
3(2). Anterior portion of median lobe of metasternumdeclivous on most of width; lacking vestigial longitudinal carina or sulcus medially. Elytral striae with punctures simple on disc, punctures less than twice the width of connecting striae, punctures rarely deeper than connecting stria ...... 4 3'. Anterior portion of median lobe of metasternum usually marginate, at least laterally; if anterior portion appears largely declivous then usually with vestigial longitudinal carina or sulcus medially.Elytral striae with punctures rounded, impressed, usually umbilicate on disc, punctures more than twice width of connecting striae ...... 8
4(3). Anterior angles of pronotum with punctures clearly larger, denser than discal punctures. Anterior edge of pronotum only slightly emarginate internally to eyes.Male pronotum (Fig. 25) with, at most, 2 contiguous small swellings on anterior fifth. Female pronotum with, at most, 2 small swellings.Male head at most armed with low conical process.Bolivia; Peru ...... 11.0. alexis (Blanchard) 4'. Anterior angles of pronotum with punctures similar in size and density to discal punctures. Anterior edge of pronotum deeply emarginate behind eyes, emargination weaker in smaller individuals.Female pronotum with 4 small swellings, or 2 small swellings and 2 transverse carinae, or 2 transverse carinae, small individuals with 2 small swellings only.Male head armed with long, curved horn, reduced to conical process in smaller individuals ...... 5
5(4'). Elytral striae similarly impressed on disc and apical declivity, feebly impressed on apical callus ...... 6 5'. Elytral striae more deeply impressed on apical declivity than on disc, normally impressed on apical callus ...... 7
6(5). Inner edges of pronotal carinae raised higher than outer edges; distance between inner edges of carinae (Fig. 31) longer than width of carina in larger males. Colombia; Ecuador ...... 14.0. incisus (Kirsch) 6'. Inner edges of pronotal carinae lower than lateral edges; distance between inner edges shorter than width of carina (Fig.33) in larger males. Peru ...... 15. 0. howdeni sp.nov.
7(5'). Elytral surface glossy, punctures minute ( 12 X 1). Male pronotal carinae (Fig. 27) almost transverse. Peru ...... 12.0. tenuistriatus sp.nov. 7'. Elytral surface alutaceous, distinctly punctate (12X).Male pronotal carinae (Fig.29) forming angle of less than 90°.Bolivia; Peru ...... 13. 0. obliquus sp.nov.
8(3'). Occurring from Colombia southward ...... 9 8'. Occurring from Panama northward ...... 14
I Magnification. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 9
9(8). Male pronotum (Figs. 39, 41) with transverse carina on each side of midline, carinae with lateral portion reduced or much lower than inner portion, sometimes with small supplementary tubercle medially, or with at least single central minute tubercle in smaller individuals. Female pronotum with 3 tubercles or single median tubercle in smaller individuals ...... 10 9'. Male pronotum (Figs. 35, 37, 47, 49) with 4 tubercles, lateral tubercles much larger than inner ones, or if with transverse carina on each side of midline, lateral portion always distinctly raised; smaller individuals with 2 minute tubercles medially, sometimes distal tubercles present, small. Female pronotum with 4 tubercles, sometimes connected with carina, or with 2 low tubercles medially in smaller individuals ...... 11
10(9). Median lobe of metastemum with at least few setae similar to those on mesosternum, especially anteriorly, along mesocoxa.Median lobe of metasternum completely marginate anteriorly, margin wide. Colombia; Ecuador ...... 19. 0. trituberculatus Balthasar 1 O'. Median lobe of metastemum with minute, appressed pubescence only, setae much finer than those on mesostemum.Median lobe of metasternum with anterior margin atrophied medially. Ecuador ...... 18. 0. pilatus sp.nov.
11(9'). Male pronotum (Fig. 37) with transverse carina on each side of midline, carinae approximate, covering almost entire width of pronotum in larger individuals, reduced to 2 transverse swellings with inner portion slightly tuberculate in smaller individuals. Male cephalic horn laterally compressed in larger individuals. Females with lateral portions of pronotum with punctures separated by less than their diameter. Ecuador ...... 17. 0. compressicornis Luederwaldt 11'. Male pronotum (Figs. 35, 47, 49, 51) with 4 tubercles, outer ones much larger than inner ones, reduced to swellings in smaller individuals. Male cephalic horn with apex acute or forkedin some larger individuals.Females with lateral portion of pronotum with punctures minute, only slightly larger on anterior angles ...... 12
12(11'). Median lobe of metastemum with few long setae along mesocoxal margin.Surface of elytra completely glossy, without any traces of alutaceous microsculpture (25 X ).Elytral striae with punctures subcircular.Colombia; Ecuador ...... 16. 0. diabolicus sp.nov. 12'. Median lobe of metastemum lacking long setae along mesocoxal margin. Surface of elytra usually strongly alutaceous (25 X). Ely tral striae with punctures circular.Bolivia; Colombia ...... 13
13(12'). Median lobe of metastemum usually with fine longitudinal carina anteriorly. Minimum width of median lobe of metasternum approximately half maximum width of mesofemur.
Colombia ...... 22. 0. lunicollis sp.nov. 13'. Median lobe of metasternum lacking fine longitudinal carina anteriorly.Minimum width of median lobe of metasternum approximately three-fourths maximum width of mesofemur. Bolivia ...... 23. 0. monilistriatus sp.nov.
14(8'). Median lobe of metasternum usually with fine longitudinal carina anteriorly.Anterior angles of pronotum with punctures confluent. Male pronotum (Fig. 51) of larger individuals with 4 tubercles, lateral ones much larger than inner ones. Costa Rica; Panama ...... 24. 0. pseudodidymus sp.nov. 14'. Median lobe of metasternum lacking fine longitudinal carina anteriorly. Anterior angles of pronotum with punctures separated by more than their diameter.Male pronotum (Figs.43, 45) of larger individuals with 2 longitudinal carinae medially, 2 tubercles laterally .... 15
15(14'). Elytral intervals strongly impressed along strial punctures. Male pronotal carinae (Fig.43) subparallel in larger individuals. Mexico; Guatemala ...... 20. 0. mexicanus Harold 15'. Elytral intervals slightly impressed along strial punctures. Male pronotal carinae (Fig.45) forming angle of approximately 35° in larger individuals.Costa Rica ...... 21. 0. sextuberculatus sp.nov. 10 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
16(2'). Median lobe of metastemum with conspicuous setae similar to those on lateral lobes.Male anterior tibia with apical tooth triangular, similar to penultimate tooth in aspect.Females with anterior angles of pronotum granulate ...... 17 16'. Median lobe of metastemum lacking conspicuous setae or setae shorter than those on lateral lobes. Male anterior tibia with apical tooth slender and pointing forward.Females with anterior angles of pronotum punctate ...... 19
17(16). Size small, less than 12 mm. Gena! surface with at least some coarse rugose areas.Brazil; Peru ...... 3. 0. laminiferBalthasar 17'. Size large, over 17 mm.Genal surfacesmooth, with at most few scattered, weakly impressed punctures anterior to eye ...... 18
18(17'). Elytral striae distinctly crenulate.Ecuador ...... 1. 0. hadros sp.nov. 18'. Elytral striae not crenulate, lacking distinct puncture.Ecuador ..... 2. 0. magnus sp.nov.
19(16'). Cephalic tubercle usually low (less than width of eye), located behind carinate, arcuate clypeofrontal suture.Males with posterior edge of posterior femur with apical tooth.Elytra not strongly alutaceous ...... 20 19'. Cephalic tubercle usually high (more than width of eye), on subangulate clypeofrontal suture, carina atrophied (except 0. aequatorius tubercle behind suture and then stria! punctures rounded, well-impressed, much wider than stria, occurring in Ecuador). Males with posterior edge of posterior femur unmodified.Elytra alutaceous or not ...... 21
20(19). Elytral striae not crenulate on disc. Median lobe of metasternum usually with few con spicuous long setae.Colombia ...... 9. 0. kirschii Harold 20'. Elytral striae crenulate on disc.Median lobe of metasternum lacking long setae, with at most minute pubescence.Colombia ...... 10. 0. sanctaemartae sp.nov.
21(19'). Anterior femur with anterioventral sulcus well-delimited apically, almost reaching apex, abruptly terminated ...... 22 21 '. Anterior femur with anterioventral sulcus vaguely delimited apically, gradually becoming atrophied toward apex ...... 23
22(21). Median lobe of metasternum (Fig.62) moderately wide (L/m W = 2.6). Meta sternum flat or convex on disc.Pronotum lacking conspicuous punctures posteriorly. Venezuela ...... 4. 0. gilli sp.nov.
22'. Median lobe of metastemum (Fig.63) wide (L/mW = 1.9). Metastemum broadly concave on disc. Pronotum conspicuously punctate on small area posteriorly.Peru ...... 5. 0. ashei sp.nov.
23(21'). Median lobe of metastemum (Fig.66) marginate anteriorly, similar to lateral edge. Elytra strongly alutaceous.Ecuador...... 8. 0. aequatorius Bates 23'. Median lobe of metastemum (Figs.64, 65) declivous anteriorly, margin atrophied medially. Elytra smooth, at most slightly alutaceous ...... 24
24(23'). Elytral striae with punctures deeply foveolate on apical declivity (especially striae 1 and 2). Posterior coxa with median sulcus foveolate.Ecuador ...... 7. 0. politus sp.nov. 24'. Elytral striae with punctures similarly impressed on apical declivity and disc.Posterior coxa with median sulcus not foveolate.Colombia; Venezuela ...... 6. 0. brevicollis Kirsch
25(1'). Mesepisternum flat, lacking carina parallel to posterior edge. Pronotum lacking any indication of longitudinal sulcus or depression posteriorly along midline (subgenus Planontherus nov.) ...... 26 25'. Mesepistemum with carina parallel to posterior edge. Pronotum usually with distinct -longitudinal sulcus or depression posteriorly along midline (subgenus Ontherus s.str.) ...... 27 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 11
FIGS. 2-5. Abdominal sternites: 2, 3, Ontherus ulcopygus sp.nov. (2, male; 3, female); 4, 5, 0. podiceps Harold (4, male; 5, female).
26(25). Median lobe of metasternum (Fig. 67) with some conspicuous setae similar to those on mesosternum. Argentina; Bolivia; Peru ...... 25. 0. bridgesi Waterhouse 26'. Median lobe of metastemum lacking any trace of pubescence. Peru ...... 26. 0. rectus sp.nov.
27(25'). Clypeofrontal suture lacking carina, without tubercles. Argentina; Bolivia; Brazil;
Paraguay ...... 49. 0. digitatus Harold 27'. Clypeofrontal suture tuberculate or carinate, if carinate then carina broadly arcuate, low, or broadly arcuate with single central or pair of central tubercles, or broadly arcuate, trilobate. Thorax usually subequal in width to elytra, if wider, then body size less than 10 mm and clypeofrontal suture uni- or bituberculate (compare also couplet 27") ...... 28 27". Clypeofrontal suture strongly carinate, carina semicircular, trapezoidal or subquadrate, carina of some specimens quadrilobate or with 2 widely spaced denticles. Thorax usually
distinctly wider than elytra ...... 36
28(27'). Abdominal stemites setose medially, setae similar to those on lateral portion of metasternum [rarely, abdominal sternites appear smooth medially but then posterior edge of metacoxa finely serrated (character best seen when posterior coxa rotated forward to expose posterior margin)] ...... 29 28'. Abdominal sternites bare medially, with at most row of short setae along posterior edge of metacoxa. Posterior edge of metacoxa not finely serrate ...... 45
29(28). Clypeofrontal suture tuberculate, with at most fine carina not wider than adjacent punctures ...... 30
29'. Clypeofrontal suture carinate ...... 35
30(29). Posterior edge of metacoxa smooth. Most punctures on lateral lobes of metasternum oval, each with posterolateral edge feebly indicated ...... 31 30'. Posterior edge of metacoxa finely serrated.Most punctures on lateral lobes of metasternum rounded, clearly delimited ...... 34
31(30). Pygidial margin usually complete. Median lobe of metasternum with at most minute setae. Argentina; Bolivia; Brazil; Colombia; Ecuador; French Guiana; Guyana; Paraguay; Uruguay; Venezuela ...... 29.0. appendiculatus (Mannerheim) 12 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
31' . Pygidial margin incomplete, inner edge feebly indicated apically. Median lobe of metasternum usually setose anteriorly. Species with more restricted distribution, less frequently encountered ...... 32
32(31 '). Lateral lobes of metasternum densely covered with long pubescence. Posterior edge of metafemur glabrous. Size usually over 18 mm.Argentina .... 34. 0. grandis Luederwaldt 32'. Lateral lobes of metastemum scarcely pubescent, pubescence gradually shortened toward lateral margin. Posterior edge of metafemur with few short setae. Size usually less than 15 mm. Brazil; French Guiana; (doubtful Colombian record) ...... 33
33(32'). Maximum length of last abdominal segment (Figs.2, 3) subequal to minimum combined length of remaining abdominal segments.Brazil ...... 30. 0. ulcopygus sp.nov. 33'. Maximum length of last abdominal segment (Figs. 4, 5) approximately twice minimum combined length of remaining abdominal segments. Brazil; French Guiana ...... 31. 0. podiceps Harold
34(30'). Abdominal stemites alutaceous between punctures. Bolivia; Brazil; Colombia; Costa Rica; Ecuador; Mexico; Panama; Paraguay; Peru ...... 32. 0. azteca Harold 34'. Abdominal sternites smooth between punctures. Guyana; French Guiana ...... 33. 0. cambeforti sp.nov.
35(29'). Dorsum with bright metallic green sheen. Tarsal claws strongly enlarged, "J" -shaped. Brazil ...... 36. 0. virescens (Lucas) 35'. Dorsum dark brown, lacking metallic sheen.Tarsal claws much smaller than apical segment, unmodified.Brazil; Peru ...... 35. 0. carinifrons Luederwaldt
36(27"). Prosternum posterior to procoxa, completely pubescent, rarely with pubescence restricted to lateral portions only ...... 37 36'. Prostemum, behind procoxa, glabrous, smooth, with at most few erect setae along basal margin ...... 41
37(36). Clypeofrontal carina semicircular or subtrapezoidal, median portion always higher. Combined length of head and pronotum not equal in length to elytra. Bolivia ...... 38 37'. Clypeofrontal carina subquadrate, lateral angles usually raised. Combined length of head and pronotum of some specimens equal in length to elytra. Colombia; Costa Rica; Panama;
Peru; Venezuela ...... 39
38(37). Prostemum, behind procoxae, evenly and extensively covered with long erect setae. Male anterior femur unmodified. Male anterior tibia with distance between penultimate and apical teeth more than twice the distance between antepenultimate and penultimate teeth. Bolivia ...... 38. 0. raptor sp.nov. 38'. Prostemum, behind procoxae, with pubescence lacking on disc. Male anterior femur flattened on basal fourth. Male anterior tibia with teeth approximately equidistant. Bolivia ...... 39. 0. planus sp.nov.
39(37'). Combined length of head and pronotum shorter than length of elytra. Basal margin of pronotum slightly widened medially ...... 40 39'. Combined length of head and pronotum subequal in length to elytra. Basal margin of pronotum not widened medially. Colombia; Costa Rica; Panama ...... 42. 0. brevipennis Harold
40(39). Male anterior tibia with vestigial fourth apical tooth. Brazil; Colombia; French Guiana;
Venezuela ...... 40. 0. lichyi Martinez 40'. Male anterior tibia lacking fourth apical tooth.Ecuador; Peru . .. 41. 0. edentulus sp.nov.
41(36'). Clypeofrontal carina subquadrate or trapezoidal, at least with anterior edge straight in small individuals ...... 42 41' . Clypeofrontal carina semicircular ...... 44 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 13
6 7
FIGS. 6, 7. Tarsal claws: 6, Ontherus carinicollis Luederwaldt; 7, 0. zikani Luederwaldt. Line= 0.5 mm.
8 9 10
FIGS. 8-11. Median lobe of metastemum: 8, 9, Ontherus lohifrons sp.nov. (8, male; 9, female); 10, 11, O.cephalotes Harold (10, male; 11, female). Line= 2.0 mm.
42(41). Elytral striae coarsely microsculptured, punctures only slightly wider than stria. Clypeofrontal carina quadrate, wide, half width of head in larger individuals. Male anterior tibia with apical tooth truncate at apex. Male posterior femur with ventroposterior edge always complete, produced posteriorly on basal half. Brazil ...... 43. 0. rectangulidens sp.nov. 42'. Elytral striae moderately alutaceous, punctures much wider than stria. Clypeofrontal carina trapezoidal. Anterior edge of clypeofrontal carina less than half the width of head. Male anterior tibia with apical tooth acute. Male posterior femur with ventroposterior carina atrophied, at most straight on basal half in small individuals ...... 43
43(42'). Tarsal claws (Fig.6), fully extended, at least one-third as long as last tarsal segment.Th orax conspicuously wider than elytra.Pronotum at least with weak depression preceded anteriorly by transverse bulge.Argentina; Brazil; Paraguay ...... 44.0. carinicollis Luederwaldt 43'. Tarsal claws (Fig. 7), fully extended, at most one-fourth as long as last tarsal segment. Thorax approximately as wide as elytra.Pronotum convex anteriorly.Argentina; Brazil
...... 45. 0. zikani Luederwaldt
44(41'). Clypeofrontal carina lacking emargination medially. Anterior margin of median lobe of metasternum (Fig. 8) produced into longitudinal keel in males, reduced to wide, abruptly terminated carina in females (Fig.9). Brazil ...... 46. 0. lobifrons sp.nov. 44'. Clypeofrontal carina emarginate medially. Anterior margin of median lobe of metastemum (Fig. I 0) produced into large conical process in males, in small individuals apex truncated, margin always atrophied, reduced to low arcuate carina with central portion emarginate in females (Fig. 11). Brazil ...... 47. 0. cephalotes Harold
45(28'). Clypeofrontalsuture tuberculate or carinate with single or 2 contiguous tubercles or trilobate
carina ...... 46
45'. Clypeofrontal suture broadly arcuate, carinate ...... 53 14 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
46(45). Clypeofrontal suture with trilobate carina.Anterolateral margin of metacoxa greatly enlarged, strigulate ...... 47 46'. Clypeofrontal suture tuberculate or carinate with 1 or 2 contiguous tubercles medially. Anterolateral margin of metacoxa unmodified ...... 49
4 7( 46). Anterolateral portion of metastemum with anteriormost transverse carina extending beyond lateroposterior edge of strigulate area.Male posterior femur lo bate in basal two-thirds, distal edge of lobe abruptly terminated, in larger males distal edge excavate. Metasternum longitudinally sulcated on disc. Brazil ...... 58. 0. erosus Harold 47'. Anterolateral portion of metastemum with anteriormost transverse carina almost in line with lateroposterior edge of strigulate area. Male posterior femur sharply and strongly carinate through most of its length, carina not abruptly terminated at apical two-thirds. Metastemum foveolate on disc ...... 48
48(47'). Median lobe of metastemum wide (L/mW = 2.0). Lateral lobe of metasternum usually with 6-8 distinct transverse carinae. Male posterior femur carinate throughout Brazil ...... 56. 0. insolitus sp.nov.
48'. Median lobe of metastemum moderately wide (L/m W = 2.6). Lateral lobe of metastemum usually with less than 5 distinct transverse carinae. Male posterior femur carinate in basal half. Brazil ...... 57. 0. stridulator sp.nov.
49( 46'). Length of median lobe of metastemum less than twice its maximum width (similar to Fig. 12). Posterolateral portion of proepistemum densely and evenly pubescent; elytra and base of pronotum lacking metallic sheen ...... 50 49'. Length of median lobe of metasternum twice its maximum width (similar to Fig. 13). Posterolateral portion of proepisternum with single row of setae parallel to lateral edge, sometimes with fewsupplementary setae.Elytra and base of pronotum usually with green metallic sheen ...... 52
50(49). Pygidium very wide, W/L = 2.7 in males or 2.8 in female; punctures moderate in size, umbilicate, deeply impressed. Size less than 12 mm. Argentina; Bolivia; Brazil; Paraguay; Peru; Venezuela ...... 48. 0. aphodioides Burmeister
50'. Pygidium wide, W /L = 2.1 in males or 2.3 in female; punctures small, not umbilicate, weakly impressed. Size usually over 12 mm. Distributed otherwise ...... 51
51(50'). Posterior edge of posterior femur with at most few short, fine setae apically. Male posterior femur (Fig.74) with ventral surface finely sulcate throughout anteriorly.Argentina; Bolivia; Brazil; Colombia; Ecuador; French Guiana; Guyana; Paraguay; Surinam; Trinidad; Uruguay; Venezuela ...... 28. 0. sulcator (Fabricius) 51 '. Posterior edge of posterior femur usually distinctly pubescent throughout (pubescence best seen in lateral view).Male posterior femur (Fig.73) lacking fine sulcus along anterior edge, or with at most ill-defined sulcus on apical half. Argentina; Bolivia; Brazil; Colombia; Ecuador; Peru; Venezuela ...... 27. 0. pubens sp.nov.
52(49'). Clypeofrontal carina with single tubercle.Anterior edge of posterior femur deeply notched in males. Brazil ...... 50. 0. elegans Luederwaldt 52'. Clypeofrontal carina with 2 contiguous tubercles. Anterior edge of posterior femur unmodified in males.Bolivia; Brazil; Paraguay; Uruguay .....51. 0. androgynus sp.nov.
53(45'). Pygidial margin well-defined apically...... 54 53'. Pygidial margin ill-definedapically ...... 55
54(53). Posterolateral portion of proepisternum densely, evenly pubescent. Prostemum densely covered with long, erect pubescence.Brazil ...... 37. 0. irinus Balthasar 54'. Posterolateral portion of proepistemum with single row of setae parallel to lateral edge. Prostemum with scattered, appressed setae.Uruguay ...... 54. 0. atlantidis sp.nov. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 15 /
13
12
FIGS. 12, 13. Median lobe of metastemum: 12, Ontherus sulcator (Fabricius); 13, 0. androgynus sp.nov.
Line = 1.0 mm.
55(53'). Pygidium transversely swollen basally.Posterior edge of metatrochanter completely sulcate, sulcus with at least 6 long setae.Anterior edge of posterior femur angularly produced basally
in males. Brazil ...... 55. 0. dentatus Luederwaldt 55'. Pygidium not transversely swollen basally. Posterior edge of metatrochanter with short sulcus, sulcus with fewer than 5 long setae. Anterior edge of posterior femur modified at apex in males ...... 56
56(55'). Pronotum evenly punctate. Argentina; Brazil; Paraguay ...52. 0. erosioides Luederwaldt 56'. Pronotum finely punctate on disc, punctures much smaller than on anterior angle. Brazil ...... 53. 0. amplector sp.nov.
CLAVE PARA SEPARAR LAS ES PECIES DE 0NTHERUS
OBSERVACIONES ACERC A DE LA CLAVE. Esta clave es para adultos machos y hembras. La forma mas confiable y facil de separar machos de hembras es examinando la forma del ultimo estemito abdominal. En los machos (Figs. 2, 4) el estemito es siempre claramente mas angorto medialmente, mientras que en las hembras (Figs. 3, 5) es paralelo o ligeramente mas ancho medialmente. Los machos pueden ser distinguidos de las hembras usando caracteres sexuales secundarios tales como desarrollos cefalicos o pronotales y la modificaci6n de las patas anteriores y posteriores. Estos caracteres no son constantes en todo el genera. La clave ha sido hecha usando solo caracteres extemos. En algunos casos la extracci6n de! edeago puede ser necesaria para confirmar la identificaci6n, de individuos pequefios o desgastados.
1. Sutura mesometaesternal (Figs. 62- 66) recta, en algunos especfmenes ligeramente arqueada, no angular medialmente; 16bulo medio del metaesterno generalmente carente de margen anteriormente.Sutura clfpeofrontal siempre tuberculada.Especie generalmente restringida a elevaciones altas (subgenero Caelontherus nov.) ...... 2 l'. Sutura mesometaestemal (Figs.8-13, 67 -70) angular medialmente (si la union parece recta entonces la sutura clipeofrontal es carinada); 16bulo medio del meteastemo con margen anteriormente, similar al margen coxal, margen raramente poco visible, en algunos especimenes el borde anterior se eleva mas alto que el mesoestemo. Especie generalmente restringida a elevaciones bajas...... 25
2(1'). Protibia con 3 dientes en el macho, 4 en la hembra.Caracteres sexuales secudarios del macho siempre localizados en la cabeza y el pronoto. Metafemur nunca modifiecado. Cabeza de la hembra con par de tuberculos cercanamente colocados o cuemo transversal (excepcionalmente en hem bras de 0. compressicornis el proceso parece c6nico pero el ancho mfnimo del l6bulo medio del mesoesterno es igual a la mitad del ancho maxima del
mesofemur) ...... 3 16 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
2'. Protibia con 4 dientes en el macho y la hembra.Caracteres sexuales secundarios del macho restringidos a la forma y orientaci6n del diente apical de la protibia (Fig.59) y/o la forma del metafemur (Figs.71, 72) o raramente al disco del pronoto si el diente apical de la protibia es similar al de la hembra.Cabeza de la hem bra con proceso c6nico o con carina clipeofrontal ampliamente interrumpida medialmente ...... 16
3(2). Porci6n anterior del 16bulo medio del metaestemo en declive en la mayoria de su ancho; careciendo medialmente de carina o surco longitudinal vestigial. Estrias elitrales con puntuaciones sin umbilicas en el disco, puntuaciones menos de 2 veces el ancho de la estria que los conecta, puntuaciones raramente mas profundas que el resto de la estria .. 4 3'. Porci6n anterior del 16bulo medio del metaestemo generalmente con margen, al menos lateralmente, si la porci6n anterior parece en declive entonces generalmente con carina o surco longitudinal vestigial medialmente. Estrias elitrales con puntuaciones redondeadas, impresas y generalmente umbilicadas en el disco, puntuaciones mas de 2 veces el ancho de las estrias que las conectan ...... 8
4(3). Angulos anteriores del pronoto con puntuaciones claramente mas grandes y densas que las puntuaciones discales. Borde anterior del pronoto solo ligeramente emarginado internamente a los ojos. Pronoto del macho (Fig. 25) con, a lo mas, 2 pequefias y contiguas elevaciones sobre el quinto anterior. Pronoto de la hembra con, a lo mas, 2 muy pequefias elevaciones. Cabeza del macho armada a lo mas con 1 proceso c6nico pequefio.Bolivia; Peru ...... 11. 0. alex is (Blanchard) 4'. Angulos anteriores del pronoto con puntuaciones similares en tamafio y densidad a las puntuaciones discales. Borde anterior del pronoto profundamente emarginado deras de los ojos, emarginaci6n mas debil en los individuos mas pequefios.Pronoto de la hembra con 4 pequefias elevaciones 6 2 pequefias elevaciones y 2 carinas transversas 6 2 carinas transversas, individuos muy pequefios con solo 2 pequefias elevaciones. Cabeza del macho armada con 1 cuerno largo y curvado, reducido a 1 proceso c6nico en los individuos mas pequefios ...... 5
5(4'). Estrias elitrales similarmente impresas en el disco yen el area apical, atrofiadas en los callos apicales...... 6 5'. Estrias elitrales mas profundamente impresas en el area apical, normalmente impresas en los callos apicales...... 7
6(5). Bordes internosde las carinas pronotales se elevan mas alto que los bordes laterales; distancia entre los bordes internos de las carinas (Fig.31) mas grande que el ancho de la carina en los machos mas grandes. Colombia; Ecuador...... 14. 0. incisus (Kirsch) 6'. Bordes internos de las carinas pronotales mas bajos que los bordes laterales; distancia entre los bordes internos mas corta que el ancho de la carina (Fig.33) en los machos mas grandes. Peru ...... 15. 0. howdeni sp.nov.
7(5'). Superficie elitral lustrosa, punctuaciones diminutas (12X). Carinas pronotales del macho (Fig. 27) casi transversas. Peru ...... 12. 0. tenuistriatus sp.nov. ' 7 . Superficie elitral con microescultura y con puntuaciones claras (12X). Carinas pronotales del macho (Fig. 29) formando un angulo de menos de 90 grados. Bolivia; Peru ...... 13. 0. obliquus sp.nov.
8(3'). Especies que se encuentran de Colombia hacia el sur...... 9 8'. Especies que se encuentran de Panama hacia el norte ...... 14
9 (8). Pronoto del macho (Figs.39, 41) armado con carina transversa a cada lado de la lfnea media, carinas con porci6n lateral atrofiada o mucho mas baja que la porci6n interna, algunas veces medialmente con tuberculo pequefio suplementario, o con al menos tuberculo pequefio central en los individuos mas pequefios. Pronoto de la hembra armado con 3 tuberculos o 1 solo tuberculo medio en individuos mas pequefios ...... 10 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 17
9'. Pronoto del macho (Figs.35, 37, 47, 49) armado con 4 tuberculos, tuberculos laterales mucho mas grandes que los intemos, o si hay carina transversal a cada lado de la linea media la porci6n lateral siempre claramente elevada, individuos mas pequefios con 2 tuberculos pequefios medialmente, agunas veces los tuberculos distales estan presentes y muy pequefios. Pronoto de la hembra armado con 4 tuberculos, algunas veces conectados por una carina, o con 2 tuberculos bajos medialmente en los individuos mas pequefios .. . 11
10(9). L6bulo medio del metaesterno con al menos unas pocas setas similar a las del mesoesterno, especialmente en la parte anterior y a lo largo de la mesocoxa. L6bulo medio del metastemo con margen completo anteriormente, margen ancho.Colombia; Ecuador ...... 19. 0. trituberculatus Balthasar 10'. L6bulo medio del metaesterno solo con pubescencia muy pequefia y recostada contra la superficie, setas mucho mas finas que las del mesoestemo. L6bulo medio del metaesterno con el margen anterior poco visible medialmente.Ecuador ...... 18. 0. pilatus sp.nov.
11(9'). Pronoto del macho (Fig.37) con carina transversal a cada lado de la lfnea media, carinas cercanas y cubriendo casi todo el ancho del pronoto en los indivfduos mas grandes, reducidas a 2 protuberancias transversales con la porci6n interna ligeramente tuberculada en los indivfduos mas pequefios.Cuerno cefalico del macho comprimido lateralmente en los indivfduos mas grandes. Hembra con las porciones laterales del pronoto con puntuaciones separadas por una distancia menor a su diametro.Ecuador ...... 17. 0. compressicornis Luederwaldt 11'. Pronoto del macho (Figs. 35, 47, 49, 51) con 4 tuberculos, los mas extemos mas grandes que los intemos, reducidos a elevaciones en los indivfduos mas pequefios.Cuernos cefalicos de los machos con el apice agudo o bifurcado en algunos indivfduos mas grandes. Hembra con la porci6n lateral del pronoto con puntuaciones diminutas, solo ligeramente mas grandes en los angulos anteriores ...... 12
12(11'). L6bulo medio del metaesterno con hilera de setas largas a lo largo del margen mesocoxal. Superficie de los elitros completamente pulido, sin ningun vestigio de micro escultura (25X).Estrfas elitrales con puntuaciones casi circulares. Ecuador; Colombia ...... 16. 0. diabolicus sp.nov. 12'. L6bulo medio del metaesterno sin hilera de setas largas a lo largo del margen mesocoxal. Superficie de los elitros generalmente con microescultura fuerte(25 X ).Estrfas elitrales con puntuaciones circulares. Bolivia; Colombia ...... 13
13(12'). L6bulo medio del metaesterno generalmente con carina anterior longitudinal fina.Ancho mfnimo del 16bulo medio del metaesterno de alrededor de la mitad del maximo ancho del mesofemur. Colombia ...... 22. 0. lunicollis sp.nov. 13'. L6bulo medio del metaesterno sin carina anterior longitudinal fina.Mfnimo ancho del16bulo medio del metaestemo de alrededor de tres cuartos del ancho maxima del mesofemur. Bolivia ...... 23. 0. monilistriatus sp.nov.
14(8'). L6bulo medio del metaesterno generalmente con carina longitudinal fina anterior.Angulos anteriores del pronoto con puntuaciones casi confluentes.Pronoto del macho (Fig. 51) de los indivfduos mas grandes armada con 4 tuberculos, los laterales mucho mas grandes que los intemos. Costa Rica; Panama ...... 24 . 0. pseudodidymus sp.nov. 14' L6bulo medio del metaesterno sin carina longitudinal fina anterior.Angulos anteriores del pronoto con puntuaciones separadas por mas de su diametro. Pronoto del macho (Figs.43, 45) de los indivfduos mas grandes armada medialmente con 2 carinas longitudinales y 2 tuberculos lateralmente ...... 15
15(14'). Intervalos elitrales fuertemente impresos circa de las puntuaciones de las estrfas. Carinas pronotales del macho (Fig.43) paralelas en los indivfduos mas grandes. Mexico; Guatemala ...... 20.0. mexicanus Harold 18 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
15'. lntervalos elitrales ligeramente impresos circa de las puntuaciones de las estrias. Carinas pronotales del macho (Fig.45) formando un angulo de aproximadamente 35Q en los indivfduos mas grandes.Costa Rica ...... 21. 0. sextuberculatus sp.nov.
16(2'). L6bulo medio del metaestemo con setas conspicuas como en los 16bulos laterales.Protibia del macho con el diente apical mas o menos triangular, similar en aspecto al penultimo. Hembra con los angulos anteriores del pronoto granulares ...... 17 16'. L6bulo medio del metaestemo sin setas conspfcuas o con setas mas cortas que las de los 16bulos laterales.Protibia del macho con el diente apical mas delgado y apuntando hacia adelante.Angulos anteriores del pronoto de la hembra con puntuaciones ...... 19
17(16). Tamafiopequefio, menos de 12 mm.Superficie genal con al menos algunas areas toscamente rugosas. Brasil; Peru ...... 3. 0. laminifer Balthasar 17'. Tamafio grande, sobre 17 mm. Superficie genal lisa, a lo mas con unas pocas puntuaciones dispersas, ligeramente impresas anteriores a los ojos ...... 18
18(17'). Estrfas elitrales claramente crenuladas.Ecuador...... 1. 0. hadros sp.nov. 18'. Estrfas elitrales sin crenulaciones, carentes de puntuaciones claras. Ecuador ...... : ...... 2. 0. mag nus sp.nov.
19(16'). Tuberculo cefalico generalmente bajo (menor que el ancho del ojo) y localizado detras de la sutura clipeofrontal, la cual es carinada y arqueada. Macho con el borde posterior del femur posterior armado apicalmente con diente. Elitros nunca con microescultura fuerte ...... 20 19'. Tuberculo cefalico generalmente alto (mas que el ancho de un ojo) y localizado sobre la sutura clipeofrontal, la cual es poco marcada y algo angular medialmente (excepto 0. aequatorius con el tuMrculo detras de la sutura y con las puntuaciones de las estrfas redondeadas, bien impresas, mucho mas anchas que las estrfas y de Ecuador). Macho con el borde posterior del femur posterior no modificado...... 21
20(19). Estrfas elitrales del disco sin crenulationes.L6bulo medio del metaestemo generalmente con
unas pocas setas conspicuas. Colombia ...... 9. 0. kirschii Harold 20'. Estrfas elitrales del disco crenuladas.L6bulo medio del metaestemo sin setas largas, a lo mas con una pubescencia diminuta.Colombia ...... 10. 0. sanctaemartae sp.nov.
21(19'). Femur anterior con surco anteroventral bien delimitado apicalmente, casi alcanzando el apice, terminando abruptamente ...... 22 21'. Femur anterior con el surco anteroventral vagamente delimitado apicalmente, haciendose invisible gradualmente hacia el apice ...... 23
22(21). L6bulo medio del metaesterno (Fig.62) moderadamente ancho (L/Am = 2.6).Metaesterno plano o convexo en el disco. Pronoto sin puntuaciones conspicuas posteriormente. Venezuela ...... 4. 0. gilli sp.nov.
22'. L6bulo medio del metaestemo (Fig. 63) ancho (L/Am = 1.9). Metaestemo ampliamente concavo en el disco.Pronoto con puntuaciones conspicuas sobre una area pequefia posteral-
mente.Peru ...... 5. 0. ashei sp.nov.
23(21'). L6bulo medio del metaestemo (Fig.66) anteriormente con un margen, similar al borde lateral.Eliros con microescultura fuerte. Ecuador ...... 8. 0. aequatorius Bates 23'. L6bulo medio del metaestemo (Figs.64, 65) sin margen anteriormente, margen se desvanece medialmente.Elitros pulidos, a lo mas solo con microescultura fina ...... 24
24(23'). Estrfas elitrales con puntuaciones en forma de fovea profunda en el declive apical (especialmente las estrias 1 y 2). Coxa posterior con surco medio puntuado.Ecuador ...... 7. 0 . politus sp.nov. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 19
24'. Estrfas elitrales con puntuaciones similarmente impresas sobre el declive apical y el disco. Coxa posterior con el surco medio sin puntuaciones profunda.Colombia; Venezuela ...... 6. 0. hrevicollis Kirsch
25(1'). Mesepisterno plano, sin carina paralela a la sutura posterior. Pronoto sin ninguna indi caci6n de surco longitudinal o depresi6n posterior a lo largo de la lfnea media (subgenero Planontherus nov.) ...... 26 25'. Mesepisterno con carina paralela a la sutura posterior. Pronoto generalmente con surco longitudinal claro o con depresi6n posterior a lo largo de la lfnea media (subgenus Ontherus
s.str.) ...... 27
26(25). L6bulo medio del metaesterno (Fig.67) con algunas setas conspfcuas similares a las del mesosterno.Argentina; Bolivia; Peru ...... 25. 0. hridgesi Waterhouse 26'. L6bulo medio del metaesterno sin trazas de pubescencia Peru ..... 26. 0. rectus sp.nov.
27(25'). Sutura clipeofrontal sin carina, sin trazas de tuberculos.Argentina; Bolivia; Brasil;Paraguay ...... 49. 0. digitatus Harold 27'. Sutura clipeofrontal con tuberculo o con carina, si es carinada, la carina es ampliamente arqueada y baja, o ampliamente arqueada con 1 6 2 tuberculos centrales, o amliamente arqueada y trilobulada. T6rax generalmente similar en ancho a los elitros, si es mas ancho, entonces el tamafio es de menos de 10 mm y la sutura clipeofrontal con 1 6 2 tuberculos (comparar ademas la alternativa 27") ...... 28 27". Sutura clipeofrontal fuertemente carinada, carina semicircular, trapezoidal o cuadrangular, carina algunas veces cuadrilobulada o con dos dentfculos ampliamente espaciados. T6rax generalmente claramente mas anchos que los elitros ...... 36
28(27). Esternitos abdominales setosos medialmente, setas similares a las de la porci6n lateral del metaesterno (excepcionalmente los esternitos abdominales aparecen lisos medialmente pero entonces el borde posterior de la metacoxa es finamente serriforme.Este caracter se ve mejor cuando la coxa posterior es rotada hacia adelante para exponer el margen posterior) ... 29 28'. Esternitos abdominales sin setas medialmente, a lo mas con una hilera de setas cortas a lo largo del borde posterior de la metacoxa.Borde posterior de la metacoxa nunca finamente
serriforme...... 45
29(28). Sutura clipeofrontal tuberculada, a lo mas con carina muy fina no mas ancha que las puntuaciones cercanas ...... 30 29'. Sutura clipeofrontal solo con carina ...... 35
30(29). Borde posterior de la metacoxa parejo. La mayorfa de las puntuaciones en los lobulos laterales del metaesterno ovales, cada uno con el borde posterolateral desvanecido .. .. 31 30'. Borde posterior de la metacoxa finamente aserrado. La mayorfa de las puntuaciones de los l6bulos laterales del metaesterno redondeadas y claramente delimitadas ...... 34
31 (30). Margen pigidial generalmente completo. L6bulo medio del metaesterno a lo mas con setas diminutas.Argentina; Bolivia; Brasil; Colombia; Ecuador; Guayana Francesa; Guyana; Paraguay; Uruguay; Venezuela ...... 29. 0. appendiculatus (Mannerheim) 31 '. Margen pigidial incompleto, su borde interno poco evidente apicalmente.L6bulo medio del metaesterno generalmente con setas anteriormente.Especies con ambito de distribuci6n re ducido, menos frecuentemente encontradas ...... 32
32(31 '). L6bulos laterales del metaesterno densamente cubiertos con pubescencia larga. Borde posterior del metafemur sin setas. Tamafio generalmente sobre los 18 mm. Argentina ...... 34. 0. grandis Luederwaldt 32'. L6bulos laterales del metaesterno escazamente pubescente, pubescencia gradualmente mas corta hacia el margen lateral.Borde posterior del metafomur con pocas setas cortas.Tamafio generalmente menor de 15 mm. Brasil; Guayana Francesa; (registro dudoso de Colombia) ...... 33 20 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
33(32'). Maxima longitud del ultimo segmento abdominal (Figs. 2, 3) casi igual a la minima longitud combinada de los restantes segmentos abdominales. Brasil ......
...... ·...... 30. 0. ulcopygus sp.nov. 33'. Maxima longitud del ultimo segmento abdominal (Figs. 4, 5) casi 2 veces la minima longitu combinada de los restantes segmentos abdominales. Brasil; Guayana Francesa ...... 31. 0. podiceps Harold
34(30'). Esternitos abdominales con microescultura entre las puntuaciones. Bolivia; Brasil; Colom bia; Costa Rica; Ecuador; Mexico; Panama; Paraguay; Peru...... 32. 0. azteca Harold 34'. Esternitos abdominales lisos entre las puntuaciones. Guyana; Guayana Francesa...... 33. 0. cambeforti sp.nov.
35(29'). Dorso con lustre verde metalico brillante.Ufias tarsales fuertemente alargadas con forma de J. Brasil ...... 36. 0. virescens (Lucas) 35'. Dorso pardo oscuro, sin lustre metalico. Ufias tarsales mucho mas pequefias que el ultimo segmento, no modificadas.Brasil; Peru ...... 35. 0. carinifrons Luederwaldt
36(27"). Proesterno posteriormente a la procoxa, completamente pubescente, raramente con la pubescencia restringida solo a las porciones laterales...... 37 36'. Proesterno, detras de la procoxa, sin setas y liso, a lo mas con unas pocas setas erectas a lo largo del margen basal ...... 41
37(36). Carina clipeofrontal semicircular o trapezoidal, porci6n media siempre mas elevada.Longi tud combinada de la cabeza y el pronoto nunca igual en longitud a los elitros. Bolivia... 38 37'. Carina clipeofrontal cuadrangular, angulos laterales generalmente elevados. Longitud combinada de la cabeza y el pronoto casi igual en longitud a los elitros.Colombia; Costa
Rica; Panama; Peru; Venezuela ...... 39
38(37). Proestemo, detras de la procoxa, igualmente y ampliamente cubierto con setas Jargas erectas. Profemur del macho no modificado.Protibia del macho con la distancia entre el diente pen ultimo y el apical mas de dos veces la distancia de entre el antepenultimo y penultimo dientes.Bolivia ...... 38. 0. raptor sp.nov. 38'. Proestemo, detras de la procoxa, con pubescencia ausente en el disco. Profemur del macho aplanado en el cuarto basal. Protibia del macho con los dientes aproximadamente equidistantes. Bolivia ...... 39. 0. planus sp.nov.
39(37'). Longitud combinada de la cabeza y el pronoto mas corta que la longitud de los elitros. Margen basal del pronoto ligeramente mas ancho medialmente ...... 40 39'. Longitud combinada de la cabeza y el pronoto semejante en longitud a los elitros. Margen basal del pronoto no expandido medialmente. Colombia; Costa Rica; Panama
...... 42. 0. brevipennis Harold
40(39). Protibia del macho siempre con un cuarto diente apical vestigial.Brasil; Colombia; Guayana
Francesa; Venezuela ...... 40. 0. lichyi Martinez 40'. Protibia del macho completamente sin el cuarto diente apical.Peru . 41. 0. edentulus sp.nov.
41(36'). Carina clipeofrontalcuadrangular o trapezoidal, al menos con el borde anterior mas o menos
recto en indivfduos pequefios ...... 42 41'. Carina clipeofrontal semicircular ...... 44
42(41). Estrfas elitrales dispersamente microlabradas, puntuaciones solo ligeramente mas anchas que la estrfa. Carina clipeofrontal cuadrada, ancha, la mitad del ancho de la cabeza en los individuos mas grandes. Protibia del macho con el diente apical truncado en el apice. Metafomur del macho con el borde ventroposterior siempre completo y mas o menos sobresaliente posteriormente en la mitad basal. Brasil ..... 43.0. rectangulidens sp.nov. 42'. Estrfas elitrales con microescultura moderada, puntuaciones mucho mas anchas que la estrfa. Carina clipeofrontal a lo mas casi cuadrada, mas o menos trapezoidal.Borde anterior menos A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 21
de la mitad del ancho de la cabeza.Protibia del macho con el diente apical agudo.Metafemur del macho con la carina ventroposterior poco evidente, a lo mas recta en la mitad basal en indivfduos muy pequefios ...... 43
43(42'). Ufias tarsales (Fig. 6), cuando estan totalmente extendidas, al menos un tercio tan largas como el ultimo segmento tarsal. T6rax conspicuamente mas ancho que los elitros. Pronoto al menos con depresi6n ligera precedida por una protuberancia transversal anteriormente. Argentina; Brasil; Paraguay ...... 44. 0. carinicollis Luederwaldt 43'. Ufias tarsales (Fig. 7), cuando estan totalmente extendidas, a lo mas un cuarto tan largas como el ultimo segmento tarsal. T6rax casi tan ancho como los elitros. Pronoto convexo anteriormente.Argentina; Brasil ...... 45. 0. zikani Luederwaldt
44(41 '). Carina clipeofrontal entera medialmente.Margen anterior del 16bulo medio del metaesterno (Fig.8) sobresaliente en quilla longitudinal en los machos, reducido a una carina amplia y abruptamente terminada en las hembras (Fig.9). Brasil ...... 46. 0. lohifrons sp.nov. 44'. Carina clipeofrontal emarginada medialmente. Margen anterior del 16bulo medio del metaestemo (Fig. 10) sobresaliente en proceso grande y c6nico en los machos, en indivfduos pequefios el apice es truncado, margen siempre desvanecido, reducido a una carina baja y arqueada con la porci6n central emarginada en las hembras (Fig.11 ) .Brasil ...... 47. 0. cephalotes Harold
45(28'). Sutura clipeofrontal con tuMrculo o carinado con 1 6 2 tuMrculos contfguos 6 1 carina trilobulada ...... 46 45'. Sutura clipeofrontal ampliamente arqueada y con 1 carina ...... 53
46(45). Sutura clipeofrontal con carina trilobulada. Margen anterolateral de la metacoxa muy
agrandada y con microestructura en forma de estrfas ...... 47 46'. Sutura clipeofrontal con 1 tuberculo o carina con 1 6 2 tuberculos contfguos medialmente.
Margen anterolateral de la metacoxa no modificado ...... 49
47(46). Porci6n anterolateral del metaestemo con la carina transversal mas anterior claramente mas alla del borde lateroposterior del area microestructurada. Femur posterior del macho lobulado en sus dos tercios basales, borde distal del 16bulo siempre abruptamente terminado, en los machos mas grandes el borde distal es excavado.Metaestemo longitudinalmente con surcos longitudinales en el disco. Brasil ...... 58. 0. erosus Harold 47'. Porci6n anterolateral del metaestemo con la carina transversal mas anterior mas o menos en lfnea con el borde lateroposterior del area finamente estrigulada.Femur posterior del macho aguda y fuertemente carinado en la mayorfa de su longitud, carina nunca terminada abruptamente en los dos tercios apicales. Metastemo foveolado en el disco ...... 48
48(47'). L6bulo medio del metaestemo ancho (L/Am = 2.0). L6bulo lateral del metaesterno generalmente con 6 a 8 carinas claramente visibles. Metafemur del macho carinado a todo su largo. Brasil ...... 56. 0. insolitus sp.nov.
48'. L6bulo medio del metaesterno moderadamente ancho (L/mW = 2.6). L6bulo lateral del metaesterno generalmente con menos de 5 carinas transversales claras. Metafemur del macho carinado en la mitad basal. Brasil ...... 57. 0. stridulator sp.nov.
49(46'). Longitud del 16bulo medio del metaestemo menos de dos veces su maximo ancho (similar a Fig.12). Porci6n posterolateral del proepisterno densa e igualmente pubescente; elitro y base del pronoto sin lustre metalico ...... 50 49'. Longitud del 16bulo medio del metaestemo 2 veces su maximo ancho (similar a Fig.13). Porci6n posterolateral del proepistemo con I hilera de setas paralelas al borde lateral, algunas veces con unas pocas setas suplementarias.Elitros y base del pronoto generalmente con lustre verde metalico ...... 52
50(49). Pigidio muy ancho, AIL = 2.7 en macho 6 2.8 en hembra; puntuaciones moderadas, umbilicadas y profundamente impresas. Tamafio siempre menor de 12 mm. Argentina; Bolivia; Brasil; Paraguay; Peru; Venezuela ...... 48. 0. aphodioides Burmeister 22 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA N0.170
50'. Pigidio ancho, AIL = 2.1 en macho 6 2.3 en hembra; puntuaciones pequefias, sin umbilicationes y poco impresas. Tamafio generalmente sobre los 12 mm. Distribuci6n diferente ...... 51
51(50'). Borde posterior del femur a lo mas con pocas setas cortas y finas apicalmente. Femur posterior del macho (Fig.7 4) con superficie ventral con surcos finospor toda la parte anterior. Argentina; Bolivia; Brasil; Colombia; Ecuador; Guayana Francesa; Guyana; Paraguay; Surinam; Trinidad; Uruguay; Venezuela ...... 28. 0. sulcator (Fabricius) 51'. Borde posterior del femur posterior generalmente claramente pubescente a todo lo largo (pubescencia se ve mejor de vista lateral).Femur posterior del macho (Fig.73) sin surcos finos a lo largo del borde anterior, o a lo mas con surco poco evidente en la mitad apical. Argentina; Bolivia; Brasil; Colombia; Ecuador; Peru; Venezuela .. 27. 0. pubens sp.nov.
52(49'). Carina clipeofrontal con 1 tuberculo.Borde anterior del femur con muesca profunda en macho.Brasil ...... 50. 0. elegans Luederwaldt 52'. Carina clipeofrontal con 2 tuberculos contfguos.Borde anterior del femur sin modificaci6n en macho.Bolivia; Brasil; Paraguay; Uruguay ...... 51. 0. androgynus sp.nov.
53(45'). Margen pigidial bien definido apicalmente ...... 54 53'. Margen pigidial desvanecido apicalmente ...... 55
54(53). Porci6n posterolateral del proepisterno densa y parejamente pubescente.Proesterno densamente cubierto con pubescencia larga y erecta. Brasil...... 37. 0. irinus Balthasar 54'. Porci6n posterolateral del proepisterno con 1 hilera de setas parelelas al borde lateral; proestemo con setas dispersas e inclinada. Uruguay ...... 54. 0. atlantidis sp.nov.
55(53'). Pigidio abultado transversalmente basalmente; borde posterior del metatrocanter completamente surcado, surco con al menos 6 setas largas; borde anterior del metafemur angularmente sobresaliente basalmente en macho.Brasil ... 55. 0. dentatus Luederwaldt 55'. Pigidio mas o menos plano; borde posteror del metatrocanter poco surcado, surco con menos de 5 setas largas; borde anterior del metafemur modificado en el apice en macho ...... 56
56(55'). Pronoto cubiero parejamente de puntuaciones.Argentina; Brasil; Paraguay ...... 52. 0. erosioides Luederwaldt 56'. Pronoto muy finamente punteado en el disco, puntuaciones mucho mas pequefias que sobre el angulo anterior.Brasil ...... 53. 0. amplector sp.nov.
GENUS ONTHERUS ERICHSON, 1847
Ontherus Erichson, 1847: 107; 1848b: 761; Harold, 1867: 96; 1868a: 82; 1868b: 80; Gemminger and Harold, 1869: 1008; Harold, 1869a: 59; 1869b: 98; l 869c: 503; Kirsch, 1871: 356; Burmeister, 1874: 126; Harold, 1875a: 64; 1875b: 181; 1880: 21; Bates, 1887: 50; 1891: 25; Waterhouse, 1891: 356; Bruch, 1911: 186; Gillet, 1911a: 319; 1911b: 57; Bruch, 1915: 541; Luederwaldt, 1930: 105; 1931: 363; Balthasar, 1938: 220; Paulian, 1938: 233; Pessoa and Lane, 1941: 456; Blackwelder, 1944: 206; Lange, 1947: 310; Martinez, 1947: 45; Pereira, 1955: 461; Roze, 1955: 44; Pereira and d'Andretta, 1955: 251; Martinez, 1959: 69; Martinez and Pereira, 1960: 79; Pereira and Martinez, 1960: 49; Howden and Young, 1981: 121; Halffter and Edmonds, 1982: 89.
Type species: Scarabaeus sulcator Fabricius: Designated by Luederwaldt, 1931: 364.
DIAGNOSIS. Presence of sensory pit (Fig. 15) on antenna! segment 7 and 8, combined with presence of intervening setae (Fig. 14) on ventrnmedial carina of anterior tibiae, extreme posterior position of posterior coxae reducing median width of abdominal segments (Fig. 2-5), and absence of transverse carinae on median and posterior tibiae separates Ontherus from all closely related genera. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 23
REDESCRIPTION. Body. Oblong, moderately to strongly convex, dorsum usually entirely glabrous, or with at most minute setae. Size. Small to large (5-23 mm). Head. Anterior edge feebly to moderately bisinuate medially, never distinctly bidentate. Clypeus marginate anteriorly, margin usually wider in females. Clypeofrontal suture usually with conical process(es) and/or carina or otherwise modified (exceptsimple in 0. digitatus). Eyes large and always visible fromabove, ventral area large and globose. Posterior edge of ocular canthus never reaching occipital margin. Anteroventral portion of clypeus with short trans verse ridge medially, in few species ridge dentate medially. Labium transverse, distinctly emarginateanteriorly, surface rugose and moderately to densely coated with long erect setae throughout. Labial palpi 3-segmented, first segment oval and larger than second, with long setae, second segment broadly oval to rounded, with setae similar to first segment, third segment slender, subcylindrical and glabrous. Antennae 9-segmented, club 3-segmented, segments 7-9 elongate, segments 7 and 8 with deep pit that opens on external surface. Pronotum. Moderately to strongly convex, variously modified, especially in males of species belonging to Hadros, Laminifer, Alexis and Mexicanus groups. Lateral declivities with feeble to pronounced reniform gibbosity and usually, small depressed area above, lacking any distinct lateral ridge as in Copris (except for 0. lamin(fer). Edges with distinct marginal bead. Elytra. With 9 striae including 1 along epipleural margin, but eighth stria atrophied on basal third and partly to entirely fused to ninth on posterior two-thirds. Striae narrow to wide and slightly to deeply impressed, usually distinct throughout, punctures varying frombarely discernablefrom stria to large, rounded and umbilicate. Intervals slightly to moderately convex. Epipleura narrow, usually with aligned row of setae anteriorly. Thoracic sterna. Proepisternum lacking depression to receive anterior femur anteriorly, with fine transverse carina on anterior third not reaching lateral edge, medioposterior surface smooth. Prosternum with posteror edge straight. Mesosternum transverse, disc transversly impressed, surface usually coarsely punctate and pubescent. Mesometasternal suture vari able, straight or obtusely to acutely angular medially, sometimes longitudinal carina present on mesosternum. Metastemum with median lobe usually much longer than wide. Median coxal cavities subparallel, distinctly marginate. Legs. Anterior tarsi always present. Anterior spur movable, variable in shape and usually smaller in males. Anterior tibiae with lateral edges variably dentate, but usually quadridentate. Apical edge always obliquely truncate. Ventromedial carina (Fig. 14) with at least 3 intervening setae, usually 10 or more. Middle and posterior tibiae rectangular in cross-section, expanded apically and lacking transverse carina on lateral margins. Middle tibiae with 2 elongate spurs, spur adjacent to tarsus almost twice as long as other spur. Posterior tibiae with single long spur. Middle and posterior tarsi long, each subequal in length to tibia, segments triangular in shape, decreasing in length toward apex, apical segment with small but distinct membranous projection ventrally between claws. Claws usually small and unmodified, except large and "J" -shaped in 0. virescens and retractable in some species of Brevipennis group. Abdomen. Sternites (Figs. 2-5) strongly reduced in length medially, fused. Pygidium wider than high, marginate or not apically. Male genitalia. Parameres usually symmetrical, rarely slightly asymmetri cal, internal sac with double flagellum (Fig. 16), lambda (Fig. 17), sigmoid (Figs. 259-314) and occasionally 1 or 2 accessory sclerites (Figs. 18, 19) and usually small area covered with velum. Female genitalia. Spermatheca (Fig. 20) simple, V-shaped.
SUBGENUS CAELONTHERUS NOV. Type species: Ontherus alexis (Blanchard)
ETYMOLOGY. Caelontherus, (heavenly + Ontherus) Latin adjective, refers to species being usually found at higher elevations throughout Central and South America. 24 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
DESCRIPTION. Body usually elongate, almost parallel-sided and convex dorsally. Head variably armed with single tubercle, or pair of closely set tubercles, or transverse horn, or variously shaped and elongate horn. Pronotum usually slightly declivous anteriorly, or variously armed with swellings, tubercles and/or carinae, convex in small individuals. Elytra convex on disc, almost flat in species of Brevicollis group; striae variously shaped, eighth stria indistinguishable fromninth, except on apical declivity in some species, distinct in species of Brevicollis group; intervals variously shaped and sculptured. Mesosternum lacking short longitudinal carina posteriorly. Mesepimera always carinate parallel to posterior edge. Median lobe of metastemum straight or slightly arcuate medially. Anterior tibia with 3 or 4 teeth in males, 4 in females. Posterior femur simple, except in species of Kirschii group. Parameres in lateral view triangular, variously modified apically in species of Mexicanus group.
HADROS GROUP
DIAGNOSIS. Large body size (19-23 mm), presence of granules on head and anterior portion of pronotum, and presence of fine granules on apex of parameres, separates the following 2 species from all others in genus.
1. Ontherus (Caelontherus) hadros sp.nov. (Figs. 21, 22, 91-93, 259; Map 1)
ETYMOLOGY. Hadros, (large) a Greek adjective, refers to the size of this species.
DESCRIPTION. Colour. Body reddish brown to dark brown, venter and elytral margin slightly lighter. Pubescence dark yellowish. Length. 19-23 mm. Head (Figs. 21, 22). Anterior edge broadly arcuate and almost flat, bisinuate medially, notched at clypeogenal junction. Clypeus transversely and rather coarsely wrinkled on apical fourth with remainder weakly wrinkled in males or coarsely wrinkled throughout in females. Clypeofrontal suture arcuate, with laterally compressed horn, apex transverse and truncate in larger males, or finely carinate with small tubercle posterior to suture in femalesand smaller males. Anterior genal edge subequal in length to lateral, genal surface almost entirely smooth in males or transversely and finely wrinkled in females. Vertex smooth basally, weakly granulate in males, or granulate in females anterior to eyes. Pronotum (Figs. 21, 22). Transverse, W/L
= 1.5, anterior edge slightly sinuate medially; punctation minute on disc and most of surface in larger males, or minute on disc and becoming granulate on anterior and lateral declivities in females and smaller males. Anterior half widely excavated, posterior edge of excavation delimited by sinuous bulge, sublobate laterally in larger males, or convex, except for 1 small flatarea in females and 3 in small males. Elytra. Subquadrate, L/W = 1.1. Striae moderately wide, alutaceous, shallowly impressed throughout their length, punctures almost twice the width of stria, deeper than stria, separated by their diameter. Elytral intervals slightly convex, finely and evenly punctate. Thoracic sterna. Proepistemum with posterolateral portion moderately pubescent, external third almost smooth. Prosternum behind procoxa densely covered with long pubescence. Mesosternum deeply impressed posteriorly, densely punc tate, pubescence fine and short, few rows of coarse and long setae along anterior edges of mesocoxa. Mesepistemum moderately punctate and strongly carinate. Metastemum with disc finely punctate, deeply impressed in males or with elongate oval depression on posterior two-thirds in females; lateral lobes densely punctate and strongly alutaceous throughout, anteriormost punctures each with anterior edge finelycarinate, posterior third with punctures simple and rounded; median lobe wide, L/mW = 1.6, lateral margins convergent throughout their length, surface with setiferous punctures shallowly impressed and restricted to anterior half and lateral portion, separated by approximately their diameter, surface between entirely and finely alutaceous, junction with mesostemum straight and finely carinated. Legs. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 25
/
• 0. hadros
*o. magnus • 0. laminifer
MAP 1. Known distribution of Ontherus hadros, 0. magnus and 0. laminifer.
Anterior legs with tibia quadridentate, similar in males and females, ventromedial carina lacking intervening setae on basal third, apical spur acute and inwardly bent. Posterior legs unmodified. Abdomen. Sternites 2-5 alutaceous, with dense and setiferous punctures on anterior two-thirds laterally, stemite 6 finely punctate and alutaceous, suture between stemites 5-6 unmodified. Pygidium, W/L = 2.1 in males or 2.6 in females, punctures fine, shallowly impressed and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 91-93). Tubus approximately 2.5 times as long as parameres, apicoventral portion flattened and sulcate. Parameres in lateral view with apices rounded, ventral portion obliquely truncate, apical third finely granulate, dorsal portion with 2 small denticles directed posteriorly on apical fourth. Sigmoid sclerite of internal sac as in Figure 259.
MATERIAL STUDIED. 2 a a, 1 9 (HAHC, MNHN).
Holotype O' (MNHN): Macas, Ecuador/ Museum Paris, ex. coll., R. Oberthur/ HOLOTYPE, Ontherus hadros, F. Genier, 1992. Dissected.
Allotype 9 (MNHN): Zarayacu ( =Sarayacu, Napo?), Ecuador or./ Museum Paris, ex. coll., R. Oberthur/ ALLOTYPE, Ontherus hadros, F. Genier, 1992. Dissected.
Paratypes: ECUADOR: Napo: 27km. N. W. Baeza, 2-6.III.1976, 2700m., S. Peck, dung tp. 10, (1 a, HAHC).
2. Ontherus (Caelontherus) magnus sp.nov. (Figs. 94-96, 260; Map 1)
ETYMOLOGY. Magnus, (large) a Latin adjective, refers to the size of this species.
DESCRIPTION. Colour. Body dark brown, ventrum and elytral margin dark reddish brown. Pubescence reddish. Length. 19-21 mm. Head. Anterior edge broadly arcuate and 26 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 almost flat, bisinuate medially, notched at clypeogenal junction. Clypeus transversely and rather coarsely wrinkled on apical fourth in males and throughout in females, remainder weakly wrinkled in males. Clypeofrontal suture arcuate, with laterally compressed horn, apex acute to truncate in males or finely carinate with small tubercle posterior to suture in females. Anterior genal edge subequal in length to lateral, genal surface finely wrinkled on anterior half in males or coarsely wrinkled in females. Vertex smooth basally, weakly granulate in males and granulate in females anterior to eyes. Pronotum. Transverse, W/L = 1.5, anterior edge slightly sinuate medially; punctation minute on disc, distinctly larger on posterior angles, anterior half of declivity almost completely smooth, remainder finely granulate in males or moderate on disc and becoming granulate on anterior and lateral declivities in females. Anterior third widely excavated, posterior edge of excavation delim ited by sinuous bulge in males or convex, except for 1 small depression in females. Elytra. Subquadrate, L/W = 1.1. Striae wide, finely alutaceous, shallowly impressed throughout their length, punctures almost equal to width of stria, separated by 1-2 diameters. Elytral intervals slightly convex, finely and evenly punctate, interval 2-3 alutaceous on apical declivity. Thoracic sterna. Proepisternum with posterolateral portion moderately pubes cent, external third smooth. Prosternum behind procoxa densely covered with long pubes cence. Mesosternum deeply impressed posteriorly, densely punctate, pubescence fine and short, few rows of coarse and long setae along anterior edges of mesocoxa. Mesepisternum moderately punctate and carinate. Metasternum with disc finely punctate, deeply impressed in males or with elongate oval depression on posterior half in females; lateral lobes densely punctate and strongly alutaceous throughout, anteriormost punctures each with anterior edge finely carinate, posterior third with punctures simple and rounded; median lobe wide, L/mW = 1.4, lateral margins convergent throughout their length, surface with setiferous punctures shallowly impressed and restricted to anterior half and lateral portion, separated by approxi mately their diameter, entirely and finely alutaceous, junction with mesosternum straight and finely carinate. Legs. Anterior legs with tibia quadridentate, similar in males and females, ventromedial carina lacking intervening setae on basal third, apical spur acute and inwardly bent. Posterior legs unmodified. Abdomen. Sternites 2- 5 finely alutaceous, with dense and setiferous punctures on anterior half laterally, sternite 6 finely punctate and alutaceous, suture between sternites 5-6 unmodified. Pygidium, W/L = 2.1 in males or 2.4 in females, punctures fine, shallowly impressedand almost evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 94-96). Tubus approximately 2.0 times as long as parameres, apicoventral portion deeply sulcated. Para meres in lateral view with apices rounded, ventral portion obliquely truncate, apical third finely granulate. Sigmoid sclerite of internal sac as in Figure 260.
MATERIA L STUDIED. 1 ry, 1 9 (CMNC, ZMHB).
Holotype O' (ZMHB): Ecuador, Fritsche V./ Zool. Mus., Berlin/ HOLOTYPE, Ontherus magnus, F. Genier, 1992. Dissected.
Allotype 9 (CMNC): ECUADOR: PICHINCHA, 2lkm. E. Tandapi ( =CornejosAstorga), 2600m., 7-14.V I.1976, S.Peck, moss for. dng. tp. 7./ ex. collection, H. and A. Howden/ ALLOTYPE, Ontherus magnus, F. Genier, 1992. Dissected.
LAMINIFER GROUP
DIAGNOSIS. Small body size (7-11 mm), presence of granules on head and pronotum and absence of fine granules on parameres, separates the following species from all others in genus. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 27
3. Ontherus (Caelontherus) laminiferBalthasar, 1938 comb.nov. (Figs. 23, 24, 97-99, 261; Map 1)
Ontherus lamin!fer Balthasar, 1938: 221 (description) Ontherus laminifer Balthasar: Blackwelder, 1944: 206 (list) Ontherus laminifer Balthasar: Pereira, 1955: 462 (comment)
REDESCRIPTION. Colour. Body uniformly dark brown to black. Pubescence fulvous. Length. 7-11 mm. Head (Figs. 23, 24). Margin almost semicircular, upturned, bisinuate and broadly notched medially, clypeogenal junction finely notched. Clypeus transversely and coarsely wrinkled, except narrow band along anterior edge of hornin males or becoming granulate in females. Clypeofrontal suture broadly arcuate, almost straight, with transverse and backward-curved lamella covering two-thirds of distance between eyes, lateral edges parallel on basal fourth and gradually narrowing toward apex in larger males, reduced to lobe broadly notched apically in smaller males, or low carina interrupted medially in females. Anterior genal edge distinctly longer than lateral, genal surface strongly and transversely wrinkled on anterior half, remainder strongly granulate in both sexes. Vertex completely smooth except finely punctate small area adjacent to eyes and weakly granulate posterior surface of lamella in males, or densely granulate, except narrow band along posterior edge, in females. Pronotum (Figs. 23, 24). Strongly transverse, W/L = 1.7, anterior edge strongly widened on median third, deeply and widely emarginate behind eyes, gap of emargination membranous in males or widened medially on median tenth, barely emarginate behind eyes and narrowly membranous behind head insertion in females; disc smooth, granulation restricted to lateral declivities in males or anterior and lateral declivities in females, narrow band of fine granules along posterior edge. Anterior third declivous behind head insertion, upper edge in line with inner eyes margin, with 2 small triangular tubercles, area below tubercles distinctly depressed in larger males, declivity restricted to anterior fourth and tubercles reduce to sinuous bulges in smaller males or convex in females. Elytra. Subquad rate, L/W = 0.9. Striae moderately wide, finelyalutaceous, shallowly impressed throughout their length, punctures less than twice the width of stria, separated by approximately twice their diameter. Intervals almost flat, punctures uneven in diameter, almost evenly distributed, finelyalutaceous along stria, basal and apical portion. Thoracic sterna. Proepistemum with posterolateral portion moderately pubescent, lateral margin with row of coarse setiferous punctures. Prostemmp behind procoxa densely pubescent, posterior edge densely fringed with yellowish seta. Mesostemum deeply impressed posteriorly, punctation more dense on disc, pubescence fine and short, few long setae along anterior edges of mesocoxa. Mesepisternum finely punctate, strongly alutaceous and carinate. Metasternum with disc finely punctate, with shallow longitudinal sulcus on posterior two-thirds; lateral lobes moderately punctate, punctures shallow, weakly delimited, transverse, anteriormost punc tures each with anterior edge carinate, pubescence short and apposed anteriorly, surface strongly alutaceous, more so on anterior half; median lobe wide, L/mW = 1.4, lateral edges convergent throughout their length, surface with punctures restricted to anterior half, shallowly impressed, separated by approximately their diameter, finely and unevenly alu taceous on anterior third, junction with mesostemum straight, carina atrophied. Legs. Anterior legs with tibia quadridentate and similar in both sexes, in larger individuals apical tooth approximately half total length of protibia, ventromedial carina facking inter vening setae on basal half, apical spur acute and inwardly bent. Posterior legs unmodified. Abdomen. Stemites 2-5 alutaceous, with dense and setiferous punctures throughout, stemite 6 setiferously punctate and alutaceous throughout, suture between stemites 5-6 unmodified. Pygidium, W IL = 2.1 in males or 2.4 in females, punctures fine, shallowly impressed and slightly more numerous on apical half, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 97 -99). Tu bus approximately 1.5 times as long as parameres, ventral portion flat. Parameres in lateral view with apices 28 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
rounded, ventral portion slightly obliquely truncated, surface smooth. Sigmoid sclerite of internal sac as in Figure 261.
MATERIAL STUDIED. 200" ct, 16 Q Q (AMNH, BMNH, CASC, ICCM, MCZC, MNHN, NMPC, WDEC).
TYPE MATERIAL Holotype O" (by monotypy)(NMPC): Amazonas, Manaos/ Coll. C. Felsche, Kauf 20, 1918/ TYPUS/ Ontherus laminifer, n. sp., Dr. V. Balthasar det./ Laminifer m./ Nat. Mus. Pragae, inv. 65458./ HOLOTYPE, Ontherus laminifer, Balthasar, 1938, vidit F. Genier, 1991. Holotype studied.
NON- TYPE MATERIAL Distribution: BRAZIL: Amazonas: Ega (=Tefe?); Japuyas (=Japura?); Tefe; Uypizanga (Rio Negro, 14 km. from Manaus) (90m). Para: Obidos; Santarem. Rondonia: Manoa; Porto Velho; Rio Madeira (Madeira-Mamore R. R. Co. Camp 39). PERU: Loreto: Iquitos (Mishuyacu); Middle Rio Ucayali. Madre de Dios: Iberia (150m). San Martin: Achinamisa.
Additional data: Specimens collected in May, August to December.
BREVICOLLIS GROUP
DIAGNOSIS. Straight or slightly arcuate, anterior edge of median lobe of mesostemum and large conical process located on clypeofrontal suture (slightly behind suture in 0. aequatorius) unique for both sexes. In males, slender and forwardly directed apical tooth of tibia, unmodified posterior femur and large, inflatable membranes on parameres, distinguishes the following 4 species.
4. Ontherus (Caelontherus) gilli sp.nov. (Figs. 62, 101, 102, 262; Map 2)
ETYMOLOGY. Gilli, a patronym in gratitude to Bruce D. Gill, collector of most of the specimens studied of this species.
DESCRIPTION. Colour. Body dark brown to black. Pubescence reddish brown. Length. 8-14 mm. Head. Anterior edge arcuate and almost flat, slightly sinuate medially, clypeo genal junction lacking indentation. Clypeus transversely and rather coarsely wrinkled on apical fourth in males or two-thirds in females, remainder weakly wrinkled to shallowly punctate. Clypeofrontal suture ill-defined, broadly arcuate, with large conical process medially on suture, reduced to swelling in small individuals. Anterior genal edge subequal in length to lateral, genal surface finelypunctate in males or transversely wrinkled on anterior half and remainder finely punctate in females. Vertex smooth on disc, remainder finely punctate. Pronotum. Transverse, W/L = 1.5; anterior edge slightly wider, narrowly mem branous and almost straight behind head insertion; disc almost smooth in larger males, punctures slightly larger on anterior half of pronotum. Anterior fifthslightly declivous behind head insertion, almost convex in small individuals. Elytra. Subquadrate, L/W = 1.0. Striae fine, almost smooth, moderately impressed throughout their length, punctures approximately 3 times as wide as stria on disc, separated by 3-4 diameters. Elytral intervals slightly convex, finely and evenly punctate, surface smooth. Thoracic sterna. Proepisternum with postero lateral portion rather scarcely pubescent, external third smooth. Prosternum behind procoxa densely covered with long pubescence on disc. Mesosternum impressed posteriorly, punc tation dense, except 2 small almost smooth areas laterally, pubescence fine and short, few long setae along anterior edge of mesocoxa. Mesepisternum densely punctate, punctures fine, moderately carinate. Metasternum with disc smooth, finely punctate in smaller A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 29
• O.gilli
*o. ashei • 0. aequatorius
MAP 2. Known distribution of Ontherus gilli, 0. ashei and 0. aequatorius.
individuals, with shallow longitudinal sulcus on posterior eighth; lateral lobes scarcely punctate, punctures oval and well-impressed, pubescence moderately long and obliquely oriented laterally, surface slightly alutaceous on anterior half, remainder smooth; median lobe (Fig. 62) moderately wide, L/mW = 2.6, lateral edges convergent on posterior three-fourths and on anterior fourth straight to slightly divergent, surface finely and evenly punctate and alutaceous, junction with mesosternum slightly arcuate and carinate. Legs. Anterior legs with tibia quadridentate in both sexes, apical tooth slender, straight and directed slightly obliquely outward in males or unmodified in females, ventromedial carina lacking intervening setae on basal half, apical spur slender, acute and apex slightly bent inwardly in males or strongly bent inwardly on apical third in females, Posterior legs unmodified. Abdomen. Sternites 2-5 smooth, with row of coarse punctures along anterior edge, few scattered and setiferous punctures laterally, stemite 6 finely punctate throughout, suture between stemites 5-6 deeply grooved. Pygidium, W/L = 1.3 in males or 1.5 in females,punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 100-102). Tubus approximately 2.3 times as long as parameres. Parameres in lateral view with apices rounded, ventral portion straight, dorsal portion widely membranous on basal two-thirds. Sigmoid sclerite of internal sac as in Figure 262.
MATERIAL STUDIED. 90' a, 7 9 9 (BDGC, CMNC, FGIC, HAHC, IZAV, MNHN).
Holotype a (CMNC): VENEZUELA: Tach., San Cristobal, 10-18 VIII 83, B. Gill, 1200m/ HOLOTYPE, Ontherus gilli, F. Genier, 1992. Dissected.
Allotype 9 (BDGC): Same data as holotype.
Paratypes: VENEZUELA: locality unspecified (1 a MNHN). Merida: ?Merida (1 O', 2 9 9 MNHN); 25kmN. W.Merida onJajiRoad(1800m),Chorrera Gonzales, 28.VI-3.VIII.89, 30 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 forest carrion tps, S. and J. Peck (2ct ct, 2 9 9 HAHC). Tachira: same data as holotype (2ctct, 1 9 BDGC) (1ct, 19 FGIC) (1ct IZAV ).
5. Ontherus (Caelontherus) ashei sp.nov. (Figs. 63, 103-105, 263; Map 2)
ETYMOLOGY. Ashei, a patronym in acknowledgement of James S. Ashe's important collecting efforts for beetles in Latin America.
DESCRIPTION. Colour. Body dark reddish brown. Pubescence fulvous. Length. 13 mm. Head. Anterior edge arcuate and upturned, bisinuate medially, clypeogenal junction lacking indentation. Clypeus transversely wrinkled on apical fourthin males, remainder moderately to coarsely punctate. Clypeofrontal suture finely carinate laterally, broadly arcuate, with moderate conical process medially on suture. Anterior genal edge longer than lateral, genal surface rather coarsely punctate in males. Vertex finely and sparsely punctate throughout.
Pronotum. Transverse, W/L = 1.8; anterior edge slightly wider, narrowly membranous and almost straight behind head insertion; disc finely punctate, punctures larger on anterior third of pronotum. Anterior fifth declivous behind head insertion, posterior portion delimited by bulge medially. Elytra. Subquadrate, L/W = 1.0. Striae fine,moderately impressed through out their length, punctures approximately 3 times as wide as stria on disc, separated by 4-7 diameters. Elytral intervals slightly convex, finely and evenly punctate, surface finely alutaceous. Thoracic sterna. Proepisternum with posterolateral portion moderately pubes cent, external third smooth. Prosternum behind procoxa moderately pubescent, with few long setae on disc. Mesosternum impressed posteriorly, punctation dense and coarse except 2 small and almost smooth areas laterally, pubescence fine and short, few long setae along anterior edge of mesocoxa. Mesepisternummoderately punctate, punctures fineto moderate, strongly carinate. Metasternum with disc smooth, finely punctate, deeply foveolate on anterior two-thirds; lateral lobes moderately punctate, punctures oval and well-impressed, pubescence moderately long and obliquely oriented laterally, surface strongly alutaceous on
anterior two-thirds, remainder smooth; median lobe (Fig. 63) wide, L/mW = 1.9, lateral edges convergent on posterior two-thirds, anterior third parallel to slightly divergent, surface finely and evenly punctate and alutaceous, junction with mesosternum slightly arcuate and finely carinate throughout. Legs. Anterior legs with tibia quadridentate, apical tooth slender and inwardly bent on apical half in males, ventromedial carina lacking intervening setae on basal two-fifths, apical spur spiniform, slightly bent inwardly in males, posterior tibia longitudinally impressed ventrally, edges of impression sharply delimited. Abdomen. Sternites 2-5 finely alutaceous, with row of coarse punctures along anterior edge, few scattered and setiferous punctures laterally, sternite 6 finely punctate throughout, suture between sternites 5- 6 with few deep punctures laterally. Pygidium, W/L = 2.1, punctures fine and evenly distributed, apical margin clearly delimited throughout its length. Male genitalia (Figs. 103-105). Tubus approximately 2.3 times as long as parameres. Parameres in lateral view with apices dentate ventrally and dorsally, ventral portion straight, dorsal portion widely membranous throughout. Sigmoid sclerite of internal sac as in Figure 263.
MATERIAL STUDIED . 1 ct (SEMC).
Holotype ct (SEMC): Peru, S. A., Aug. 3, 1936, F. Woytkowski, no. 3756/ Collected on way to Chachapoy as from San Idelfonso, Dept. Amazonas/ HOLOTYPE, Ontherus ashei, F. Genier, 1993. Dissected.
REMARKS. Ontherus ashei sp.nov. is described from a single male. It is closely related to 0. gilli sp.nov., but readily distinguishable fromit by the characters given in the key. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 31
6. Ontherus (Caelontherus) brevicollis Kirsch, 1871 comb.nov. (Figs. 59, 64, 106-108, 264; Map 3) 2 Ontherus brevicollis Kirsch, 1870 (1871): 356 (description) Ontherus brevicollis Kirsch: Harold, 1880: 23 (list and comment) Ontherus brevicollis Kirsch: Gillet, 191 lb: 57 (catalogue) Ontherus brevicollis Kirsch: Luederwaldt, 1931: 393 (key) Ontherus brevicollis Kirsch: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body uniformly dark reddish brown to black. Pubescence fulvous. Length. 11-18 mm. Head. Anterior edge arcuate, slightly upturned and bisinuate medially, clypeogenal junction finely notched. Clypeus transversely and coarsely wrinkled on anterior fourth in males or two-thirds in females, remainder smooth in males or asperulate in females. Clypeofrontal suture ill-defined, broadly arcuate, with large conical process medially on suture, reduced to finely tuberculate swelling in smaller individuals. Anterior genal edge slightly longer than lateral, genal surface finely punctate in males or coarsely wrinkled on anterior half and finely to coarsely punctate on remainder of surface in females. Vertex finely punctate throughout, except small area posterior to apex of conical process with coarse and dense punctures in larger females. Pronotum. Transverse, W/L = 1.7; anterior edge slightly wider, narrowly membranous and almost straight behind head insertion; disc almost smooth, punctation slightly larger on anterior angles in males or anterior angles and posterior edge of declivity in females. Anterior fourth declivous behind head insertion, almost convex in smaller individuals. Elytra. Subquadrate, L/W = 1.1. Striae moderately wide, almost smooth, shallowly impressed throughout their length, punctures approximately 3 times as wide as stria on disc, separated by 2-3 diameters. Elytral intervals slightly convex, finely and evenly punctate, finely alutaceous on disc and apical declivity. Thoracic sterna. Proepistemum with posterolateral portion rather scarcely pubescent in males or moderately pubescent in females, external fourth smooth. Prostemum behind procoxa densely covered with long pubescence. Mesosternum impressed posteriorly, punctation dense except 2 small smooth areas laterally, pubescence fine and short, few long setae along anterior edge of mesocoxa. Mesepisternum densely punctate, punctures of 2 different sizes, finely carinate. Metasternum with disc finely punctate, with wide and longitudinal sulcus on posterior half in males or fine longitudinal sulcus on posterior half in females, lateral lobes moderately punctate, punctures broadly oval to rounded, pubescence moderately long and obliquely oriented, surface alutaceous, more so anteri orly; median lobe (Fig. 64) moderately wide, L/mW = 2.2, lateral edges convergent on posterior four-fifths and parallel to slightly divergent on anterior fifth, surface finely and evenly punctate and alutaceous, junction with mesosternum straight and lacking carina. Legs. Anterior legs with tibia quadridentate in both sexes, apical tooth slender and inwardly bent on apical half in males (Fig. 59) or unmodified in females, ventromedial carina usually lacking intervening setae on basal half, apical spur spiniform straight and shorter than apical tooth in males or slender, inwardly bent on apical third and longer than apical tooth in females. Posterior legs unmodified. Abdomen . Sternites 2-5 finely alutaceous, with row of setiferous punctures along anterior edge, some scattered and larger setiferous punctures laterally, sternite 6 finely punctate throughout, suture be tween sternites 5- 6 finely grooved laterally. Pygidium, W/L = 1.9 in males or 2.3 in females, punctures fine and evenly distributed, apical margin clearly delimited through out its length in both sexes. Male genitalia (Figs. 106-108). Tubus approximately 2.5 times as long as parameres. Parameres in lateral view with apices truncate, ventral
2 Technically the description is on page 340. 32 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. brevicollis
• 0. politus
MAP 3. Known distribution of Ontherus brevicollis and 0. politus.
portion slightly concave, dorsal portion widely membranous on basal half. Sigmoid sclerite of internal sac as in Figure 264.
MATERIAL STUDIED. 145 a a, 127 9 9 (AMNH, BDGC, CMNC, CNCI, DEIC, HAHC, ISNB, IZAV, MZLU, MNHN, NMPC, SMFD, SMTD, USNM, ZMHB).
TYPE MATERIAL Holotype O' (SMTD): Bogota, m. Kirsch (green paper)/ Ontherus kirschi, Har., Dr.V.Balthasar det./ Staatl. Museum fiir, Tierkunde Dresden/ HOLOTYPE, Ontherus brevicollis, Kirsch, 1871, vidit F. Genier, 1991. Holotype studied.
REMARKS. I assume that this specimen is the holotype; the locality label is identical to the label of the holotype of Ontherus incisus (Kirsch), the specimen fits the original description in all respects and there are no indication of variation in the original Description.
NON-TYPE MATERIAL Distribution: BOLIVIA (doubtful): No locality. BRAZIL (doubtful): No locality. COLOMBIA: Antioquia: Santa Rosa de Osos. Boyaca: Muzo. Cauca: 1 lmi. E Silvia (3000m); 16km E Silvia (2750m); Popayan. Cundinamarca: Esmeralda; Pacho; Tecon dama Falls (30km S W Bogota); Viani. Distrito Especial: Agnas Largas (near Bogota); Bogota. Meta: Villavicencio. Narifio: Pasto. Norte de Santander: l 2k S Pamplona (3000m); 30km S Chinacota (2600m); 35km S Chinacota (3000m). ECUADOR: No locality. VENEZUELA: Merida: Hechicera; Monte Zerpa (2000m) (Merida); La Mucuy (lOkm E Tabay) (2450m); Los Chorros; Merida; Telef[erico]. Est[acion]. La Montana (2450m) (Merida). Tachira: 45km NE San Cristobal (2600m).
Additional data: Specimens were collected in cloud forests using flight intercept traps, and in dung and carrion traps set in forestsand montane forests. Specimens collected in February to August. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 33
7. Ontherus (Caelontherus) politus sp.nov. (Figs. 65, 109-111, 265; Map 3)
ETYMOLOGY. Politus, (glossy) Latin adjective, refers to the appearance of the dorsum of this species.
DESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 11-16 mm. Head. Anterior edge arcuate, slightly upturned and slightly sinuate medially, clypeogenal junction entire. Clypeus weakly wrinkled on anterior fourth in males or coarsely wrinkled on anterior three-fourths in females, remainder smooth in males or asperulate in females. Clypeofrontal suture with lacking carina in males or finely carinate in females, with large conical process medially on suture, reduced to finely tuberculate swelling in smaller individuals. Anterior genal edge subequal to lateral, genal surface smooth, finely punctate in smaller males or wrinkled on anterior half and punctate on remainder of surface in females. Vertex finely punctate throughout. Pronotum. Transverse, W/L = 1.7; anterior edge slightly wider, narrowly membranous and almost straight behind head insertion; disc smooth, finely punctate in smaller males, anterior and lateral declivities finely punctate in males or disc finely punctate and anterior half moderately punctate in females. Anterior fourth declivous on median third. Elytra. Subquadrate, L/W = 1.0. Striae fine, smooth, moderately impressed throughout their length, punctures rounded, deeply impressed on apical declivity, approximately 3 times as wide as stria, separated by 2-4 diameters on disc. Elytral intervals slightly convex, finely and evenly punctate. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent, external fourth smooth. Prosternum behind procoxa densely covered with long pubescence. Mesosternum impressed posteriorly, punctation dense, pubescence fine and short, a few long setae along anterior edge of mesocoxa. Mesepisternum moderately punctate, punctures rounded and of 2 different sizes, sharply carinate. Metasternum with disc finely punctate and with a shallow longitudinal sulcus; lateral lobes moderately punctate, punctures rounded posteriorly, becoming oval anteriorly, anteriormost punctures each with anterior edge carinate, pubescence moderately long and obliquely oriented, surface finely alutaceous, more so anteriorly; median lobe (Fig.
65) moderately wide, L/mW = 2.2, lateral margins convergent on posterior three-fourths and parallel to slightly divergent on anterior fourth, surface moderately punctate and alutaceous throughout, junction with mesosternum slightly arcuate, declivous and finely carinate. Legs. Anterior legs with tibia quadridentate in both sexes, apical tooth slender and inwardly bent on apical third or unmodified in females, ventromedial carina lacking intervening setae on basal half, apical spur spiniform, straight and shorter than apical tooth in males or slender, inwardly bent on apical third and longer than apical tooth in females. Posterior legs unmodified.Abdomen. Sternites2- 5 finely and scarcely punctate, with row of punctures along anterior edge, some scattered setiferous punctures laterally, stemite 6 finely punctate throughout, suture between sternites 5- 6 with row of coarse and deep punctures. Pygidium, W /L = 1.9 in males or 2.3 in females, punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 109-111). Tu bus approximately 1.3 times as long as parameres. Parameres in lateral view with apices truncate, ventral portion straight and subdentate apically, dorsal portion largely membranous on median half. Sigmoid sclerite of internal sac as in Fig ure 265.
MATERIAL STUDIED. 17 CJ' CJ', 10 Q Q (CMNC, FGIC, HAHC, MNHN, QCAZ).
Holotype CJ' (CMNC): ECU[ADOR]: NAPO, 6600' (2010m), 15km NW Baeza, 2-6 III [19]76, S. Peck, cloud forest, dung trap 12/ ex collection H. and A. Howden/ HOLO TYPE, Ontherus politus, F. Genier, 1992. Dissected. 34 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Allotype 9 (CMNC): ECUADOR: Napo Prov., 2000m, 7 km S. Baeza, 20-28.II.1 979, H. and A. Howden/ dung, cup trap/ 23.II.[ 19]79/ ALLOTYPE, Ontherus politus, F. Genier, 1992.
Paratypes: ECUADOR: Locality unspecified, Buckley, 1880 (1 et MNHN). Carchi: Sebun doi* (=Sibundoy, Putumayo, Colombia?), 11-15.IX.1977, 2600m, L. Pefia (1 et HAHC); Env[iron]. de Tulcan, Dr G. Rivet, 1902 (2 9 9 MNHN). Napo: 24 km. N. W. Baeza, 2400m, 22. VII.1 994, F. Genier, pastureland, ex. cow dung ( 1 et, 3 9 9 FGIC); same data as holotype (1 et, 1 9 FGIC) (9 et et, 3 9 9 HAHC) (1 et QCAZ); same data as Allotype (1 et FGIC) (1 et HAHC).
8. Ontherus (Caelontherus) aequatorius Bates, 1891 comb.nov. (Figs. 66, 112-1 14, 266; Map 2)
Ontherus aequatorius Bates, 1891: 25 (description) Ontherus aequatorius Bates: Gillet, 191 lh: 57 (catalogue) Ontherus aequatorius Bates: Luederwaldt, 1931: 364 (comment) Ontherus aequatoris (sic) Bates: Luederwaldt, 1931: 400 (key) Ontherus aequatorius Bates: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body reddish brown to black. Pubescence fulvous. Length. 12-17 mm. Head. Anterior edge arcuate, slightly upturned and slightly bisinuate medially, clypeogenal junction shallowly notched. Clypeus strongly wrinkled on anterior third in males or completely, except anterior declivity of tubercle in females, remainder becoming shallowly punctate in males or anterior declivity of tubercle asperulate in females. Clypeofrontal suture, arcuate, finely carinate, with moderate conical process medially posterior to suture, reduced to small tubercle in smaller individuals. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior third in males or three-fourths in females and punctate on remainder of surface. Vertex finely punctate, punctures larger in females and smaller individuals. Pronotum. Transverse, W/L = 1.6; anterior edge slightly wider, slightly more behind lateral edges of eyes, narrowly membranous and almost straight behind head insertion; disc finely punctate, punctation becoming larger on anterior and lateral declivities. Anterior fourth declivous behind head insertion, posterior edge slightly sinuate and swollen medially, almost convex in smaller individuals. Elytra. Subquadrate, L/W = 1.0. Striae fine, moderately impressed on disc, slightly more impressed on apical declivity, broadly oval to rounded, umbilicate, slightly deeper than stria, deeply impressed on apical declivity, approximately 3-4 times as wide as stria, separated by 2-4 diameters on disc. Intervals moderately convex, strongly alutaceous throughout, punctures moderate and evenly distributed, denser on sutural interval. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent, external third smooth. Prosternumbehind procoxa moderately covered with long pubescence. Mesosternum impressed posteriorly, punctation large and dense, pubescence fine and short. Mesepisternum moderately punctate, punctures broadly oval to rounded and of 2 different sizes, strongly carinate. Metasternum with disc finely punctate and with shallow longitudinal sulcus on posterior half and large circular concavity on anterior half in males or shallow longitudinal sulcus in females; lateral lobes moderately punctate on anterior half and along anterior edge of metacoxa, punctures rounded and well-impressed, pubescence moderately long and obliquely oriented, surface strongly alutaceous, more so anteriorly; median lobe (Fig. 66) moderately wide, L/mW = 2.2, lateral margins convergent on posterior half and parallel to slightly divergent on anterior half, surface punctate and strongly alutaceous throughout, junction with mesosternum slightly arcuate and strongly carinate. Legs. Anterior tibia quadridentate in both sexes, apical tooth slender and inwardly bent on apical third or unmodified in females, ventromedial carina lacking intervening setae on basal half, apical spur spiniform, straight and shorter than apical tooth in males or slender, inwardly bent on apical third and longer than apical tooth in A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 35 females. Posterior legs unmodified. Abdomen. Stemites 2-5 with row of small setiferous punctures along anterior edge, few setiferouspunctures laterally, surface alutaceous through out, stemite 6 finelypunctate throughout, suture between stemites 5- 6 with row of punctures laterally. Pygidium, W/L = 1.7 in males or 2.2 in females, punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 112-114). Tubus approximately 2.0 times as long as parameres. Parameres in lateral view with apices rounded, ventral portion slightly concave, dorsal portion largely membranous on basal two-thirds. Sigmoid sclerite of internal sac as in Figure 266.
MATERIAL STUDIED. 69 CJ' a, 57 Q Q (BMNH, CASC, CNCI, FGIC, HAHC, ICCM, ISNB, MNHN, NMPC, SMFD, SMTD, USNM,WDEC, ZMHB, ZMUC).
TYPE MATERIAL Holotype CJ' (MNHN): (recto) Ecuador., feet., Ed. Whymper., (verso) 5, G,/ Ontherus aequatorius Bates/ Museum Paris, ex coll., R. Oberthi.ir, 1952/ HOLOTYPE, Ontherus aequatorius, Bates, 1891, vidit F. Genier, 1991. Holotype studied.
NON-TYPE MATERIAL Distribution: ECUADOR: Pucay*; Sebundoy* (=Sibundoy, Putumayo, Colombia?). Azuay: 8km N E Giron (2600m); Tarqui. Cafiar: 5km E Zhud (3000m). Chimborazo: Riobamba. Napo: 24 km NW Baeza (2400m). Pichincha: Gualea.
Additional data: Some specimens were collected under cow dung in pasture at an elevation of 2400 m. Specimens collected in March, June, July, November.
KIRSCHII GROUP
Diagnosis. Both sexes distinguished by slightly arcuate anterior edge of median lobe of mesostemum, combined with tuberculate head, tubercles being located behind clypeofrontal suture. In males, slender and forwardly directed apical tooth of anterior tibia, combined with modified posterior femur, and simple parameres, separates the following 2 species.
9. Ontherus (Caelontherus) kirschii Harold, 1867 comb.nov. (Figs. 14-20, 71, 115-117, 267; Map 4)
Ontherus kirschii Harold, 1867: 96 (description) Ontherus kirschi Harold: Gemminger and Harold, 1869: 1008 (unjustified emendation)(catalog) Ontherus kirschi Harold: Harold, l 869a: 59 (comment) Ontherus kirschi Harold: Harold, 1869c: 503 (comment) Ontherus kirsch ii Harold: Kirsch, 1871: 357 (comment) Ontherus kirschi Harold: Harold, 1880; 21, 22 (distribution and comment) Ontherus elongatus Waterhouse, 1891: 357 (description) new synonymy. Ontherus elongatus Waterhouse: Gillet, 191 lb: 58 (catalog) Ontherus kirschi Harold: Gillet, 191 lh: 58 (catalog) Ontherus kirschi Harold: Luederwaldt, 1931: 396, 399 (key) Ontherus elongatus Waterhouse: Luederwaldt, 1931: 364, 396, 399 (comment and key) Ontherus elongatus Waterhouse: Blackwelder, 1944: 206 (list) Ontherus kirschi Harold: Blackwelder, 1944: 206 (list) Ontherus elongatus Waterhouse: Roze, 1955: 44 (list)
REDESCRIP TION. Colour. Body reddish brown to black, sometimes with moderate to strong iridescent tinge on elytra. Pubescence fulvous to reddish brown. Length. 11-16 mm. Head. Anterior edge arcuate, slightly upturned and bisinuate medially, clypeogenaljunction straight. Clypeus strongly wrinkled throughout in both sexes, wrinkles feeble basally in smaller individuals. Clypeofrontal suture arcuate, finely carinate, with small conical process medially posterior to suture, reduced to small tubercle in smaller individuals. Anterior genal edge subequal to lateral, genal surface moderately to strongly wrinkled on anterior half and 36 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 punctate on remainder of surface, punctures fine along eye margins. Vertex finely punctate except smooth area medially on posterior half. Pronotum. Transverse, WIL = 1.6; anterior edge narrowly membranous and almost straight behind head insertion; posterior margin slightly sinuate medially; disc finely punctate, punctures becoming larger and denser on anterior and lateral declivities. Anterior fifth with small transverse swelling as wide as third distance between eyes, convex in smaller individuals. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide, alutaceous and shallowly impressed throughout, punctures oval, slightly deeper than stria, less than 2 times as wide as stria, separated by 2-4 diameters on disc. Intervals moderately convex, punctures extremely fine, evenly distributed on disc and lateral declivities, larger and more numerous on apical declivity. Thoracic sterna. Proepistemum with posterolateral portion densely pubescent, external fourth smooth. Prosternum behind procoxa densely covered with pubescence, posterior edge with fewrows of setae approximately 2 times as long as pubescence on disc. Mesostemum impressed posteriorly, punctures large and dense, except small area along anterior edge of mesocoxa, pubescence fine and short. Mesepistemum coarsely punctate, punctures broadly oval to round, impressed, strongly carinate. Metastemum with disc finely punctate and alutaceous, with shallow longitudinal sulcus on anterior third and oval concavity on posterior two-thirds in males or shallow longitudinal sulcus slightly wider on posterior half in females; lateral lobes densely punctate and strongly alutaceous on anterior half, punctures oval, anterior punctures each with anterior edge sharply carinate, posterior half with punctures scarcer, broadly oval to rounded, surface finely alutaceous, pubescence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.3, lateral margins convergent on posterior three-fourths and parallel to slightly divergent on anterior fourth, surface finely and densely punctate, finely alutaceous throughout, junction with mesostemum slightly arcuate and carinate. Legs. Anterior femora flattened on basal half in males or unmodified in females. Anterior tibia quadridentate in both sexes, apical tooth slender and slightly bent inwardly on apical third in males or unmodified in females, ventromedial carina lacking intervening setae on basal third to half, apical spur spiniform, fine on apical half and shorter than apical tooth in males or slender, slightly bent inwardly on apical fourth and subequal to apical tooth in females. Posterior femur (Fig. 71) slender, posterodorsal edge triangularly dentate apically, posteroventral edge largely interrupted medially, never lobed basally. Posterior tibia flattened and widened in larger males or unmodified in females and smaller males. Abdomen. Sternites 2-5 with row of small setiferous punctures along anterior edge, few setiferous punctures laterally, surface finely alutaceous, stemite 6 finely and densely punctate throughout, suture between stemites 5-6 with row of small foveolate punctures laterally. Pygidium, W/L = 2.0 in males or 2.4 in females, punctures moderate and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 115-117). Tubus approximately 2.5 times as long as parameres. Parameres in lateral view with apices subtruncate, ventral and dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 267.
MATERIAL ST UDIED. 231 u u, 203 9 9 (AMNH, BDGC, BMNH, CASC, CNCI, CMNC, FGIC, HAHC, ISNB, IZAV, MCZC, MNHN, NMPC, PAIC, SMFD, SMTD, USNM, UZMH, ZMAS, ZMHB, ZMUC, ZSMC).
TYPE MATERIAL Ontherus kirschii Harold: Lectotype 9 (MNHN): Bogota/ Kirschi, t. Harold/ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthiir, 1952/ LECTOTYPE, Ontherus kirschii, Harold, 1867, Des. F. Genier, 1991. Type material studied and lectotype here designated. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 37
Ontherus elongatus Waterhouse: Holotype ct (BMNH): Venezuela (recto), 47.26 (verso)/ Type/ Ontherus elongatus, ct, (�) Waterh./ Holotype, Ontherus elongatus, Waterhouse, 1891, vidit F. Genier, 1991I Ontherus kirschii, Harold, Det. F. Genier, 1991. Holotype studied.
NON-TYPE MATERIAL Distribution: COLOMBIA: Josomoco*. Boyaca: Muzo. Cundinamarca: Fomeque; Fusagasuga; Monteredondo; Susumuco (23km W Villavicencio) (1 OOOm); Ub:ique. Distrito Especial: Bogota. Meta; Villavicencio. Norte de Santander: 3km NChinacota (lOOOm); Mesa Rica (2500m)*; Ocafia. Putumayo: Upper Putumayo R. Valle: Cali. VENEZUELA: Aragua: Portapan (Turmero) (1500m); Rancho Grande (l lOOm) (l 150m) (1460m); Tiara (50km S W Caracas) (2000m). Barinas: La Chimenea (5km S La Soledad) (1500m)*. Distrito Federal: Caracas; Env[iron]. de Caracas. Tachira: Pregonero (Las Trampitas) (1240m); San Cristobal (1200m); 20km NE San Cristobal (1200m). Trujillo: Bocon6.
Additional data: Some specimens were collected using carrion traps set in an unspecified habitat, carrion traps set in submontane forest, human feces trap, and in mule dung. Specimens collected in February, March, May to December.
REMARKS. The holotype (male) of 0. elongatus Waterhouse has been studied and is identical in all respects to 0. kirschii Harold.
10. Ontherus (Caelontherus) sanctaemartae sp.nov. (Figs. 72, 118-120, 268; Map 4)
ETYMOLOGY. Sanctaemartae, latinized form of Santa Marta, name of the Sierra in northern Colombia where this species has been collected.
DESCRIPTION. Colour. Body reddish brown to dark brown. Pubescence fulvous. Length. 11-16 mm. Head. Anterior edge arcuate, slightly upturned, sinuate in males or slightly bisinuate in females medially, clypeogenal junction finely notched. Clypeus strongly wrin kled on anterior half, remainder punctate in males or strongly wrinkled throughout in females. Clypeofrontal suture arcuate and finely carinate, with small conical process medially posterior to suture, reduced to small tubercle in smaller individuals. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior half and punctate on remainder of surface, punctures fine along anterior edge of eyes. Vertex punctate except smooth area
medially on posterior half. Pronotum. Transverse, W IL = 1.6; anterior edge narrowly membranous and almost straight behind head insertion; posterior margin slightly lobate medially; disc finely punctate, punctures becoming larger and denser on anterior and lateral declivities. Anterior fourth declivous on median third, with transverse swelling approxi mately as wide as distance between eyes, width of transverse swelling reduced to approxi mately half distance between eyes in smaller individuals, almost convex in small individuals.
Elytra. Subquadrate, L/W = 1.0. Striae fine, alutaceous and moderately impressed through out, punctures rounded, deeper than stria, approximately 2 times as wide as stria, separated by 2-4 diameters on disc. Intervals slightly convex, punctures fine, evenly distributed, slightly more numerous on sutural interval. Thoracic sterna. Proepisternum with postero lateral portion moderately pubescent, external fourth smooth. Prosternum behind procoxa densely covered with fine and erect setae on anterior two-thirds and with long and erect setae on posterior third. Mesosternum impressed posteriorly, punctures large and dense, except small area along anterior edge of mesocoxa, pubescence fine and short. Mesepisternum finely punctate, punctures broadly oval to rounded and impressed, strongly carinate and alutaceous. Metasternum with disc finely punctate, with shallow longitudinal concavity on posterior three -fourths in males or shallow and narrow sulcus in females; lateral lobes densely punctate and strongly alutaceous on anterior half, punctures oval, anteriormost 38 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
\l � 0
• 0. kirschii
* 0. sanctaemartae
MAP 4. Known distribution of Ontherus kirshii and 0. sanctaemartae.
punctures each with anterior edge finely carinate, posterior half with punctures scarcer, broadly oval to round, surface finely alutaceous, pubescence moderately long and obliquely
oriented; median lobe moderately wide, L/mW = 2.1, lateral margins convergent on posterior four-fifths and parallel to slightly divergent on anterior fifth, surface finely and densely punctate, finely alutaceous throughout, junction with mesosternum slightly arcuate and carinate. Legs. Anterior femur flattened on basal half in larger males or unmodified in females and small males. Anterior tibia quadridentate in both sexes, apical tooth slender and slightly bent inwardly on apical third in males or unmodifiedin females, ventromedial carina lacking intervening setae on basal third, apical spur spiniform, fine on apical half and shorter than apical tooth in males or slender, slightly bent inwardly on apical fourth and slightly longer than apical tooth in females. Posterior femur (Fig. 72) slender, posterodorsal edge triangularly dentate apically, posteroventral edge shortly interrupted apically and basal half lobed in larger males, posteroventral edge fine but complete apically in smaller males. Posterior tibia unmodifiedin both sexes. Abdomen. Stemites 2-5 with row of finesetiferous punctures along anterior edge, few setiferous punctures laterally, surface finely alutaceous, stemite 6 finely and densely punctate throughout, suture between sternites 5-6 with row of
small irregular punctures laterally. Pygidium, W/L = 2.0 in males or 2.2 in females, punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 118-120). Tubus approximately 2 times as long as parameres. Parameres in lateral view with apices subtruncate, ventral and dorsal portion slightly concave, dorsal portion sharply emarginate before apex. Sigmoid sclerite of internal sac as in Figure 268.
MATERIAL STUDIED. 8 a a, 8 9 9 (CMNC, FGIC, HAHC, ICCM, MCZC, MNHN).
Holotype O' (MCZC): N. W. Sierra N[evada]., de Sta. Marta, Colombia VII.20 [19]28, 5-9000 ft (1500-2700m), Darlington/ Ontherus kirschi, Har., P. Pereira det. 958/ HOLOTYPE, Ontherus sanctaemartae, F. Genier, 1992. Dissected. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 39
Allotype 9 (CMNC): COLOM[BIA] Magd., 3000' (9 lOm), Campana, 25 km. S., Sta. Marta, IV-29-1973, Howden and Campbell./ ex. Collection, H. and A. Howden/ ALLOTYPE, Ontherus sanctaemartae, F. Genier, 1992.
Paratypes: COLOMBIA: Magdalena: same data as holotype (1ct, 19 FGIC) (2 ctct, 29 9 MCZC); Bonda, June (19 ICCM); Campana, 24 km S. Sta. Marta, 3000' (910m), 14.V.1973, Campbell and Howden (1ct HAHC); Campana, 26 km S. Sta. Marta, 3500' (1060m) (2ctct HAHC); El Libano, April, 6000' (1830m) (1ct, 19 ICCM); Nevada Sta Marta, F. Simons (1 9 MNHN); Valparaiso, April, 2500' (760m) (19 ICCM).
ALEXIS GROUP
DIAGNOSIS. Both sexes in this species group possess the following combination of characters: anterior edge of median lobe of metasternum straight or slightly arcuate and pronotum tuberculate and/or carinate. Males distinguished by presence of laterally com pressed cephalic horn [except for 0. ale.xis (low conical process) and 0. diabolicus (almost circular)], tridentate anterior tibia and simple parameres. Females distinguished by 2 closely set tubercles on clypeofrontal suture.
11. Ontherus (Caelontherus) alexis (Blanchard), 1845 comb.nov. (Figs. 25, 26, 121-123, 269; Map 5)
Copris alexis Blanchard, 1845 (1843): 180 (description) Ontherus didymus Erichson, 1847: 108 (description) new synonymy. Ontherus glaucinus Erichson, 1847: 108 (description) new synonymy. Ontherus didymus Erichson: Gemminger and Harold, 1869: 1008 (catalog) Ontherus glaucinus Erichson: Gemminger and Harold, 1869: 1008 (catalog) Pinotus alexis (Blanchard): Gemminger and Harold, 1869: 1009 (catalog) Copris alexis Blanchard: Harold, 1869a: 59 (comment) Ontherus glaucinus Erichson: Harold, 1880: 21, 22 (redescription, distribution and comment) Ontherus didymus Erichson: Bates, 1887: 50 (redescription, distribution and comment) Ontherus nevinsoni Waterhouse, 1891: 357 (description) new synonymy. Ontherus alexis (Blanchard): Gillet, 191lh:57 (catalog) Ontherus didymus Erichson: Gillet, 191 lh: 57 (catalog) Ontherus glaucinus Erichson: Gillet, 191 l h: 58 (catalog) Ontherus nevinsoni Waterhouse: Gillet, 191lh: 58 (catalog) Ontherus didymus Erichson: Lucas, 1920: 459 (catalog) Ontherus didymus Erichson: Luederwaldt, 1931: 364, 395, 398 (comment and key) Ontherus glaucinus Erichson: Luederwaldt, 1931: 364, 394, 395, 397 (comment and key) Ontherus nevinsoni Waterhouse: Luederwaldt, 1931: 364, 395, 397, 400, 401 (comment and key) Ontherus alexis (Blanchard): Luederwaldt, 1931: 395, 400 (comment and key) Ontherus glaucinus Erichson: Balthasar, 1938: 221 (comment) Ontherus didymus Erichson: Balthasar, 1938: 221 (comment) Ontherus alexis (Blanchard): Blackwelder, 1944: 206 (list) Ontherus didymus Erichson: Blackwelder, 1944: 206 (list) Ontherus glaucinus Erichson: Blackwelder, 1944: 206 (list) Ontherus nevinsoni Waterhouse: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body uniformly light brown to black, occasionally with moderate iridescent tinge on elytra. Pubescence reddish brown. Length. 10-14 mm. Head (Figs. 25, 26). Anterior edge broadly arcuate and slightly upturned, bisinuate medially, clypeogenal junction finelynotched. Clypeus transversely wrinkled on apical fourth in males and apical half in females, remainder shallowly and evenly punctated. Clypeofrontal suture broadly arcuate, with large conical process in males or transverse conical process bituber culate apically in females or suture with 2 adjacent low tubercles in small individuals. Anterior genal edge subequal to lateral, genal surface finely punctate, transversely wrinkled on anterior fourth in males and small individuals, on anterior half in larger females. Vertex almost smooth on posterior half, only slightly depressed internally to eyes in larger males. 40 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Pronotum (Figs. 25, 26). Transverse, W/L = 1.7; anterior edge expanded on middle half, only slightly emarginate internally to eyes in larger individuals; punctures becoming larger on anterior angles, disc almost smooth in larger individuals. Anterior fourth with 2 adjacent and small tubercles medially, strongly declivous in larger males. Elytra. Subquadrate, L/W
= 1.0. Striae fine, alutaceous, shallowly impressed, only slightly more impressed on apical declivity, punctures approximately twice the width of stria, separated by approximately 4 diameters. Elytral intervals almost flat, finely and evenly punctate, punctures denser on sutural interval. Thoracic sterna. Proepisternum with posterolateral portion densely pubes cent. Prosternum behind procoxa densely punctate and pubescent. Mesosternum impressed posteriorly, punctures deep and dense, pubescence fine and short, row of coarse and long setae along anterior edge of mesocoxa. Mesepisternum densely punctate and strongly carinate. Metasternum with disc finely and densely punctate; lateral lobes moderately punctate, punctures oval, anteriormost punctures each with anterior edge sharply carinate, pubescence moderately long and obliquely oriented, surface alutaceous, more so on anterior
half; median lobe narrow, L/mW = 3.2, lateral margins divergent on anterior fifth, surface moderately punctate, punctures becoming larger and scarce anteriorly, finely alutaceous on anterior fourth, junction with mesosternum straight, declivous, sometimes finely carinate. Legs. Anterior legs with tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on posterior fourth, apical spur obliquely truncate and dentate internally at apex, longer than apical tooth. Metathoracic legs unmodified. Abdo men. Sternites 2-5 with few round and deeply impressed setiferous punctures laterally, surface finely alutaceous, sternite 6 finely punctate and alutaceous throughout, suture between sternites 5-6 weakly grooved and with row of umbilicate punctures laterally.
Pygidium, WIL = 2.1 in males or 2.3 in females, punctures moderate, shallowly impressed and evenly distributed, apical margin clearly delimited in both sexes. Male genitalia (Figs. 121-123). Tubus approximately 3 times as long as parameres, apicoventral portion flattened. Parameres in lateral view with apices subquadrate and denticulate ventrally at apex. Sigmoid sclerite of internal sac as in Figure 269.
MATERIAL ST UDIED. 148 a a, 119 9 9 (BMNH, CASC, CNCI, DEIC, HAHC, LACM, ICCM, ISNB, MACN, MNHN, NMPC, PA IC, SEMC, SMFD, SMTD, USNM, UZMH, WDEC, ZMHB, ZMUC, ZSMC).
TYPE MATERIAL Copris alexis Blanchard: Lectotype O' (MNHN): 1281/ green disk, 6280, 34/ Museum Paris, Bolivie (Yungas), D'Orbigny 1834/ alexis Bl./ type/ LECTOTYPE, Copris, alexis, Blanchard 1845, Des. F. Genier, 1991. Type material studied and lectotype here designated.
Ontherus didymus Erichson: Holotype 9 (ZMHB): 26625/ Peru Mont. Phil[lippi?]/ didymus Er./ typus/ HOLOTYPE, Ontherus didymus, Erichson, 1847, vidit F. Genier, 1991. Holotype studied.
Ontherus glaucinus Erichson: Lectotype a (ZMHB): 26626/ Ontherus glaucinus Er., Peru M[ont?]. Phil[ippi?]./ Typus/ glaucinus Er. Peru, N. Granad./ LECTOTYPE, Ontherus glaucinus, Erichson, 1847, Des. F. Genier, 1991. Type material studied, lectotype and paralectotypes here desig nated. Paralectotype: Peru mont, philippi, Nr. 26626/ Typus/ PARALECTOTYPE, Ontherus glau cinus, Erichson, 1847, Des. F. Genier, 1991. 1 a (ZMHB). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 41
• 0. alexis • 0. incisus
MAP 5. Known distribution of Ontherus alexis and 0. incisus.
Ontherus nevinsoni Waterhouse: Holotype a (BMNH): Type/ 339./ 6745/ Ontherus nevinsoni, (�) Waterh./ Copris, Coptodera Reiche, Yungos Bolivia/ Holotype, Ontherus nevinsoni, Waterhouse, 1891, vidit F. Genier, 1991. Holotype studied. Allotype 9 (BMNH): Type/ 340/ 67.45/ coptodera, Reiche, Yungos, Bolivia/ Ontherus nevinsoni, 9 (�) Waterh./ Allotype, Ontherus nevinsoni, Waterhouse, 1891, vidit F. Genier, 1991. Allotype studied.
NON-TYPE MATERIAL Distribution: BOLIVIA: Cordico*; Cristal Mayu*; Fanampayo*; Yungas. Cochabamba: Rio Chapare (Alto palmar) (llOOm); Yungas del Palmar (2000m). La Paz: Calisaya (Rio Boopi); Chaco; Coroico; Songo ( =Zongo); Suapi. Santa Cruz: Buena Vista. COLOMBIA (doubtfull): Narifio: Pasto. PERU: Ayachuco: Opara*. Cuzco: Callanga; Marcapata; Pau cartambo; Rio Oxabamba (=Rio Ocabamba?); Rio Tono. Huanuco: Leonpaupa (800m)*. Junin: La Merced; Sani Beni (840m)*. Loreto: Jurimaguas (=Yurimaguas).
Additional data: Specimens collected in January to March, May, June, August to December.
REMARKS. Ontherus didymus Erichson and 0. glaucinus Erichson were described at the same time based on 3 specimens, 2 males and 1 female. Erichson misinterpreted the secondary sexual characters for specific characters, and named the male form 0. glaucinus and the femaleform 0. didymus. I also had the opportunity to study Waterhouse type material of 0. nevinsoni. After careful comparison with the type of 0. alexis Blanchard it is clear that they all represent a single species.
12. Ontherus (Caelontherus) tenuistriatus sp.nov. (Figs. 27, 28, 124-126, 270; Map 6)
ETYMOLOGY. Tenuistriatus, (thin+ stria) Latin adjective, refers to the elytral striae of this species. 42 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. teniustriatus • 0. diabolicus
MAP 6. Known distribution of Ontherus tenuistriatus and 0. diabolicus.
DESCRIPTION. Colour. Body uniformly black. Pubescence fulvous. Length. 13-14 mm. Head (Figs. 27, 28). Anterior edge arcuate and almost flat, slightly sinuate medially in both sexes, clypeogenal junction straight. Clypeus transversely wrinkled on apical two-fifths, remainder finely punctate. Clypeofrontal suture arcuate, with laterally compressed and posteriorly curved horn. Anterior genal edge longer than lateral, genal surface finely punctate, wrinkled on anterior fifthin males or anterior half in females.Vertex finely punctate
throughout, convex. Pronotum (Figs. 27, 28). Transverse, W/L = 1.6; anterior edge deeply emarginate and widely membranous behind eyes in males or slightly emarginate internally to eyes and membranous behind head insertion in females; punctures fine throughout. Anterior portion of disc with 2 transverse, slightly to moderately sinuous, carina, interval between carinae depressed and width subequal to length of carina in males, or anterior fourth with transverse row of 4 tubercles, lateral tubercles distinctly lower and located slightly
behind median tubercles in females.Elytra. Subquadrate, L/W = 1.1. Striae extremely fine, moderately impressed on disc and lateral declivities, deeply impressed on apical declivity, punctures rounded, umbilicate and deeper than stria, especially on apical declivity, approxi mately 3 times as wide as stria, separated by2-4 diameters on disc. Intervals slightly convex, punctures fine and evenly distributed. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent, external fourth smooth. Prosternum behind procoxa moder ately pubescent, with few setae twice as long as those on disc along posterior margin. Mesosternum impressed posteriorly, punctures dense on posterior third, small smooth area along anterior edge of mesocoxa, pubescence fine and short except few long setae along mesocoxa. Mesepisternumwith punctures rounded and of2 differentsizes, strongly carinate and finely alutaceous. Metasternum with disc smooth and with shallow and ill-defined longitudinal depression in males or convex in females; lateral lobes moderately punctate and alutaceous on anterior half, punctures oval, posterior half with punctures mostly restricted to posterior edge, surface finely alutaceous, pubescence moderately long and obliquely
oriented; median lobe narrow, UmW = 2.6, lateral margin convergent on their posterior three-fourths and parallel to slightly divergent on anterior fourth, surface moderately and A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 43 densely punctate, finely alutaceous throughout, few long setae along laterally on posterior third, junction with mesosternum slightly arcuate, declivous medially. Legs. Anterior tibia tridentate in males or quadridentate in females, ventromedial carina lacking intervening setae on basal half in males or third in females, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with row of deep punctures along anterior edge, few setiferous punctures laterally, surface finely alutaceous, sternite 6 finely and densely punctate throughout, suture between sternites5- 6 with row of small irregular punctures laterally. Pygidium, W /L = 1.6 in males or 1. 9 in females, punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 124-126). Tubus approximately 3 times as long as parameres, apicoventral portion deeply sulcate. Parameres in lateral view, with apices quadrate, ventral and dorsal portion slightly concave, ventral portion denticulate apically. Sigmoid sclerite of internal sac as in Figure 270.
MATERIAL STUDIED. 2 a a, 1 9 (MNHN, ZMHB).
Holotype O' (MNHN): Peru/ Ex. Musaeo, E. Harold/ Museum Paris, ex. coll., R. Oberthur/ HOLOTYPE, Ontherus tenuistriatus, F. Genier, 1992. Dissected.
Allotype 9 (MNHN): Perou, Chanchamayo, Thamin/ Museum Paris, ex Coll., R. Oberthur/ Allotype, Ontherus tenuistriatus, F. Genier, 1992.
Paratype: PERU: HUANUCO: Huanuco (1 a ZMHB).
13. Ontherus (Caelontherus) obliquus sp.nov. (Figs. 29, 30, 127-129, 271; Map 7)
ETYMOLOGY. Obliquus, (oblique) Latin adjective, refers to orientation of the pronotal carinae in males of this species.
DESCRIPTION. Colour. Body uniformly dark reddish brown to black, occasionally with faint iridescent tinge on elytra. Pubescence fulvous. Length. 10-15 mm. Head (Figs. 29, 30). Anterior edge arcuate and slightly upturned medially, slightly sinuate in males or bisinuate in females medially, clypeogenal junction straight. Clypeus transversely wrinkled on apical two-fifths in males or apical half in females, remainder finely punctate in both sexes. Clypeofrontal suture arcuate and lacking carina in males or finely carinate laterally in females, with laterally compressed and posteriorly curved horn in males, reduced to low conical process with apex bituberculate in smaller males, or 2 well-separated low tubercles in females. Anterior genal edge subequal to lateral, genal surfacefinely punctate throughout in males or wrinkled on anterior fourth and remainder finely punctate in females. Vertex finely punctate on anterior half, remainder smooth and convex, feebly depressed along internal margin of the eyes in males. Pronotum (Figs. 29, 30). Transverse, W/L = 1.6; anterior edge deeply emarginate and widely membranous behind the eyes in males or slightly emarginate internally to eyes and membranous behind head insertion in females; punctures finethroughout. Anterior portion of disc with 2 oblique and almost straight carinae, interval between carinae feebly to moderately depressed and subequal in width to length of carina in males or anterior fifth with 2 transverse and slightly arcuate bulges with inner edge tuberculate, anterior portion distinctly concave in females, smallestindividuals with prono tum almost convex and with 2 median and small swelling on anterior fifth. Elytra.
Subquadrate, L/W = 1.0. Striae fine, slightly alutaceous, moderately impressed on disc and lateral declivities, deeply impressed on apical declivity, punctures broadly oval to rounded and slightly deeper than stria, approximately 2 times as wide as stria, separated by 3-5 diameters on disc. Interval slightly convex, alutaceous throughout, punctures fineand evenly distributed. Thoracic sterna. Proepisternum with posterolateral portion moderately 44 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. obliquus
• 0. compressicornis
MAP 7. Known distribution of 0ntherus obliquus and 0. compressicornis.
pubescent, external fifth smooth. Prosternum behind procoxa moderately pubescent, with few setae twice as long as those on disc along posterior margin. Mesosternum impressed posteriorly, punctures dense on posterior third, small smooth area along anterior edge of mesocoxa, pubescence fineand short except few long setae along mesocoxa. Mesepisternum with punctures rounded and of 2 different sizes, strongly carinateand alutaceous. Metaster num with disc finely punctate and strongly alutaceous throughout, with shallow and narrow longitudinal sulcus on posterior two-thirds in males or posterior third in females; lateral lobes moderately punctate throughout, except row of smaller and denser punctures along anterior edge of metacoxa, punctures oval on anterior half and broadly oval to rounded on posterior half, surface strongly alutaceous on anterior half, pubescence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2. 7, lateral margins convergent on posterior four-fifthsand divergent on anterior fifth, surfacemoderately and densely punctate, strongly alutaceous throughout, junction with mesosternum straight, declivous medially, finely carinate laterally. Legs. Anterior tibia tridentate in males or quadridentate in females, ventromedial carina lacking intervening setae on basal half, apical spur inwardly bent on apical third, longer than apical tooth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few setiferous punctures laterally, surface strongly alutaceous, sternite6 finely punctate and alutaceous throughout, suture between sternites 5-6 with few superficial and small punctures laterally. Pygidium, W/L = 1.8 in males or 1.9 in females, punctures fine and dense especially along basal margin, alutaceous on basal two-thirds, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 127-129). Tubus approximately 2.5 times as long as parameres, apicoventral portion enlarged. Parameres in lateral view, with apices quadrate, ventral and dorsal portions slightly concave, ventral portion denticulate apically. Sigmoid sclerite of internal sac as in Figure 271.
MATERIAL STUDIED. 1040' a, 719 9 (CNCI, FGIC, ISNB, MCZC, MNHN, MZLU, PAIC, SMTD, ZMUC, ZSMC). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 45
Holotype a (ZSMC): Bolivia - Yungas -, Incachaca, 2100m - 10.II.1959, Leg. Zischka/Zool. Staatsslg.,Mtinchen/ HOLOTYPE, Ontherus obliquus,F. Genier, 1992. Dissected.
Allotype Q(CNCI): Bolivia, Yungas Del Palmar,2000m. - Zischka, II.1950/ ALLOTYPE, Ontherus obliquus, F. Genier, 1992.
Paratypes: No locality (1a MCZC) (1 QZSMC). COLOMBIA (doubtful): No locality (2Q ISNB). BOLIVIA: No locality (2aa ZMUC); Chiquilas* (8a a, 1 Q SMTD); Yungas (5 aa, 10 QQ ISNB) (1a, 1 QSMTD). Cochabamba: No locality, P. Germain, 1889 (8 aa, 3a a MNHN) (2a a FGIC); Chapare, 2000m,X.1950 (2 aa, 1 QPAIC); Yungas del Palmar,2000m, leg. Zischka (21a a, 17Q Q ZSMC) (2 a a, 2 QQ FGIC), same as before except: II.1950 (5 a a, 8 QQ ZSMC) ( 1a CNCI),2.II.1950 ( 4 a a, 1 Q ZSMC), 15.111.1953 (4a a ZSMC), II.1954 (4aa, 8 QQ ZSMC) (1a, 1 Q CNCI), 8.II.1954 (2aa ZSMC), III.1956 (9aa,5Q Q ZSMC) (la, 1QFGIC),15.III.1958 (2 a a, 2 QQ ZSMC); Yungas de Totora-Schuencas, 2lOOm, leg. Zischka (1a ZSMC); Yungas Incachaca, 2100m,10.II.1959, leg. Zischka (5a a, 2 QQ ZSMC). La Paz: Chaco (6aa, 1 QMNHN) (1a ISNB), same as before except: Garlepp (1a SMTD), Yungas, 3000m,G. Garlepp (1a MNHN); San Antonio* (1a MNHN). ECUADOR: No locality (1a ISNB). PERU: Cuzco: Agua Caliente*, 28.XII.1983,Leg. L. Huggert (1 QMZLU); Vilcanota (Cordillera de?) (1Q MNHN). Junin: Chanchamayo (1a ISNB).
14. Ontherus (Caelontherus) incisus (Kirsch), 1871 comb.nov. (Figs. 31, 32,130-132, 272; Map 5)
Pinotus incisus Kirsch, 1871 (1870): 357 (description) Ontherus incisus (Kirsch): Harold, 1880: 23 (distribution and comment) Ontherus thoracicus Waterhouse, 1891: 356 (description) new synonymy. Pinotus incisus Kirsch: Waterhouse, 1891: 356 (comment) Ontherus incisus (Kirsch): Gillet, 191 lb: 58 (catalog) Ontherus thoracic us Waterhouse: Gillet, 1911b: 58 (catalog) Ontherus incisus (Kirsch): Luederwaldt, 1931: 364, 398, 400 (comment and key) Ontherus thoracicus Waterhouse: Luederwaldt, 1931: 364, 395, 399 (comment and key) Ontherus incisus (Kirsch): Balthasar, 1938: 221 (comment) Ontherus incisus (Kirsch): Blackwelder, 1944: 206 (list) Ontherus thoracicus Waterhouse: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body uniformly reddish brown to dark brown. Pubescence reddish. Length. 11-15 mm. Head (Figs. 31, 32). Anterior edge arcuate and slightly upturned, bisinuate, median emargination v-shaped and stronger in females,clypeogenal junction straight to feebly notched. Clypeus transversely wrinkled on apical third in males or apical half in females, remainder finely punctate in both sexes. Clypeofrontal suture broadly arcuate, feeble in males or finely carinate laterally in females,with laterally compressed and posteriorly curved horn in males,reduced to low conical process with apex bituberculate in smaller males or 2 well-separated low tubercles in females. Anterior genal edge slightly longer than lateral, genal surface finely punctate throughout in males or wrinkled on anterior half and remainder finely punctate in females. Vertex smooth and convex in both sexes, slightly to moderately depressed along internalmargin of eyes in males.
Pronotum (Figs. 31, 32). Transverse,W/L = 1.6; anterior edge deeply emarginate and widely membranous behind eyes in males or slightly emarginate internally to eyes and membranous behind head insertion in females; punctures fine throughout. Anterior portion of disc with 2 oblique and almost straight carinae in males or anterior fourth with row of 4 small tubercles in females and smallest males, lateral tubercles lacking in smaller females.
Elytra. Subquadrate, L/W = 1.0. Striae fine,wider on apical declivity,alutaceous, moder ately impressed, feebly impressed on humeral and apical cali, punctures oval,less than 2 times as wide as stria,separated by 3- 6 diameters on disc. Intervals flat,sometime alutaceous 46 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 along stria, feebly alutaceous on apical declivity, punctures minute, denser on apical declivity. Thoracic sterna. Proepisternum with posterolateral portion moderately pubes cent, external fourth smooth. Prosternum behind procoxa moderately pubescent, with few longer setae on disc. Mesosternum impressed posteriorly, punctures dense throughout, pubescence fine and short except 2-4 longer setae along mesocoxa. Mesepisternum irregu larly punctate, strongly carinate and alutaceous. Metasternum with disc finely punctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures mostly restricted to anterior half, fewpunctures along edge of metacoxa and on disc, punctures oval on anterior half and broadly oval to rounded on posterior half, surface alutaceous, slightly more on anterior half, pubescence moderately long and obliquely oriented; median lobe narrow,
L/mW = 2. 7, lateral margins convergent on posterior four-fifths and divergent on anterior fifth, surface finely and densely punctate, punctures larger anteriorly, alutaceous throughout, junction with mesosternum slightlyarcuate and declivous, lacking any traces of carina. Legs. Anterior tibia tridentate in males or quadridentate in females, ventromedial carina lacking intervening setae on basal third, apical spur inwardly bent on apical fourth in both sexes, posterior legs unmodified. Abdomen. Sternites 2-5 with fewsetiferous punctures laterally, surface slightly alutaceous and with few fine punctures, sternite 6 finely punctate and alutaceous throughout, suture between sternites 5-6 with row of superficial and small punctures laterally. Pygidium, W/L = 1.9 in males or 2.3 in females, punctures fine and dense, surface slightly alutaceous, apical margin clearly delimited in both sexes. Male genitalia (Figs. 130-132). Tubus approximately 2.0 times as long as parameres, apicoventral portion slightly concave on apical sixth. Parameres in lateral view with apex obliquely truncated, apicodorsal angle of truncation rounded, ventral portion straight, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 272.
MATERIAL STUDIED. 51 u u, 41 9 9 (AMNH, BMNH, CASC, HAHC, ISNB, MNHN, NMPC, PAIC, SMFD, SMTD, USNM, UZMH, ZMAS, ZMHB, ZMUC, ZSMC).
TYPE MAT ERIAL Pinotus incisus Kirsch: Holotype O" (by monotypy)(SMTD): Bogota, m. Kirsch/ Typus!/ Staatl. Museum for, Tierkunde Dresden/ HOLOTYPE, Pinotus, incisus, Kirsch, 1871, vidit F. Genier, 1991. Holotype studied. Ontherus thoracicus Waterhouse: Holotype u (BMNH): Type/ 67.45/ Colombia/ 337./ Ontherus thoracicus, u(Type) Waterh./ Copris, baterodera Reiche, Nova Granada/ Holotype, Ontherus thoracicus, Waterhouse, 1891, vidit F. Genier, 1991./ Ontherus incisus (Kirsch), 1871, Det. F. Genier,1991. Holotype studied. Allotype 9 (BMNH): 338/ 67.45/ Type/ Ontherus thoracicus, 9, (�) Waterh./ Allotype, Ontherus thoracicus, Waterhouse, 1891, vidit F. Genier, 1991/ Ontherus incisus (Kirsch), 1871, Det. F. Genier, 1991. Allotype studied.
NON-TYPE MATERIAL Distribution: Chiquilas*; Quebrada Negra*. COLOMBIA: Boyaca: Muzo. Caldas: Aguadas. Cundinamarca: Fusagasuga; Pacho. Distrito Especial: Bogota. Putumayo:
Sebondoy ( = Sibundoy) ECUADOR: Normandia*. Morona-Santiago: Macas. Napo: Baeza; 17km N.E. Baeza (1280m); 15km N.W. Baeza (2010m); 7km S Baeza (2000m). Pastaza: Canelos. Tungarahua: Santa Inez.
Additional data: Specimens were collected using cup dung traps, dung traps and mega dung traps set in forest and cloud forest. Specimens collected in February, March, December.
REMARKS. Waterhouse described 0. thoracicus, assuming that he had correctly identified a male specimen (fromMr. Nevinson's collection) with 4 tubercles on the thorax as 0. incisus A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 47
Kirsch. If this specimen was a male, and was from Colombia, it could only be 0. lunicollis sp.nov. The male specimen which he had at hand, had "two slightly prominent processes (separated from each other by a somewhat wide emargination), truncated at their apex, the angles of truncature slightly and obtusely dentiform". The type of 0. incisus (Kirsch), a small teneral male, was dissected and, both the external morphological characters, as well as the genitalic characters, are identical to Waterhouse's species. Therefore, 0. thoracicus Waterhouse is a junior synonym of 0. incisus (Kirsch).
15. Ontherus (Caelontherus) howdeni sp.nov. (Figs. 33, 34, 133-135, 273; Map 8)
ETYMOLOGY. Howdeni, a patronym in honour of Henry F. Howden who encouraged me to undertake this revision.
DESCRIPTION. Colour. Body uniformly reddish brown to dark brown. Pubescence reddish. Length. 11-14 mm. Head (Figs. 33, 34). Anterior edge arcuate and slightly upturned, bisinuate, median emargination v-shaped and stronger in females, clypeogenal junction moderately to deeply notched. Clypeus transversely wrinkled on apical third in males or apical half in females, remainder finely punctate in both sexes. Clypeofrontalsuture broadly arcuate, carina lacking in males or finely to moderately carinate laterally in females, with laterally compressed and posteriorly curved horn in males, reduced to low conical process with apex bituberculate in smaller males or 2 well-separated low tubercles in females. Anterior genal edge slightly longer than lateral, genal surface wrinkled on anterior fourth remainder finely punctate. Vertex finely punctate on anterior third, convex in both sexes, shallowly to moderately depressed along internalmargin of eyes in males. Pronotum
(Figs. 33, 34). Transverse, W/L = 1.5; anterior edge deeply emarginate and widely mem branous behind eyes in males or slightly emarginate internally to eyes and membranous behind head insertion in females;punctures finethroughout, slightly larger on anterior angles in females. Anterior portion of disc with 4 tubercles, outer ones lobiform and stronger than inner ones in larger males or anterior fourth with transverse row of 4 small tubercles in
females and smallest males. Elytra. Subquadrate, L/W = 1.0. Striae fine, wider on apical declivity, alutaceous, moderately impressed, less impressed on humeral and apical cali, punctures oval, approximately twice width of stria, separated by 3-6 diameters on disc. Intervals flat, finelyto moderately alutaceous throughout, punctures minute, denser on apical declivity. Thoracic sterna. Proepisternum with posterolateral portion moderately pubes cent, external fourth smooth. Prosternum behind procoxa moderately pubescent, with few longer setae on disc. Mesosternum impressed posteriorly, punctures dense throughout, except small area along edge of mesocoxa, pubescence fine and short except 1-4 longer setae along mesocoxa. Mesepisternum irregularly punctate, strongly carinate and alutaceous. Metasternum with disc finely punctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures mostly restricted to anterior half, few punctures adjacent to edge of metacoxa and on disc, punctures oval on anterior half and broadly oval to rounded on posterior half, surface alutaceous, slightly more on anterior half, pubescence moderately
long and obliquely oriented; median lobe narrow, L/mW = 2.5, lateral margins convergent on posterior four-fifths and divergent on anterior fifth, surface finely and densely punctate, punctures larger anteriorly, alutaceous throughout, junction with mesosternum slightly arcuate and declivous medially, lacking any traces of carina. Legs. Anterior tibia tridentate in males or quadridentate in females, ventromedial carina lacking intervening setae on basal third, apical spur inwardly bent on apical fourth in both sexes, posterior legs unmodified. Abdomen. Sternites2-5 with fewsetiferous punctureslaterally, surface slightly alutaceous and with few fine punctures, sternite 6 finely punctate and alutaceous throughout, suture between sternites 5-6 with a row of superficial and small irregular punctures laterally. 48 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
l
• 0. howdeni
• 0. pilatus
MAP 8. Known distribution of Ontherus howdeni and 0. pilatus.
Pygidium, WIL = 1.9 in males or 2.1 in females, punctures fine and dense, surface slightly alutaceous, apical margin clearly delimited in both sexes. Male genitalia (Figs. 133-135). Tubus approximately 2.0 times as long as parameres, apicoventral portion slightly tubercu late on apical sixth. Parameres in lateral view with apex rounded, ventral portion straight and denticulate at apex, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 273.
MATERIAL STUDIED. lOaa, 6 9 9 (CMNC, CNCI, HAHC, MNHN, SMTD, ZMAS, ZMHB).
Holotype a (CMNC): Chanchamayo, Peru, XI 15, 1968, J. Schunke L./ ex collection, H. and A. Howden/ HOLOTYPE, Ontherus howdeni, F. Genier, 1992. Dissected.
Allotype 9 (CMNC): Same data as holotype except: I 10, 1969.
Paratypes: CHILE: No locality (1a CNCI). ECUA DOR: No locality (1 a SMTD). PERU: No locality (2aa, 1 9 SMTD); "Inner Peru" (1 9 ZMAS). Junin: Chanchamayo (1 a MNHN), same as before except: Thamin. (1a MNHN), 10.1.1969, J Schunke L. (1 a, 1 9 FGIC) (19 HAHC), 12.1.1969 (1 a HAHC). Piura: Huancabamba (2aa, 1 9 ZMHB).
16. Ontherus (Caelontherus) diabolicus sp.nov. (Figs. 35, 36, 136-138, 274; Map 6)
ETYMOLOGY. Diabolicus, (diabolical) Latin adjective, refers to male head and pronotum aspect when viewed frontally.
DESCRIPTION. Colour. Body uniformlyreddish to black, with strong iridescent tinge on elytra in clean specimens. Pubescence fulvous to reddish brown. Length. 9-13 mm. Head (Figs. 35, 36). Anterior edge arcuate and slightly upturned, bisinuate, median emargination A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 49 v-shaped and stronger in females, clypeogenal junction entire. Clypeus transversely wrin kled on apical fourth in males or apical half in females, remainder finely punctate in both sexes. Clypeofrontal suture broadly arcuate, feeble in larger males, slightly carinate in females, with moderately posteriorly curved horn in well-developed males or 2 well separated low tubercles in females and smaller males. Anterior genal edge slightly longer than lateral, genal surface wrinkled on anterior fifth, remainder finely punctate. Vertex finely punctate on anterior half, convex in both sexes, slightly depressed along internal margin of eyes in males. Pronotum (Figs. 35, 36). Transverse, WIL = 1.6; anterior edge deeply emarginate and widely membranous behind eyes in males or slightly emarginate internally to eyes and membranous behind head insertion in females; punctures fine throughout, slightly larger on anterior angles. Anterior portion of disc with 4 tubercles, lateral ones much stronger than median ones located posteriorly, in larger males lateral tubercles lobiform and as close, or closer, to anterior edge of pronotum than posteromedian tubercles or with transverse row of 4 small, subequally sized, tubercles on anterior fourth, lateral tubercles lacking in extremely small females. Elytra. Subquadrate, L/W = 0.9. Striae fine,alutaceous, moderately impressed on disc and lateral declivities, wider and much deeper on apical declivity, punctures oval anteriorly and broadly oval to rounded posteriorly, approximately 3 times as wide as striae, separated by 2-5 diameters on disc. Intervals slightly convex, surface glossy, punctures fine and evenly distributed. Thoracic sterna. Proepisternum with posterolateral portion pubescent, external fifth smooth. Prosternum behind procoxa moder ately pubescent, with few longer setae on disc. Mesosternum impressed posteriorly, punc tures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, strongly carinate and alutaceous. Metasternum with disc finely punctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures moderately dense, punctures oval on anterior half rounded on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubescence moderately long and obliquely oriented; median lobe narrow, L/mW = 2.8, lateral margins convergent on posterior four-fifths and divergent on anterior fifth, anteriormost portion much thicker, surface finely and densely punctate, punctures larger anteriorly, finely alutaceous throughout, usually with row of thin and long setae along edge of mesocoxa, junction with mesosternum straight, narrowly declivous medially. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal third in males or basal fourth in females, apical spur inwardly bent on apical fifth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few setiferous punctures laterally and row of fine punctures along anterior edge, surface slightly alutaceous, sternite 6 finely punctate and alutaceous throughout, suture between sternites 5-6 shallowly carinate laterally, finely and coarsely microsculptured. Pygidium, W IL = 2.1 in males or 2.5 in females, punctures fineand dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 136- 138). Tubus approximately 2.0 times as long as parameres, apicoventral portion flat. Parameres in lateral view with apex narrow and obliquely truncate, ventral portion straight and denticulate at apex, dorsal portion straight and slightly declivous apically. Sigmoid sclerite of internal sac as in Figure 27 4.
MATERIAL STUDIED. 1900" a, 147 9 9 (AMNH, CMNC, CNCI, FGIC, HAHC, ISNB, MNHN, NMPC, SEMC, SMFD, SMTD, USNM, ZMHB).
Holotype a (CMNC): ECUADOR: PA ST., lOOOm, Llandia (17 km N. Puyo), 19.V II.1994, F. Genier, remnant rain for., feces tp./ HOLOTYPE, Ontherus diabolicus, F. Genier, 1994. Dissected.
Allotype 9 (CMNC): Same data as holotype. 50 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Paratypes: BOLIVIA: Chiquilas* (doubtful) (3 O' O' SMTD). COLOMBIA: Amazonas: Leticia, 700' (220m), 20-25.II.(19)72, Peck and Howden, dung trap (90'O', 3 9 9 HAHC), same as before except: 23.II-2.III.[19]74, Howden and Nealis, (1 O' HAHC), 23.II- 2.III.[19]74, Los Alpes, S. Peck, (3 O'O', 3 9 9 HAHC), 23.II-2.III.[19]74, Road End, S. Peck (1 O' HAHC). Meta: Villavicencio (1O' MNHN), same as before except: F. A. M., V. 1919, (1O' ISNB); 23km W. Villavicencio, Susamuco, lOOOm, 3-5.III.[19]72, S. and J. Peck, forest (19 HAHC). ECUADOR: No locality (20'O', 1 9 SMTD); Aguamo*, R. Haensch S. (1O' ZMHB); Chaca* (1 O' SMTD); Jarugui* (19 NMPC). Loja: Loja,AbM Gaujon (1 9 MNHN). Morona-Santiago: Macas (20'O' ZMHB). Napo: No locality, R. Haensch S. (1O' ISNB); Coca, R. Haensch S. (1O' SMTD) (1O',19 ZMHB); Limon cocha, 10-15.III.1975, J. M. Campbell (3O' O', 1 9 HAHC), same as before except; 800' (250m), 11.III.1976 (3 O' O', 3 9 9 CNCI), 15.III.1976 (1 9 CNCI), 18-24.VI.[19]76, S. Peck, 3 dng. tps. (14-16), nature trail for. (19 HAHC), 22-28.VI.[19]76, S. Peck, dng. tps. 22-24, virg. for., 2km N (14 O' O', 119 9 CMNC) (2O' O', 2 9 9 FGIC) (9 O'O', 129 9 HAHC); Rio Jatun Yacu, Rio Napo Watershed*, 5.III.[19]37, W. C. Macintyre (19 USNM); Km 7.3 Sarayacu-Loreto Rd (1200m), 11.VII.1994, F. Genier, cloud forest, feces tp. (1O', 3 9 9 FGIC), same as before except: 14.VII.1994 (6 O'O', 1 9 FGIC), 20.VII.1994 (1O' FGIC); Km 11.1 Sarayacu-Loreto Rd. (1200m) 20.VII.1994, F. Genier, cloud forest, feces tp. (1 O' FGIC); Km 25.4 Sarayacu-Loreto Rd. (950m), 12.VII.1994, F. Genier, cloud forest,feces tp.(11 O'O', 9 9 9 FGIC),same as before except: 14. VII.1994 (60'0', 49 9 FGIC); Sta. Cecilia, 340m, 8.VI-1.VIII.1968, William G. Saul (29 9 SEMC); 12km W. S. W. Tena, 600m, 6-11.VII.[19]76, S. Peck, for. dung traps, traps 30-32 (2O' O',2 9 9 HAHC),same as beforeexcept: traps 30-33 (2 O'O', 2 9 9 HAHC). Pastaza: Llandia (17 km. N. Puyo) (lOOOm), 19.VII.1994, F. Genier, remnant rain for.,feces tp. (68 O'O', 549 9 FGIC), same as before except: carrion tp. (3O' O', 4 9 9 FGIC), 20.VII.1994 (15O' O', 10 9 9 FGIC); lkm E. Mera, 1 lOOm, 13-17.VII.[19]76, S. Peck, ravine edge for., car. dng. tps. 44 (19 HAHC); 22km S. E. Puyo, 900m, 12-16.VII.[19]76, S. Peck, for. dng. tps. 39-41 (1 O' CMNC); 25krn N. N. E. Puyo, lOOOm, 4-13.VII.1976, S. Peck, for. dng. tps. 25-27 (5 O'O', 4 9 9 HAHC); 8km E. Rio Negro, 1400m, 1 Okm W. Pastaza (=Shell), 13-17.VII.[19]76, S. Peck, for. dng. tps. 46-47 (1O' CMNC); 9km E. S. E. Ve racruz, 900m, 22-24.VIII.1975, R. Webster, forest dung traps - 2 (1 O', 1 9 HAHC). Sucumbfos: Dureno, 150m,23-28.IX. l 977, Luis Pefia (1O' HAHC), same as before except: 23-30.IX.1977 (1 O', 19 HAHC). Tungurahua: Ambato (19 AMNH). Zamora Chinchipe: Sabanilla, F. Ohaus S., 24.IV.[19]05 (1O' USNM), same as before except: 26.IX.[19]05 (1 O' SMFD) (19 USNM), 28.IX.[19]05 (1O' SMFD) (1 O' USNM), 30.IX.[19]05 (1 O' NMPC), Loja Ostcordill., 27.IX.[19]05 (19 MNHN) (19 SMTD), 29.IX.[19]05 (1 9 MNHN) (19 NMPC); Zamora, Loja Ostcordill., Arsen, 26.IX.[18]95 (1 O' SMTD).
Additional data: Specimens were collected in dung traps set in forest,cloud forest and virgin forest and carrion-dung traps set in ravine edge forest.
MEXICANUS GROUP
DIAGNOSIS. The following combination of characters separates the next group of species: straight or slightly arcuate anterior edge of median lobe of metasternum; presence of tubercles and/or carina on pronotum in both sexes; presence of modified cephalic horn (compressed laterally or anteroposteriorly, bituberculate or widened apically) in well developed males, or 1 or 2 closely set tubercles or transverse horn in females; tridentate anterior tibia and dorsally expanded apical portion of parameres. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 51
REMARKS. Species of this group are similar externally to those of the previous group. However, the modified shape of parameres easily separates them from those of the Alexis group.
17. Ontherus (Caelontherus) compressicornis Luederwaldt, 1931 comb.nov. (Figs. 37, 38, 139-141, 275; Map 7)
Ontherus compressicornis Luederwaldt, 1931: 401 (description) Ontherus compressicornis Luederwaldt: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body uniformly reddish to dark brown. Pubescence fulvous. Length. 12-16 mm. Head (Figs. 37, 38). Anterior edge broadly arcuate in males or rounded in females, slightly upturned medially, bisinuate, median emargination v-shaped and stronger in females, clypeogenal junction notched. Clypeus reticulate throughout in males or transversely wrinkled throughout in females. Clypeofrontalsuture ill-defined, with a long sabre-like horn posteriorly with apex truncate, hornreduced to a laterally compressed conical process in smaller males and females. Anterior genal edge longer than lateral, genal surface shallowly punctate throughout in males or wrinkled on anterior three-fourths and remainder punctate in females. Vertex almost smooth in larger males, finely punctate anteriorly and smooth posteriorly in smaller males and females, deeply impressed along internal margin of the eyes in well-developed males, depressions much weaker, but always distinct, in smaller males and females. Pronotum (Figs. 37, 38). Transverse, W/L = 1.6; anterior edge slightly emarginate behind the eyes, margin almost vertical on short distance internally to the eyes, narrowly membranous behind head insertion in both sexes. Disc with 2 slightly oblique carinae covering two-thirds width of pronotum, lateral edges subdenticulate, inner edges tuberculate, carina sharp on lateral half, reduced to 2 arcuate carinae covering half width of pronotum, with lateral edges subquadrate and inner edges finely tuberculate in moderately sized males and females, smaller males and females armed with 2 closely set denticles on anterior portion of pronotum, punctures moderate and dense, confluent on anterior angles, fine and well-separated throughout in larger males. Elytra. Subquadrate, L/W = 1.0. Striae fine, shallowly impressed, slightly deeper on apical declivity, punctures rounded and umbilicate, much deeper than stria, especially on apical declivity, approximately 5 times as wide as stria, separated by 2-3 diameters on disc and by approximately their diameter on apical declivity. Intervals slightly convex, surface alutaceous, finely and densely punctate throughout. Thoracic sterna. Proepistemum with posterolateral portion moderately pubes cent, external fourth smooth. Prosternum behind procoxa moderately pubescent, several long setae along posterior edge. Mesosternum impressed posteriorly, punctures coarse and dense, except a small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternumirregularly punctate, alutaceous and strongly carinate. Metasternum with disc finely punctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures dense and oval on anterior half, scattered and broadly oval to rounded on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubescence moderately long, obliquely oriented; median lobe moderately wide, L/mW = 2.6, lateral margins convergent on posterior nine-tenths, parallel to slightly divergent on anterior tenth, surface finely and densely punctate, punctures larger anteriorly, finely alutaceous throughout, junction with mesostemum slightly arcuate, carinate and raised on its full width. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal fourth to third, apical spur inwardly bent on apical fourth in both sexes. Hinds legs unmodified. Abdomen. Sternites 2-5 with few coarse setiferous punctures laterally and row of scattered punctures along anterior edge, surface slightly alutaceous throughout, sternite 6 finely punctate and alu taceous throughout, suture between stemites 5-6 with few small and coarse punctures.
Pygidium, W/L = 1.9 in males or 2.1 in females, punctures fine and dense, surface glossy, 52 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 apical margin clearly delimited in both sexes. Male genitalia (Figs. 139-141). Tubus approximately 1.8 times as long as parameres, apicoventral portion concave. Parameres in lateral view with apex obliquely truncated, ventral portion straight, dorsal portion straight basally and produced into hook apically. Sigmoid sclerite of internal sac as in Figure 275.
MATERIAL STUDIED. 45 CJ' CJ', 34 9 9 (CASC, CMNC, HAHC, ISNB, MNHN, MZSP, NMPC, PAIC, SMTD, WDEC).
TYPE MATERIAL Ontherus compressicornis Luederwaldt: Holotype a (MZSP): Ecuador, Baron/ Staudinger, 1927I Ontherus nevinsoni, Wat./ TIPO/ Ontherus a, compressicornis, Lueder., Lueder. det. 29/ 16855. Holotype studied.
NO N- TYPE MATERIAL Distribution: ECUADOR: Caiiar: J avin (900m). Cotopaxi: 112km W Latacunga ( =14km W Pilalo) (1550m). Pichincha: 23km E Alluriquin (Chiriboga Road) (1400m); 28km E Alluriquin (Chiriboga Road) (1580m); 3 lkm NE Alluriquin (Chiriboga Road) (l770m); 3km E Tandapi (=Cornejo Astorga) (1310m); 1 lkm E Tandapi.
Additional data: Specimens were collected using carrion and dung traps set in forests, moss forests and wet ravines. Specimens collected in February, June, July.
18. Ontherus (Caelontherus) pilatus sp.nov. (Figs. 1, 39, 40, 142-144, 276; Map 8)
ETYMOLOGY. Pilatus, ("armed with a javelin") Latin adjective, refers to peculiar aspect of apex of cephalic development in well-developed males.
DESCRIPTION. Colour. Body uniformly reddish to black, upper side usually with moderate iridescent tinge. Pubescence fulvous. Length. 9-13 mm. Head (Figs. 1, 39, 40). Anterior edge broadly arcuate, almost flat, strongly bisinuate, median emargination v-shaped, clypeogenal junction notched. Clypeus transversely and rather moderately reticu late throughout. Clypeofrontal suture ill-defined, with long backwardly curved horn, com pressed laterally on basal half and spatulate on apical half, reduced to laterally compressed conical process in moderately developed males and to swelling in smallest males or with transverse plate with apex arcuate, apex notched in moderately developed females, reduced to weakly bituberculate swelling in smallest females. Anterior genal edge subequal to lateral, genal surface strongly alutaceous, shallowly punctate throughout in males or weakly wrinkled on anterior three-fourths in females. Vertex strongly alutaceous and finely punctate anteriorly, glossy posteriorly, deeply impressed along internal margin of eyes in well-developed males, depressions much weaker but always distinct in smaller males and females. Pronotum (Figs. 1, 39, 40) Transverse, W /L= 1.6; anterior edge deeply emarginate behind eyes, margin almost vertical on short distance internallyto eyes, widely membranous behind eyes in both sexes. Disc with 2 transverse and slightly oblique carinae each covering less than one-fourth width of pronotum, lateral edges subdenticulate, inner edges tuberculate, carinae sharp and irregular on lateral three-fourths, reduced to 2 sublobate tubercles laterally on anterior third in addition to small tubercle position slightly posteriorly on midline, or transverse row of 3 small and sharptubercles on anterior fourthin females,convex in smallest individuals, surface usually strongly alutaceous throughout, punctures fine and evenly distributed. Elytra. Subquadrate, L/W = 1.0. Striae fine, shallowly impressed, slightly deeper on apical declivity, punctures rounded and slightly umbilicate, as deep as stria, especially on apical declivity, approximately 5- 6 times as wide as stria, separated by 2-3 diameters on disc and by less than their diameter on apical declivity. Intervals slightly convex, surface alutaceous, finely and densely punctate throughout. Thoracic sterna. Proepisternum A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 53 with posterolateral portion moderately pubescent, external fourth smooth. Prosternum behind procoxa moderately pubescent, several long setae along basal third. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of meso coxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, alutaceous and strongly carinate. Metasternum with disc finelypunctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures moderately dense and oval, each with anterior edge finely carinate on anterior half, broadly oval to rounded on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubescence moderately long, obliquely oriented; median lobe moderately wide, L/mW = 2.4, lateral margins convergent throughout, surface finely and densely punctate, punctures larger anteriorly, alutaceous throughout, junction with mesosternum slightly arcuate, carinate laterally, declivous medially. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal half, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2- 5 with few large and coarse setiferous punctures laterally and along anterior edge, surface alutaceous throughout, sternite 6 finely punctate and alutaceous throughout, suture between sternites 5-6 with few moderate and coarse irregular punctures. Pygidium, W/L= 1.9 in males or 2.2 in females, punctures fine and dense, surface slightly alutaceous, apical margin clearly delimited in both sexes. Male genitalia (Figs. 142-144 ). Tubus approximately 1.7 times as long as parameres, apicoventral portion unmodified. Parameres in lateral view with apex truncated, ventral portion straight and with small denticle apically, dorsal portion concave basally and with hook apically. Sigmoid sclerite of internal sac as in Figure 276.
MATERIAL STUDIED. 19 a a, 11 9 9 (CMNC, FGIC, HAHC).
Holotype a (CMNC): ECUADOR: PINCHINCHA, 23km E. Alluriquin, Chiriboga Rd., 4600' (1400m), 19-27.VI.1975, S. Peck, dung trap 37I ex. collection, H. and A. Howden/ HOLOTYPE, Ontherus pilatus, F. Genier, 1992. Dissected.
Allotype 9 (CMNC): Same data as holotype.
Paratypes: ECUADOR: Pichincha: 23km. E. Alluriquin, Chiriboga Rd, 4600' (1400m), 19-27.VI.1975, S. Peck, dung trap 37 (8 a a, 5 9 9 HAHC) (1 a, 1 9 FGIC); 14km N. E. Alluriquin, Chiriboga Rd., 19-27.VI.1975, S. Peck, wet ravine, dng. tp. 36 (8 a a, 3 9 9 HAHC) (1 a, 1 9 FGIC).
Additional data: Specimens were collected using a dung trap set in a wet ravine.
19. Ontherus (Caelontherus) trituberculatus Balthasar, 1938 comb.nov. (Figs. 41, 42, 145-147, 277; Map 9)
Ontherus trituberculatus Balthasar, 1938: 220 (description) Ontherus trituberculatus Balthasar: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body uniformly reddish brown to black, upper side usually with moderate iridescent tinge. Pubescence fulvous. Length. 9-13 mm. Head (Figs. 41, 42). Anterior edge broadly arcuate, almost flat, strongly bisinuate, median emargination v shaped, clypeogenal junction notched. Clypeus transversely and rather moderately reticulate throughout. Clypeofrontal suture ill-defined, with long backwardly curved horn,compressed laterally with apex acuminate or obliquely truncate, with 2 broadly triangular lobes on posterolateral edges on apical third, reduced to laterally compressed conical process in moderately developed males and to swelling in smallest males or with transverse plate with apex rounded, apex notched in moderately developed females, reduced to weakly bituber culate swelling in smallest females. Anterior genal edge subequal to lateral, genal surface 54 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 finely alutaceous, shallowly punctate throughout in males or weakly wrinkled on anterior three-fourths in females. Vertex strongly alutaceous and finely punctate anteriorly, glossy posteriorly, deeply impressed along internal margin of eyes in well-developed males, depressions much weaker but always distinct in smaller males and females.
Pronotum (Figs. 41, 42). Transverse, W/L = 1.6; anterior edge deeply emarginate behind eyes, margin almost vertical on short distance internally to eyes, widely membranous behind eyes in both sexes. Disc with 3 tubercles on anterior three-fifths, lateral tubercles slightly larger and sublobate, sometimes feebly produced laterally into short carina, median tubercle sharp and position slightly behind lateral ones, reduced to straight transverse row of 3 tubercles on anterior third, median tubercle larger than lateral ones, or transverse row of 3 sharp tubercles on anterior fourth in females, smallest individuals with single median tubercle or disc convex, surface moderately alutaceous throughout, punctures fineand evenly distributed. Elytra. Subquadrate, L/W = 0.9. Striae fine, shallowly impressed, slightly deeper on apical declivity, punctures broadly oval to rounded and slightly umbilicate, as deep as stria, approximately 3 times as wide as stria, separated by 2-3 diameters on disc and by less than their diameter on apical declivity. Intervals slightly convex, surface alutaceous, finely and densely punctate throughout. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent, external fifth smooth. Prosternum behind procoxa moderately pubescent, several long setae along basal third. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, alutaceous and strongly carinate. Metasternum with disc finely punctate and alutaceous throughout, convex in both sexes; lateral lobes with punctures moderately dense and oval, each with anterior edge finely carinate on anterior half, scattered and broadly oval to rounded on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradu ally weakening posteriorly, pubescence moderately long, obliquely oriented; median lobe moderately wide, L/m W = 2.2, lateral margins convergent throughout, surface finely and densely punctate, punctures larger and setiferous anteriorly along mesocoxal edges, alutaceous throughout, junction with mesostemum slightly arcuate, widely carinate throughout. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal third to fourth, apical spur inwardly bent on apical fourthin both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few large and coarse setiferous punctures laterally and row of smaller, entire to fused, punctures along anterior edge, surface slightly alutaceous throughout, sternite 6 finely punctate and alu taceous throughout, suture between stemites 5-6 with few small irregular punctures.
Pygidium, WIL = 1.9 in males or 2.3 in females, punctures fine and dense, surface slightly alutaceous, apical margin clearly delimited in both sexes. Male genitalia (Figs. 145-147). Tubus approximately 1.4 times as long as parameres, apicoventral portion unmodified. Parameres in lateral view with apex truncate and concave, ventral portion straight and produced in denticle apically, dorsal portion concave and produced apically. Sigmoid sclerite of internal sac as in Figure 277.
MATERIAL STUDIED. 33 a a, 19 9 9 (BDGC, BMNH, CMNC, HAHC, ICCM, NMPC, SMTD).
TYPE MATERIAL Lectotype a (SMTD): Cachabe, low c., XII. 96., (Rosenberg)/ Coll. C. Felsche, Kauf 20, 1918/ TYPUS/ Ontherus 3-tuberculatus, n. sp. Typus, Dr. V. Balthasar det./ LECTOTYPE, Ontherus trituberculatus, Balthasar, 1938, Des. F. Genier, 1991. Type material studied, lectotype and paralectotypes here designated.
Paralectotypes: same data as Lectotypes (2a a, 1 9 SMTD) (1 a, 1 9 NMPC). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 55
• 0. trituberculatus
• 0. monilistriatus
MAP 9. Known distribution of Ontherus trituberculatus and 0. monilistriatus.
NON-TYPE MATERIAL Distribution: COLOMBIA: Valle del Cauca: Bajo Calima (50m); 70km E Buanaventura (Anchicaya Dam) (1200m). ECUADOR: Carchi: Chical (0° 56' N 78° 11'W) (1250m); 18km SEMaldonado (2420m). Esmeraldas: Cachabe. Pichincha: 5.3km on Via Pachija* (2800-3000m); 85km NW Quito (on Puerto Quito Rd.) (1520m); l 13km NW Quito (on Puerto Quito Rd.) (790m); 4km S E Santo Domingo (500m); 16km S E Santo Domingo (Tinalandia) (680m.).
Additional data: Specimens were collected using dung traps set in a forest during day and night, day only, and in a swamp. Specimens collected in January, June to August, December.
20. Ontherus (Caelontherus) mexicanus Harold, 1868 comb.nov. (Figs. 43, 44, 148-150, 278; Map 10)
Ontherus mexicanus Harold, 1868b: 80 (description) Ontherus mexicanus Harold: Gemminger and Harold, 1869: 1008 (catalog) Ontherus mexicanus Harold: Harold, 1869a: 59 (comment) Ontherus mexicanus Harold: Harold, 1869c: 503, 504 (redescription and distribution) Ontherus mexicanus Harold: Bates, 1887: 50 (distribution) Ontherus mexicanus Harold: Gillet, 191 lb: 58 (catalog) Ontherus mexicanus Harold: Luederwaldt, 1931: 395, 397 (key) Ontherus mexicanus Harold: Halffter and Edmonds, 1982: 86, 89-90 (biology)
REDESCRIPTION. Colour. Body uniformly reddish brown to black, sometimes with iridescent tinge on elytra. Pubescence fulvous. Length. 10-16 mm. Head (Figs. 43, 44). Anterior edge arcuate, slightly upturned and bisinuate, median emargination shallow, clypeogenal junction finely notched. Clypeus transversely wrinkled on anterior half in males or two-thirds in females, remainder of surface punctate. Clypeofrontal suture ill-defined, with backwardly curved and acute horn, trapezoidal in cross section and small and smooth 56 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 swelling laterally in males, or clypeofrontal carinate, with 2 well-separated acute tubercles medially in females, median cephalic development reduced to, 1 in males or 2 in females, small median swellings in smaller individuals. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior half, posterior half finely alutaceous and punctate. Vertex alutaceous and finely punctate on anterior half, smooth and convex on posterior half, internal edges of eyes impressed in larger males. Pronotum (Figs. 43, 44). Transverse, W/L = 1.6; anterior edge deeply emarginate and widely membranous behind eyes in males or margin slightly sinuous and narrowly membranous behind head insertion in females, anterior sulcus strongly microsculptured and clearly delimited posteriorly, finely granulose, posterior margin strongly sinuous medially in larger females. Disc with 2 almost longitudinal carinae with anterior and posterior portions feebly to moderately tuberculate, laterally with 2 small sublobate tubercles in males or with transverse row of 4 small tubercles in females, development of smaller individuals reduced to pair of vaguely delimited and well-separated swelling in males or 2 small and closely set tubercles medially on anterior fifth in females, punctures fine and evenly distributed throughout, except anterior declivity almost smooth in larger males. Elytra. Subquadrate, L/W = 1.0. Striae fine, moderately impressed on disc, deeper on apical declivity, punctures rounded and umbilicate, as deep as stria, approximately 3 times width of stria, separated by 1-2 diameters on disc and less than their diameter on posterior portion of apical declivity. Intervals convex, surface finely and evenly punctate throughout. Thoracic sterna. Proepistemum with posterolateral portion moderately pubes cent and alutaceous, external fourth smooth. Prostemum behind procoxa moderately pubes cent, several long setae along posterior edge. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, slightly alutaceous and strongly carinate. Metastemum with disc finely and scarcely punctate medially, slightly alutaceous, convex in both sexes; lateral lobes with punctures oval, posterior edge vaguely delimited, approximately evenly spaced on anterior half, rounded and clearly delimited, evenly distributed but leaving smooth area on disc, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubes cence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.5, lateral margins convergent throughout, surface finely and densely punctate, punctures larger anteriorly, strongly alutaceous anteriorly, less so posteriorly, junction with mesostemum slightly arcuate, carinate laterally. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal third, apical spur inwardly bent on apical fourthin both sexes. Posterior legs unmodified. Abdomen. Stemites 2-5 with few coarse setiferous punctures laterally and row of smaller punctures along anterior edge, surface alutaceous throughout, sternite 6 finely and densely punctate, surface alutaceous, minimum width less than half maximum width, suture between sternites 5-6 with row of small irregular punctures laterally. Py gidium, W/L = 2.1 in males or 2.2 in females, punctures fine and dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 148-150). Tubus approximately 1.8 times as long as parameres, apicoventral portion slightly depressed. Parameres in lateral view with apex enlarged and truncate, ventral portion straight, dorsal portion concave, apicodorsal angle strongly swollen and posteriormost portion produced into small denticle. Sigmoid sclerite of internal sac as in Figure 278.
MATERIAL STUDIED. 620a a, 510 Q Q (BDGC, CASC, CMNC, CNCI, DEIC, FGIC, HAHC, ICCM, ISNB, LACM, MAMC, MCZC, MNHN, NMPC, SEMC, SMTD, TAMU, USNM, WBWC, ZMHB, ZMUC, ZSMC).
TYPE MATERIAL Holotype Q (BMNH?): Mexico. Holotype not studied (Holotype not found). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 57
• O. mexicanus
• 0. sextuberculatus
MAP 10. Known distribution of Ontherus mexicanus and 0. sextuberculatus.
NON-TYPE MATERIAL Distribution: GUAT EMALA: Alta Verapaz: 3.7mi. S Coban (1370m); Senahu; 5km S Tactic (Patal) (1370m). Baja Verapaz: 4.3km E Purulha (1680m); 7km N E Purulha (1500m); 8km S PurulM (1660m); 16.5km N Salama (1650m). MEXICO: Chiapas: 5mi. E Jct. Hwy. 17 and 190; 6.6mi. WEI Bosque (1460m); 3mi.W Jitolol; Lagos de Montebello, Lago Pojoj (1500m); 8mi. N Puebla Nuevo S (l 830m); San Cristobal (las Casas) (2100- 2160m); San Cristobal las Casas (2190m); 3mi. W San Cristobal (las Casas); 3mi. SW San Cristobal (las Casas). Hidalgo: Ixtlahuaco (l 450m); lOmi. E Jacala; lOmi. NE Jacala; 20mi. NE Jacala; Laguna Atezca* (l 300m); 6.4mi. S Tenango de Dorias (3000m); Tlanchinol; Zacualtipan (2060m); 3km. S Zacualtipan. Oaxaca: km 140 on Hwy. 175* (1220m); 6mi. S Valle Nacional (600m); 12mi. S Valle Nacional (910m); 13mi. S Valle Nacional (1lOOm); 15mi. S Valle Nacional (1220m); 18mi. S Valle Nacional (1460m); 26km E Valle Nacional (1220m). Puebla: Acaxochitlan; 4.4mi. S W Huauchinango (1700m); Necaxa; Nuevo Necaxa* (1200m); Sierra de Zacapoaxtla; Teziutlan; 15mi. E Teziutlan; 6mi. W Teziutlan; 4mi. NEXicotepec de Juarez (1130m); 4mi. S Zacapoaxtla, (2070m). QuintanaRoo: Landa de Matamoros (1520m); 25mi. E Landa de Matamoros (1520m). San Luis Potosi: 13mi.W Xilitla (1460m); 20km S W Xilitla (1520m); 20mi. W Xilitla (1620m). Tamaulipas: Estacion. Biologica Canindo (nr. San Jose, 7.5 km W Gomez Farias) (1400m); 6mi. NW Gomez Farias (Rancho del Cielo) (1130m). Veracruz: Cordoba; Laguna de Catemaco (360m); Los Tuxtlas (Estacion de Biologia Tropical UNAM)*; lmi. S Pueblo Calcahualco; 20km N E Tlanchinol*.
Additional data: Specimens were collected using carrion bait traps, human dung traps, light traps and in cow dung. Some of these traps were set in cloud forests, Liquidambar forests, pine cloud forests, sycamore forests, tropical deciduous forestsand moist ravine oak forests. They were also collected using human dung traps in cloud forests and flight intercept traps in montane tropical forests. Specimens collected in May to October. 58 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
REMARKS. Ontherus mexicanus Harold and 0. sextuherculatus sp.nov. are closely related. Although similar externally, the allopatric distribution pattern and their extremely different aedeagi offer no doubt about their specific status. Ontherus mexicanus Harold can also be distinguished from 0. sextuberculatus sp.nov. by these additional characters: the apical portion of the eighth striae is deeply impressed and abruptly terminated (normally impressed in 0. sextuberculatus sp.nov. ), and its larger average size.
21. Ontherus (Caelontherus) sextuberculatus sp.nov. (Figs. 45, 46, 151-153, 279; Map 10)
ETYMOLOGY. Sextuberculatus, (six+ tubercles) Latin adjective, refers to the pronotal development aspect of males.
DESCRIPTION. Colour. Body uniformly dark brown to black, sometimes with iridescent tinge on elytra. Pubescence fulvous. Length. 8-13 mm. Head (Figs. 45, 46). Anterior edge arcuate, slightly upturned medially and subdenticulate, median emargination v-shaped, clypeogenal junction finely notched. Clypeus transversely wrinkled on anterior half in males or anterior two-thirds in females, remainder of surface punctate. Clypeofrontal suture ill-defined, with backwardly curved horn, trapezoidal in cross-section, posterior edge slightly enlarged apically and small and smooth swelling laterally in males or clypeofrontal carinate, with 2 well-separated tubercle medially in females. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior half, posterior half finely alutaceous and punctate. Vertex alutaceous and finely punctate on anterior half, smooth and convex on posterior half, internal edges of eyes impressed in larger males. Pronotum (Figs. 45, 46)
Transverse, W /L = 1.6; anterior edge deeply emarginate and widely membranous behind eyes in males or margin slightly sinuous and narrowly membranous behind head insertion in females, anterior sulcus strongly microsculptured and clearly delimited posteriorly, finely granulose, posterior edge sinuous medially in females. Disc with 2 approximately oblique carinae with anterior and posterior portions tuberculate, laterally with 2 small sublobate tubercles in males or with transverse row of 4 small tubercles on anterior fifth in females, punctures fine and evenly distributed throughout, except anterior declivity almost smooth in larger males. Elytra. Subquadrate, L/W = 0.9. Striae fine, moderately impressed on disc, slightly deeper on apical declivity, punctures rounded and umbilicate, as deep as stria, approximately 3 times as wide as stria, separated by 2-3 diameters on disc and less than their diameter on posteriormost portion of apical declivity. Intervals convex, surface finely and evenly punctate throughout. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent and alutaceous, external fifth smooth. Prosternum behind procoxa moderately pubescent, several long setae along posterior edge. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubes cence fine and short, except 2-5 longer setae along mesocoxa. Mesepisternum irregularly punctate, slightly alutaceous and strongly carinate. Metasternum with disc finely and scarcely punctate medially, slightly alutaceous, convex in both sexes; lateral lobes with punctures oval, each with posterior edge ill-defined, evenly spaced on anterior half, rounded and clearly delimited, evenly distributed but with small smooth area on disc, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubes cence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.5, lateral margins convergent throughout, surface finely and densely punctate, punctures larger anteriorly, strongly alutaceous anteriorly, less so posteriorly, junction with mesosternum slightly arcuate, strongly carinate laterally. Legs. Anterior tibia tridentate in males, quadri dentate in females, ventromedial carina lacking intervening setae on basal fourth, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few coarse setiferous punctures laterally and row of smaller entire to confluent A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 59 punctures along anterior edge, surface alutaceous throughout, sternite 6 finely and densely punctate, surface alutaceous, minimum width less than half maximum width in males, suture between stemites S- 6 with row of small irregular punctures laterally. Pygidium, W/L = 1.9 in males or 2.1 in females, punctures fine and dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 1Sl-1S3). Tubus approximately 1.8 times as long as parameres, apicoventral portion flattened. Parameres in lateral view with apex enlarged and truncate, ventral portion convex and dentate at apex, dorsal portion concave, apicodorsal angle produced into denticle. Sigmoid sclerite of internal sac as in Figure 279.
MATERIAL STUDIED. 228et et, 241 Q Q (INBC, USNM).
Holotype et(INBC): Sector Cerro Cocori, Fca, de E. Rojas, Prov. Limon, COSTA RICA, E. Rojas, Die 1990, L-N-286000,S67SOO/ INBIO CRlOOO 396734 (bar code label)/ HOLOTYPE, Ontherus sextuberculatus, F. Genier, 1994. Dissected.
Allotype Q (INBC): Same data as holotype except: 396624 (bar code label).
Paratypes: Costa Rica: Cartago: Grano de Oro, Chirrip6, Turrialba(l 120m) 8-30.VII.1992, P. Campos, L-N 2002SO, S9S900 (2etet, 3 QQ INBC), same as before except: 8- 31.VIII.1992 (1et, 1 Q INBC), IX.1992 (1et, 3 Q Q INBC), X.1992 (13 etet, lSQ Q INBC), II.1993 (12etet, 13 QQ INBC), VII.1993 (1et INBC). Guanacaste: Rfo San Lorenzo, R. F. Cord. Guanacaste (Tenorio) (lOSOm), VII.1991, C. Alvarado, L-N-287800, 427600 (1 et INBC), same as before except: Tierras Morenas, Z. P. Tenorio, XII.1992, G. Rodriguez, (1Q INBC). Heredia: Est. El Ceibo, Braulio Carrillo P. N. (400-600m), IX.1989, R. Aguilar and M. Zumbado, S27700, 2S6SOO (1et INBC), same as before except: X.1989 (9 et et INBC), XI.1989 (3 etet, 2 QQ INBC). Limon: Est. Jalova, P. N. Tortuguero (Om), VII.1990, M. Barrelier, L-N-2S86S0,602100(1 Q INBC); Hamburgfarm, Reventazon, Ebene, 27.VII.[19]3S, F. Neverman (Seta, SQQ USNM); Rio Blanco (4km S. W.) 8.V.1987, J. F. Corrales (1 et INBC); Rfo Sardinas, R. N. F. S. Barra del Colorado (lOm), 14-22.VIII.1993, F. Araya, L N 291SOO_S64700, #2306 (1 Q INBC), same as before except: 12-20.IX.1993, #23SS (3et et, 3 QQ INBC), 11-19.X.1993, #2398 (Setet, 3 QQ INBC), 10-12.XI.1993, #2446 (Seta, 6QQ INBC); Sector Cerro Cocori, Fca. de E. Rojas, XI.1990, E. Rojas, L-N-286000,S67SOO (7Q Q INBC), same data as before except: XII.1990 (82a et, 91Q Q INBC), I.1991 (9etet, 10Q Q INBC), III.1991 (70etet, 6S Q Q INBC), V.1991 (1Q INBC), VI.1991 (1et, 1Q INBC), VIII.1991 (1Q INBC), IX.1991 (1 Q INBC), I.1992 (1 Q INBC), S.VI-SVII.1992, Tp. malaise (1 Q INBC), 12-31.VIII. l 992 (l Q INBC), X.1992 (3 Q Q INBC), III. 1993 ( 1 et INBC), IX.1993 ( 1et INBC). San Jose: P. N. B. Carrillo, Lat. l0°09'7S", Long. 84°S7'1S", 14.XII.1988, Col. J. F. Corrales (2etet INBC).
REMARKS. Ontherus mexicanus seems to be restricted to the cloud foresthabitat, whereas 0. sextuberculatus has been collected from sea level to 1120 m.
22. Ontherus (Caelontherus) lunicollis sp.nov. (Figs. 47, 48, 1S4-1S6, 280; Map 11)
ETYMOLOGY. Lunicollis, (crescent-shaped + bearing) Latin adjective, refers to male pronotal development.
DESCRIPTION. Colour. Body uniformly reddish brown to black, sometimes with irides cent tinge on elytra. Pubescence fulvous. Length. 9-16 mm. Head (Figs. 47, 48). Anterior edge broadly arcuate, flat to slightly upturned, bisinuate, median emargination broad and shallow in males or broadly v-shaped in females, clypeogenal junction almost straight. Clypeus strongly and transversely wrinkled on anterior half in males or anterior three-fourths 60 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 in females, remainder moderately punctate in males or strongly punctate in females. Cly peofrontal suture ill-defined, with backwardly curved horn, bifurcate apically, transverse in larger individuals to feebly compressed laterally formoderately sized individuals in males or with 2 well-separated acute tubercles medially in females, cephalic development reduced to 1 conical process in males or 2 in females, small median tubercles in smaller individuals. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior half, posterior half finely punctate in males or with punctures almost confluent in females. Vertex moder ately punctate on anterior half, smooth and convex on posterior half, internal edges of eyes deeply foveolate in larger males. Pronotum (Figs. 47, 48). Transverse, W/L = 1.6; anterior edge slightly emarginate and membranous behind eyes, margin almost vertical in lateral view on short distance internally to eyes in males or margin slightly sinuous and narrowly membranous behind head insertion in females, anterior sulcus narrow, strongly microsculp tured and clearly delimited posteriorly. Disc with 4 tubercles, lateral tubercles large, close to anterior edge, preceded by concavity anterolaterally, median tubercles small and located posteriorly to lateral tubercles, posterior edge of declivity arcuate in males or with pair of closely set and low tubercles on anterior two-fifths, anterior declivity as wide as head insertion in females, development of smaller individuals reduced to pair of vaguely delimited transverse swellings on anterior fifth, punctures fine and evenly distributed throughout, except anterior declivity almost smooth in larger males, punctures slightly larger on anterior angles in females. Elytra. Subquadrate, L/W = 1.0. Striae fine, moderately impressed on disc, slightly deeper on apical declivity, punctures round and umbilicate, as deep as stria, approximately 5 times as wide as stria, separated by 2-3 diameters on disc and less than their diameter on posteriormost portion of apical declivity. Intervals moderately convex, surface finely and evenly punctate throughout, feebly to moderately alutaceous. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent and alutaceous, externalfourth smooth. Prosternum behind procoxa moderately pubescent, several long setae along posterior edge. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternumirregularly punctate, alutaceous and strongly carinate. Metasternum with disc finely and scarcely punctate medially, convex in both sexes; lateral lobes with punctures oval, almost evenly spaced on anterior half, rounded, restricted to metacoxal edge, leaving scarcely punctate area on disc on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubes cence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.8, lateral margins convergent throughout, surface finely and densely punctate, punctures larger anteriorly, strongly alutaceous anteriorly, less so posteriorly, usually with distinct longitudi nal carina medially on anterior third, junction with mesosternum slightly arcuate, finely carinate. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal third, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few coarse setiferous punctures laterally and row of small entire to confluent punctures along anterior edge, surface alutaceous throughout, sternite 6 finely and densely punctate, surface glossy, suture between sternites 5-6 with row of small irregular punctures laterally. Pygidium, W/L = 2.1 in males or 2.5 in females, punctures fine and dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 154-156). Tubus approximately 2.0 times as long as parameres, apicoventral portion slightly depressed. Parameres in lateral view with apex obliquely truncate, ventral portion slightly concave, dorsal portion concave, apicodorsal angle strongly produced into small denticle. Sigmoid sclerite of internal sac as in Figure 280.
MATERIAL STUDIED. 210 a a, 173 Q Q (AMNH, BDGC, BMNH, CMNC, CNCI, FGIC, HAHC, ISNB, MCZC, MNHN, MZLU, NMPC, PAIC, SMTD, USNM, UZMH, ZMAS, ZMHB, ZMUC, ZSMC). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 61
• 0. lunicollis
• 0. pseudodidymus
MAP 11. Known distribution of Ontherus lunicollis and 0. pseudodidymus.
TYPE MATERIAL Holotype O' (CMNC): 20 km. W. Silvia, 6000' (1830m), Cauca, Colombia, July 17, 1970, H. and A. Howden/ ex collection, H. and A. Howden/ HOLOTYPE,Ontherus lunicollis, F. Genier, 1992. Dissected.
Allotype Q (MNHN): Villa Elvira, Cauca/ Museum Paris, 1936, Coll. A. Boucomont/ ALLOTYPE, Ontherus lunicollis, F. Genier, 1992.
Paratypes: No locality (1O',2 Q Q MNHN). BRAZIL (doubtful): No locality (1O' NMPC). COLOMBIA: No locality (1O', 3 Q Q AMNH) (1 Q BMNH) (5O' O', 1 9 ISNB) (20'O' MCZC) (20' O', 1 9 MNHN) (40' O', 1 9 SMTD) (1O' UZMH) (2 Q Q ZMUC) (1O' ZSMC), same as beforeexcept: Felipe Ovalle Q., Ac. 33501 (2 O'O', 1 Q AMNH);Aguatal * (1 9 SMTD); Caucathal*, (20' O', 1 Q ISNB); Rio Vitaco* (1O' MNHN); Santa Rosa* (1 O' MNHN). Antioquia: Abejorral (1O' MNHN); Medellin, H. Daniel, [19]37, usnm 145398, # 65 (1O', 3 9 Q USNM); Rfonegro (1O' MNHN); Santa Barbara (1 O' MNHN). Boyaca: Muzo (3 Q Q NMPC). Caldas: Aguadas (1O' ISNB); Manizales (20' O', 1 9 MNHN), same as beforeexcept: Salamina ( 1 Q MNHN), A. M. Patino (2 O'O', 2 Q 9 FGIC) (460' O', 66 9 9 MNHN) (20' O', 2 9 Q NMPC). Cauca: No locality (100'O', 7 9 9 ISNB) (3 O'O', 1 9 NMPC) (5O' O', 2 9 Q SMTD) ( 4 O'O', 2 Q 9 ZMHB); El Tampo (=El Tambo?), Popayan, 1700m, 15.IX.1932, coll. Kj. von Sneidern (1 Q MZLU); Popayan (3 O' O' SMTD), same as before except: Abbe Gaujon, 1899 (1 O' MNHN); Silvia, 6000' (1830m),22.II.1970, H. Howden (1O' HAHC);20km W. Silvia,6000' (1830m), 22.II.1970, H. Howden (2 O' O' HAHC), same as before except: 17 .VII.1970, H. and A. Howden (3 O'O', 1 Q HAHC), 18.VII.1970 (3 Q 9 HAHC), 19.VII.1970 (1 9 HAHC); Villa Elvira* (20' O' NMPC) (1O' SMTD) (20' O' ZMHB). Cundinamarca: La Mesa* (1 9 MNHN); Sesamuco,23km W. Villavicencio, lOO m,3-5.III.[19]72, forest, S. and J. Peck (1 Q HAHC); Tecondama Falls*, 30km S.W. Bogota, 27.II-6.III.[19]72, forest dung trap, S. and J. Peck (1 O',1 Q HAHC). Distrito Especial: Bogota (IO' MNHN), same as before except: Bieht (1 9 ZMHB), Buq., 9419 (20'O' ZMHB), De Haan, 26629 (20' O', 1 9 ZMHB), 62 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Parzudacki,1839 (1 9 MNHN),de Savanna a Bogota,Dr. 0. Thieme,1877 (1a MNHN), P. Ringter, Halle (1 9 NMPC). Meta: Villavicencio (3 aa MNHN). Narillo: Pasto (2 9 9 ISNB). Norte de Santander: Guayabal (3 aa, 1 9 MNHN) (2 9 9 NMPC); Quindio: llkm E. Calarca, 7000' (2130m), 7-11.11.[19]74, S. and J. Peck, DT988-9 (2aa, 1 9 CNCI), same as before except: 9.111.[19]74, DT993-4 (3a a, 1 9 HAHC); 15km E. Calarca, 8000' (2440m), 7.III.[19]74, S. Peck, DT993-4 (8a a, 8 9 9 CNCI) (7 a a,4 9 9 HAHC), same as before except: 7-10.III.[19]74, S. and J. Peck (2aa, 3 9 9 CNCI); lkm S. Calarca, 5000' (1520m), 8.III.[19]74, S. and J. Peck (12aa, 4 9 9 HAHC), same as before except: 8-10.III.[19]74, DT990-2 (4aa, 1 9 CNCI). Tolima: No locality (1 a,19 SMTD); San Antonio (2a a, 2 9 9 FGIC) (1a MNHN) (3 9 9 NMPC) (4aa, 2 9 9 SMTD) (21aa,89 9 ZMHB). Valle del Cauca: Cali, X.1908 (2 9 9 ZMHB), same as before except: IX-XII.[18]94, Rosenberg (4aa, 2 9 9 SMTD), 1896, Burger S. (1 9 ZMHB; Maquanal, Las Pavas (1 9 MNHN); Pichinde, Xll.[18]91, 5000ft., W. Rosenberg (1 9 NMPC); n[ea]r. Pichinde, 5000' (1520m), 18.Vll.1970, H. and A. Howden (1a HAHC); Restrepo, 1600m, 6-7.Vll.1983, B. Gill (1 a BDGC); Tocota (3a a,1 9 ZMHB).
Additional data: Some specimens were collected in a dung trap.
REMARKS. The material studied included 5 specimens not considered part of the type series. Three of these specimens are from an unspecified locality in Costa Rica, (1a, 2 9 9 SMTD), one (1 a SMTD) is from an unspecified locality in Panama, and the last specimen (1 9 SMTD) is from Chiriqui (Panama). These specimens are old,and until fresh specimens are collected in Central America I consider 0. lunicollis sp.nov. to be strictly South American. The male cephalic horn in larger individuals of 0. lunicollis sp.nov. is always slender and bifurcate apically,whereas in the sister species 0. pseudodidymus sp.nov. it is usually transverse.
23. Ontherus (Caelontherus) monilistriatus sp.nov. (Figs. 49,50, 157-159, 281; Map 9)
ETYMOLOGY. Monilistriatus, (necklace+ furrow) Latin adjective,refers to the aspect of elytral striae.
DESCRIPTION. Colour. Body uniformly black, with faint iridescent tinge on elytra. Pubescence fulvous. Length. 15-16 mm. Head (Figs. 49,50). Anterior edge broadly arcuate, slightly upturned, bisinuate, median emargination shallow, broadly v-shaped,clypeogenal junction almost straight. Clypeus strongly and transversely wrinkled on anterior third in males or anterior two-thirds in females,remainder finely punctate in males or strongly punctate in females.Clypeofrontal suture ill-defined, withbackwardly curved horn,bifurcate apically, rather transverse in holotype, compressed laterally in male paratype, or with 2 well-separated acute tubercles medially in females. Anterior genal edge subequal in length to lateral,genal surface wrinkled on anterior half,posterior half finely punctate in males or with punctures almost confluent in females. Vertex finely punctate on anterior half,smooth and convex on posterior half,internal edges of eyes deeply foveolate in males. Pronotum
(Figs. 49,50). Transverse,W IL = 1.7; anterior edge slightly emarginate and membranous behind eyes,margin almost vertical in lateral view on short distance internal to eyes in males or margin slightly sinuous and narrowly membranous behind head insertion in females, anterior sulcus narrow,strongly microsculptured and clearly delimited posteriorly. Disc with 4 swellings,laterals large,close to anterior edge,preceded by shallow concavity anterolat erally, median swelling small and located posteriorly, posterior edge of declivity poorly defined in males or with pair of closely set transverse swellings on anterior fourth,anterior A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 63 declivity as wide as head insertion in females, punctures fine and evenly distributed throughout, slightly larger on anterior angles in females. Elytra. Subquadrate, L/W = 1. 0. Striae fine, moderately impressed on disc, slightly deeper on apical declivity, punctures rounded and umbilicate, slightly deeper than stria, approximately 5 times as wide as stria, separated by 1-2 diameters on disc and less than their diameter on posteriormost portion of apical declivity. Intervals moderately convex, surface alutaceous, finely and evenly punctate throughout. Thoracic sterna. Proepisternum with posterolateral portion moderately pubes cent and alutaceous, external fourth smooth. Prosternum behind procoxa moderately pubes cent, several long setae along posterior edge. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, alutaceous and strongly carinate. Metasternum with disc finely and scarcely punctate medially, convex in both sexes; lateral lobes with punctures oval, almost evenly spaced on anterior half, rounded, restricted to metacoxal edge and leaving scarcely punctate area on disc on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weaken ing posteriorly, pubescence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.3, lateral margins convergent on posterior two-thirds, divergent on anterior third, surface finely and densely punctate, punctures larger anteriorly, strongly alutaceous anteriorly, less so posteriorly, junction with mesosternum slightly arcuate, finely carinate laterally and declivous medially. Legs. Anterior tibia tridentate in males, quadridentate in females, ventromedial carina lacking intervening setae on basal third, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites2-5 with few coarse setiferous punctures laterally and row of small entire to confluentpunctures along anterior edge, surface alutaceous throughout, sternite 6 finely and densely punctate, surface alutaceous, suture between sternites 5-6 with row of small irregular punctures laterally.
Pygidium, WIL = 2. 1 in males and females, punctures fine and dense, surface slightly alutaceous, apical margin clearly delimited in both sexes. Male genitalia (Figs. 157-159). Tubus approximately 2.3 times as long as parameres, apicoventral portion flattened. Para meres in lateral view with apex obliquely truncate, ventral portion slightly concave, dorsal portion concave, apicodorsal angle strongly produced into small denticle. Sigmoid sclerite of internal sac as in Figure 281.
MATERIAL STUDIED. 3 ct ct, 1 9 (NMPC, SMTD).
Holotype ct (SMTD): Coroico, Bolivia/ W. H. Muche, Radeberg, Ankauf/ Saatl. Museum fiir, Tierkunde Dresden/ HOLOTYPE, Ontherus monilistriatus, F. Genier, 1992. Dissected.
Allotype 9 (NMPC): Coroico, Bolivia/ ALLOTYPE, Ontherus , monilistriatus, F. Genier, 1992.
Paratypes: BOLIVIA: La Paz: Chaco, Garlepp (1 ct SMTD); Same data as allotype ( 1 ct NMPC).
24. Ontherus (Caelontherus) pseudodidymus sp.nov. (Figs. 51-55, 160-162, 282; Map 11)
Ontherus didymus Erichson: sensu Bates 1887: 50 (redescription, distribution and comment) Ontherus didymus Erichson: sensu Luederwaldt, 1931: 394, 398 (key) Ontherus didymus Erichson: sensu Blackwelder, 1944: 206 (list) Ontherus didymus Erichson: sensu Howden and Young, 1981: 121-122 (key, redescription, distribution and biology)
ETYMOLOGY. Pseudodidymus, (false+ didymus) Greek adjective, refers to the fact that specimens of this species were previously misidentified as 0. didymus Erichson. 64 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
DESCRIPTION. Colour. Body uniformly reddish brown to black, usually with strong iridescent tinge on elytra. Pubescence fulvous. Length. 10-16 mm. Head (Figs. 51-55). Anterior edge arcuate, flat to slightly upturned, bisinuate, median emargination broad and shallow in males or broadly v-shaped in females, clypeogenal junction almost straight. Clypeus strongly and transversely wrinkled on anterior half in males or anterior two-thirds in females, remainder moderately punctate in males or strongly punctate in females. Clypeofrontal suture ill-defined, with backwardly curved horn, acuminate (Fig. 53) or bifurcate apically (Fig. 54), sometimes larger males with transverse lamina (Fig. 55), almost parallel laterally and broadly notched apically or with 2 well-separated acute tubercles medially in females, cephalic development reduced to 2 small median tubercles in smaller individuals. Anterior genal edge subequal to lateral, genal surface wrinkled on anterior half, posterior half finely punctate in males or with punctures almost confluentin females. Vertex moderately punctate on anterior half in males or punctures strong and confluent in females, smooth and convex on posterior half, internal edges of eyes deeply foveolatein larger males.
Pronotum (Figs. 51, 52). Transverse, W/L = 1.6; anterior edge slightly emarginate and membranous behind eyes in males or margin slightly sinuous and narrowly membranous behind head insertion in females, margin almost vertical in lateral view on short distance internally to eyes in larger individuals, anterior sulcus narrow, strongly microsculptured and clearly delimited posteriorly. Disc with 4 tubercles, laterals large, sublobate and obliquely oriented, close to anterior edge, preceded by concavity anterolaterally, median tubercles small and distinct, located in line with posterior edge of lateral tubercles, posterior edge of declivity not clearly bordered by arcuate bulge in males or with pair of closely set and low tubercles on anterior two-fifths, anterior declivity as wide as head insertion in females, development of smaller individuals reduced to pair of vaguely delimited swellings on ante rior fifth; punctures moderate in size, almost evenly distributed throughout, except anterior declivity almost smooth in larger males, punctures distinctly larger on anterior angles.
Elytra. Subquadrate, L/W = 0.9. Striae fine, moderately impressed on disc, deeper on apical declivity, punctures subcircular, broadly angulate laterally, umbilicate, as deep as stria, approximately 3 times as wide as stria, separated by 1- 2 diameters on disc and less than their diameter on apical declivity. Intervals convex, surface finely and evenly punctate throughout, slightly to moderately alutaceous. Thoracic sterna. Proepisternum with posterolateral portion pubescent and alutaceous throughout. Prosternum behind procoxa moderately pubescent, several long setae along posterior portion of disc. Mesosternum impressed posteriorly, punctures coarse and dense, except small area along edge of mesocoxa, pubescence fine and short, except few longer setae along mesocoxa. Mesepisternum irregularly punctate, alutaceous and strongly carinate. Metasternum with disc finely and densely punctate, except on midline posteriorly finely and scarcely punctate, convex in both sexes; lateral lobes with punctures oval, each puncture with anterior edge finely carinate, almost evenly spaced on anterior half, rounded and restricted to metacoxal edge leaving scarcely punctate area on disc on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly, pubescence moderately long and obliquely oriented; median lobe rather narrow, L/mW = 3.0, lateral margins convergent on posterior three-fourths, parallel to slightly divergent on anterior fourth, surface finely and densely punctate, punctures only slightly larger anteriorly, moderately alutaceous anteriorly, less so posteriorly, with at most indistinct longitudinal carina medially on anterior third, junction with mesosternum slightly arcuate, finely carinate laterally, sometimes completely carinate, lateral margins much wider than those on mesosternum. Legs. Anterior tibia tridentate in males, quadri dentate in females, ventromedial carina lacking intervening setae on basal third in males or basal fourth in females, apical spur inwardly bent on apical fourth in both sexes. Posterior legs unmodified. Abdomen. Sternites 2-5 with few coarse setiferouspunctures laterally and A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 65 row of small,entire to confluent punctures along anterior edge,surface alutaceous through out, stemite 6 finely and densely punctate, surface glossy, suture between stemites 5-6 slightly impressed and with row of small irregular punctures laterally. Pygidium, W /L = 2.3 in males or 2.6 in females, punctures fine and dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 160-162). Tubus approximately 2.0 times as long as parameres,apicoventral portion usually flat on apical half. Parameres in lateral view with apex obliquely truncate, ventral portion slightly concave, dorsal portion concave, apicodorsal angle produced into small and acute denticle. Sigmoid sclerite of internal sac as in Figure 282.
MATERIAL STUDIED. 388 a a, 3739 9 (AMNH, AVEC, BDGC, CMNC, CNCI, DEIC, FGIC, HAHC, INBC, LEMC, MCZC, MNHN, SEMC, TAMU, UNSM, USNM, WBWC, WDEC,ZMHB).
Holotype O' (CMNC): COSTA RICA: Punt., Monteverde, 1200-l 700m., 2-6.XII.1987, Genier and Bertrand/ Collection,Franc;ois Genier/ HOLOTYPE. Ontherus pseudodidy mus,F. Genier, 1992.
Allotype 9 (CMNC): Same data as holotype.
Paratypes: No locality (1 a SMFD); Maces*, Ronaima (1 a SMTD). COSTA RICA: No locality (20'O', 3 9 9 MNHN),same as beforeexcept: Rogers (5 O'O', 4 9 9 MNHN) (1 a CNCI), Hoffm. (1 a ZMHB), Schausf (19 MNHN); Coliblanco*, l 950m, W. R. Maxon (1O', 19 USNM); Piedra Negra*, Schild and Burgdorf (1 O' USNM); R. Susio*,H. Rogers (19 AMNH) (1 9 CNCI) (20' a, 19 MNHN). Alajuela: Orotina, 7.XII.1929, A. Alfaro (19 USNM); Penas Blancas, 1420m, J. Ashe, R. Brooks and R. Leschen, F.I.T., #274 (20'O' SEMC). Cartago: Irazu (19 USNM), same as before except: H. Rogers (1 a AMNH) (19 ISNB) (20' a MNHN) (29 9 USNM) (30'O' ZMHB), 6-7000ft. (1830-2130m), H. Rogers (1 O' CNCI) (19 ISNB) (29 9 MCZC) (5 ctO', 29 9 MNHN); Turialba (1 O' NMPC), same as before except: Heyne V. (1O',19 ZMHB). Guanacaste: Estac[ion]. Cacao,S. W. Side Volcan Cacao (1000-1400m), XI-XII.1989,R. Blanco and C. Chavez,URCG, 323300, 375700 (560'O', 64 9 9 INBC). Limon: Hamburg Farm, Santa Clara Pr., 1525' (470m), 30.V.1925 (19 USNM); Pacay as, C. Werckele (1O' AMNH). Puntarenas: Monteverde, 1400m, 23-28.II.[19]80, R.S. Anderson,10 cup traps (60'O', 3 9 9 HAHC),same as before except: 24-28.II.[19]80, R.S. Anderson, mega dung trap (1O',39 9 HAHC), 4500' (1370m), 21-26.V.1979, J.M. and B.A. Campbell (1 O' CNCI),9.V.1989, J. Ashe, R. Brooks and R. Leschen,F.I.T. #068 (1 Q SEMC), Campbell's Woods, 9.V.1989, 1520m, J. Ashe, R. Brooks and R. Leschen, F.I.T. #073 (19 SEMC), 9.V.1989,1570m, J. Ashe, R. Brooks and R. Leschen,pitfall trap, #076 (1 ct SEMC), Campbell's Woods, 14.V.1989, 1450m, J. Ashe, R. Brooks and R. Leschen, F.I.T., #177 (4 O'O', 59 9 SEMC), 15.V.1989, J. Ashe, R. Brooks and R. Leschen, F.I.T., #201 (3 O' O' SEMC), 1520m, 21.V.1989, J. Ashe, R. Brooks and R. Leschen,F.I.T., #316 (1O' SEMC),1520m,21.V .1989,J. Ashe,R. Brooks andR. Leschen, pitfall trap,#318 (1 O' SEMC), 21.V.1989, J. Ashe,R. Brooks and R. Leschen, F.I.T.,#321 (1 Q SEMC),Boehme House, 21. V.1989, J. Ashe, R. Brooks and R. Leschen, pitfall trap, #322 (1 Q SEMC), 1520m, 24.V.1989, J. Ashe, R. Brooks and R. Leschen, F.I.T., #433 (1O' SEMC),1550m,23.V .1989,J. Ashe,R. Brooks and R. Leschen, F.I.T., #435 (1 Q SEMC), Boehm House, 24.V.1989, J. Ashe,R. Brooks and R. Leschen, pitfall trap, #437 (1 O',3 9 Q SEMC), 1520m, 24.V.1989, J. Ashe, R. Brooks and R. Leschen, pitfall trap, #453 (7O' O', 5 Q 9 SEMC), 25.V.1989, J. Ashe, R. Brooks and R. Leschen, black light, #480 (19 SEMC), 1630m, 8.VII.1989, S. E. Roberts, F.I.T. (1 a, 19 SEMC), 14-16.VII.1989,R. E. Beer,F.I.T. (2 a a, 2 Q Q SEMC),1630m, 6.VII.1990, S. E. Roberts, F.I.T. (1 O' SEMC),Reserva de Monteverde,1500m, l-4.VI.1979,J. M. and B. A. Campbell 66 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
(7CJ' CJ', 5 9 9 CNCI), 24.V.1977, A. Forsyth (1CJ' WBWC),V.1979, A. Forsyth (60'CJ' WBWC),20.V.1979, A. Forsyth (7 CJ'CJ', 7 9 9 WBWC),20.V.1979, A. Forsyth,ex. human dung trap (5CJ' CJ', 3 9 9 WBWC), 20.V.1979,A. Forsyth,ex. human dung trap,D + N #2 trapo (40' CJ',1 9 WBWC), 23.V.1979,A. Forsyth (3CJ' CJ', 3 9 9 WBWC), 24.V.1979,a. Forsyth,ex. human dung trap (2CJ' CJ', 1 9 WBWC),1400m, 20.ll.[19]g3,B. Gill (3CJ' CJ', 7 9 9 CMNC), 13-16.Vll.19g3, 1500m,B. Gill (1 CJ',1 9 BDGC) (1 CJ' CMNC),1400m, 22.V.1979,H. and A. Howden (1 CJ' HAHC), 1400m,23.V.1979, H. and A. Howden,dung 6 cups trps 24hrs. (2 CJ' CJ' HAHC), 1400m, 29.V.1979,H. and A. Howden (1 CJ',2 9 9 HAHC), 1400m, 3.VI.1979, H. and A. Howden (1CJ' HAHC), 1400m, 4.VI.1979, H. and A. Howden,dung cup trap (3CJ' CJ' HAHC),1400m., 4.VI.1979, H. and A. Howden,Hoge finca,dung 5 cup trap (1CJ' H.AHC), 1400m, 12-14.VIII.19g7, H. and A. Howden (19 HAHC), 1400m, 15-17.VIII.19g7, H. and A. Howden, FIT (19 HAHC), 1400m, 21- 24.VIII.19g7,H. and A. Howden (20' CJ',2 9 9 HAHC),27.II.19g3, D. H. Lindeman,FIT (19 HAHC), 15.VI.19g3, D. H. Lindeman (1 CJ' HAHC), 1520m 15-23.Vll.9g3, D. H. Lindeman, FIT (1 CJ' HAHC), 1520m, l 1-1g.v 1.19g3, D. H. Lindeman, F.l.T. (20'CJ' BDGC), 20.ll.19gg, D. B. Thomas,F. D. Parker (1 CJ' HAHC), 26.V-3.VI.[19]g4, E. Riley, D. Rider and D. LeDoux (90'CJ', 10 9 9 TAMU), 11-13.III.1976, I. C. Reed (19 TAMU),Monteverde Reserve,1500m, 27.V.1979, H. and A. Howden,dung cup trap 3 day (29 9 HAHC),Monteverde reserve, l 500m,1. VI.1979,H. and A. Howden,mega dung trap (19 HAHC), Monteverde reserve, 1500m, 15.VIII.19g7, H. and A. Howden, night (20'CJ' HAHC), Sendero Chomogo (Monteverde reserve), 16gOm, 19.V11.19g3, D. H. Lindeman,6hr. dung traps (7CJ' CJ', 10 9 9 HAHC),Sendero Chomogo (Monteverde Reserve),16gom, 24.V11.19g3,D. H. Lindeman,6hr. dung traps (gryCJ', 5 9 9 HAHC), Sendero Chomogo (Monteverde Reserve), 16gom, 25.V11.19g3, D. H. Lindeman, 6hr. dung traps (3 CJ'CJ', 2 9 9 HAHC); Puntarenas, 1959, C. W. Palmer (19 AMNH); S[an] Vito, Las Cruces, 1200m, 15-1g.vm.19g2,B. Gill (5 CJ'CJ', 5 9 9 BDGC) (19 CMNC), same as before except: 12.IX.19g2 (1 CJ' BDGC); 6km S. San Vito, 27.IV-7.V.1967 (1 CJ', 1 9 WDEC); San Jose: 30km N. San Isidro (1700m),19.11.19g3, B. Gill (1CJ',19 BDGC); San Jose (1CJ' ISNB),same as before except: V.[19]21 (1CJ' USNM), V.[19]21, cow dung (1CJ' USNM), VIIl.[19]21, cow dung (1CJ' USNM), 1000-1200m,25.X.[19]25, F. Never mann (1CJ'USNM),1000-1200m, l.IX.[19]2g,F. Nevermann (1CJ'USNM),1000-1200m, X.[19]2g, F. Nevermann (1 9 USNM), 1000-1200m, X.[19]30, F. Nevermann (20' CJ' USNM), 1000-1200m, 24.XI.[19]32, F. Nevermann (19 USNM), 1000-1200m, 17.V.[19]34, F. Nevermann (1 CJ' USNM), 1000-1200m, 13.Vll.[19]34, F. Nevermann (1CJ' USNM), 1000-1200m, 15.Vll.[19]37,F. Nevermann (1CJ' USNM), C. Werckle S. G. (19 ZMHB),Henry Schmidt leg.,vend. 31.VllI.1911 (20'CJ' MNHN). PANAMA: Chiri qui: No locality (1 CJ' MNHN), same as before except: 2.X.1975, 1500hrs, 0. P. Young, opossum faeces (1CJ' HAHC), same as before except: 2-3.X.1975, tapir faeces trap (19 HAHC); Boquete (1 CJ',1 9 ZMHB), same as before except: 12.1.1940, G. C. Wood (19 WDEC); 2km E. Cerro punta, 2200m, 2g.v.-g.VI.[19]77, S. Peck, dung tps 26-29 (1CJ' CNCI) (1CJ', 29 9 HAHC); 2km W. Cerro Punta, 1720m, go 51' N g1° 36' W, 19- 23.V.[19]77,H. and A. Howden, light (1 CJ' CNCI) (60'CJ', 11 9 9 HAHC) (1 9 MCZC), same as before except: go 51' N g2° 36' W,2g_V. l977,cup trap (1CJ',2 9 9 BDGC),1300m, go 51' N g2° 36' W, 29.V.[19]77,cup trap day 1 (7CJ' CJ', 15 9 9 HAHC), 30.V.[19]77,day 2 (20'CJ' 29 9 HAHC), go 51' N g2° 36' W,15-30.V.[19]77 (1 CJ' MCZC), go 51' N g2° ' 36' W, 1-7.Vl.[19]77, cup traps (20'CJ' HAHC), 5750' (1750m), 9.1.1964, L. J. Bottimer, human dung (20'CJ', 2 9 9 CNCI), Baldwin Forest, l 760m, 30.V-g.Vl.[19]77, S. Peck, dung trap 19 (30'CJ', 2 9 9 FGIC) (60'CJ', 7 9 9 HAHC) (240'CJ', lg9 9 LEMC), l 760m, Baldwin forest, 30.V-g.VI.[19]77, S. Peck, carrion tps 20 (19 HAHC),Baldwin Forest, 3.VI.[19]77, H. Howden, 10 cup traps (20'CJ' HAHC); 2km W. Cerro Punta, Audubon For[est]., 1700m, 24-2g.V.[19]77,S. and J. Peck,Ber 375, under carrion (1CJ' A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 67
HAHC); 15km N. E. Gualaca, Cerro Hornito, 1200m, 17-19.VI.1982, B. Gill (30"0", 39 9 BDGC) (1 O" CMNC), same as before except:VIII.1982 (20"O", 1 99 BDGC); 5km S. W. Hato del Vo lcan, Lagunas, 1360m, 22-26.V.[19]77, S. Peck, dung traps (5O" O", 29 9 FGIC) (1 O", 4 99 HAHC) (35 O"O", 459 9 LEMC), same as before except: 27.V.[19]77, S. and J. Peck, ber. 373, u[nder] carrion (2 99 HAHC); 4km W. Hato del Vo lcan,Las Lagunas, 1360m, 26-27.V.[19]77,H. and A. Howden(11O'O",15 9 9 HAHC), same as before except: l.VI.[19]77 (20"O" HAHC); Lino (1 O' ZMHB); Santa Clara, Dist. Renacimiento, 4000' (1220m), 22-25.V.1982, B. C. Ratcliffe and C. and K. Messenger (19 UNSM); Santa Clara, 23-25.V.1980,E. Riley and LeDoux (3 O" O" TAMU); St. Clara, Hartmann's Finca, 15-18.VI.[19]85, Riley and Rider (40"O', 1 9 TAMU); 2km N. Sta. Clara, Hartmann's Finca, 1300m, 8° 51' N 82° 46' W, 24-25.V.[19]77,H. and A. Howden (1O", 1 9 BDGC) (8 O"O", 2 99 CMNC) (20"O" HAHC), same as before except: 20.V.[19]77,cup traps(40" O', 9 99 HAHC); 4km N. Sta. Clara,Hartmann's Finca,1500m, 27.VI-3.VII.1981, B. Gill (19 AV EC) (60"O", 49 9 BDGC) (19 HAHC) (20"O", 59 9 CMNC); 15km N. W. H[ato] del Vo lcan, Hartmann finca,1500m, 20-31.V.[19]77, S. Peck, dung tps 18 (40"O", 5 99 HAHC),same as before except: 20-25.V.[19]77,carrion tp 3 (1O", 1 9 MCZC), 1550m, 31.V.[19]77,dung tps 17 (60"O", 5 99 HAHC), 1550m, carrion tp 3+16(10",19 CNCI),1550m,dung traps 4-6(60"0",29 9 HAHC);La Fortuna Dam, 1200m, 19-21.VI.1982, B. Gill (90"O", 15 99 CMNC), same as before except: 15-17.VI.1982, trap megadung (40"O", 19 BDGC) (1O" HAHC), lOOO m, 5-6.VII.1981 (1 O" AV EC) (16 O"O", 169 9 BDGC) (1 O", 1 9 HAHC); [La] Fortuna (Dam), 13 15.VI.1985, E. G. Riley and D. Rider (1 O" TAMU); V[olcan] de Chiriqui, 25-4000' (760-1220m), Champion (1 9 MNHN); Code: Cerro Gaital, 10-12.VI.1985, E. Riley and D. Rider (1 9TAMU). Darien: Pirri Range, 12-30.IV.[l 9] 12, A. E. Goldman (1 9USNM).
Additional data: Specimens were collected using light traps,flight intercept traps and pitfall traps baited with carrion, dung and human feces. Some specimens were collected using cup traps baited with human feces and" megadung" traps. Finally,specimens were also collected from opossum and tapir feces.
SUBGENUS PLANONTHERUSNOV.
Type species: Ontherus bridgesi Waterhouse
ETYMOLOGY. Planontherus, (flat + Ontherus) Latin adjective, refers to the flattened body aspect.
DESCRIPTION. Body elongate and flattened dorsoventrally. Head tuberculate in both sexes. Pronotum transverse, declivous anteriorly behind head insertion in both sexes. Elytra depressed on disc; striae fine,eighth stria interrupted on basal third and distinct from ninth; intervals usually strongly convex. Mesosternum with short carina posteriorly. Mesepister num flat lacking carina parallel to posterior edge. Median lobe of metasternum arcuate to angulate and raised anteriorly. Anterior tibia with either tooth reduced to 5 lobes, 4 teeth with penultimate and apical widely separated, or 5 teeth in well-developed males. Posterior femur with posteroventral margin lobate apically. Parameres in lateral view triangular.
25. Ontherus (Planontherus) bridgesi Waterhouse, 1891 comb.nov. (Figs. 56 - 58, 163-165, 283; Map 12)
Ontherus bridgesi Waterhouse, 189 l: 358 (description) Ontherus bridgesi Waterhouse: Gillet, 19 l lb: 57 (catalog) Ontherus bridgesi Waterhouse: Luederwaldt, 1931: 393 (key) Ontherus bridgesi Waterhouse: Blackwelder, 1944: 206 (list) 68 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
REDESCRIPTION. Colour. Body dark reddish brown to black, pubescence fulvous. Length. 11-16 mm. Head. Anterior edge broadly arcuate, almost flat, medianfourth slight! y emarginate in males or with 2 widely separated and barely indicated teeth in females, clypeogenal junction finely notched. Clypeus finely wrinkled on anterior third, strongly wrinkled on anterior half for the Argentinean population, remainder finely punctate in males or wrinkled throughout in females. Clypeofrontal suture broadly arcuate, with small conical process medially, slightly more acute in larger males. Anterior genal edge much longer than lateral, angle rather blunt, genal surface almost smooth in larger males, finely wrinkled on anterior half in smaller males and females, strongly wrinkled on anterior half in both sexes for the Argentinean population. Vertex finely punctate throughout. Pronotum. Transverse,
W IL = 1.6; anterior edge narrowly membranous and slightly lobed medially behind head insertion; posterior angles slightly depressed; disc finely punctate throughout. Anterior sixth slightly declivous behind head insertion. Elytra. Elongate, L/W = 1.2. Striae fine, alutaceous, moderately impressed on disc and lateral declivities, much deeper on apical declivity, punctures as deep as stria, broadly oval to rounded and umbilicate, approximately 3 times as wide as stria, separated by 3-6 diameters on disc. Elytral intervals variable, flat to strongly convex, surface glossy to strongly alutaceous with fine punctures. Thoracic sterna. Proepistemum with posterolateral portion with few setiferous punctures. Prostemum behind procoxa with few setiferous punctures on basal third. Mesosternum impressed and with short longitudinal carina posteriorly, punctation coarse and dense throughout, pubes cence fine and short. Mesepisternum moderately punctate, punctures each with lateral edge ill-defined, surface flat and strongly alutaceous. Metasternum with disc finely punctate, deeply foveolate in males or almost convex in females; lateral lobes scarcely punctate, punctures on anterior half quickly becoming fine to absent toward posterior half, surface varying from completely glossy to alutaceous, pubescence short and obliquely oriented; median lobe narrow, L/mW = 2.8, lateral margins straight and convergent throughout, surface slightly alutaceous and scarcely punctate, punctures on anterior half equidistant and each with erect seta, junction with mesosternum arcuate to distinctly angulate, finely carinate throughout, elevated medially. Legs. Anterior tibia quadridentate (Argentinean form) (Fig. 56), quadridentate with accessory lobe between penultimate and apical tooth (Bolivian form) (Fig. 57) or quinquelobate (Peruvian form) (Fig. 58) in males or quadridentate in females, ventromedial carina lacking intervening setae on basal four-fifth in males or on basal half in females, apical spur inwardly bent on apical fourth. Posterior femur widening toward apex, ventroposterior edge spatulate apically, dorsoposterior edge lobate apically, posterior tibia slightly to moderately compressed on basal third in larger males or unmodified in females. Abdomen. Sternites 2-5 with anterolateral portion impressed and with row of finely setiferous punctures, surface slightly to moderately alutaceous, sternite 6 scarcely and finelypunctate, median portion produced into bulge in larger males, suture between sternites 5-6 deeply furrowed throughout, moderately furrowedfor the Argentinean population, with row of deep foveolate punctures laterally. Pygidium, W/L = 1.9 in males or 2.1 in females, punctures fine and evenly distributed, apical margin clearly delimited throughout its length in both sexes. Male genitalia (Figs. 163-165). Tubus approximately 2.2 times as long as parameres, ventral portion depressed. Parameres in lateral view triangular, junction with tubus sinuate ventrally. Sigmoid sclerite of internal sac as in Figure 283.
MATERIAL STUDIED. 85 et et, 64 9 9 (BMNH, CMNC, CNCI, HAHC, ICCM, ISNB, MNHN, PAIC, SMTD, USNM, ZSMC).
TYPE MATERIAL Holotype et (BMNH): Type/ Bolivia (recto), 48., 75 (verso)/ Ontherus bridgesi, et (type) Waterh./ Holotype, Ontherus bridgesi, Waterhouse, 1891, vidit F. Genier, 1991. Holotype studied. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 69
• 0. bridgesi
* 0. rectus
MAP 12. Known distribution of Ontherus hridgesi and 0. rectus.
Allotype Q (BMNH): Peru (recto), 73. 33. (verso)/ Type/ Allotype, Ontherus bridgesi, Waterhouse, 1891, vidit F. Genier, 1991. Allotype studied.
NON-TYPE MATERIAL Distribution: ARGENTINA: Salta: 20km N La Caldera (El Ucumar) (780m); 22km N La Caldera (El Ucumar) (1550m). BOLIVIA: Cochabamba: Carrasco (=Provincia de); Cochabamba; Yungas del Palmar (2000m); Yungas de Totora-Schuencas (21 OOm); Yungas Incachaca (21OOm). La Paz: Chaco; Coroico; Yungas de Arepucho (Sihuencas) (2200- 2500m); Yungas de Corani (2500m). COLOMBIA (doubtful): No locality. PERU: Cusco: Marcapata; Pillahuata. VENEZULA (doubtful): No locality.
Additional data: Specimens were collected using a malaise-flightintercept trap in a subtropical humid forest. Specimens collected in February to April, September, October, December.
REMARKS. Under this specific name I am including what appear to be three distinct populations (Peruvian, Bolivian and Argentinean). They have identical male genitalia. They differ mostly by the male secondary sexual characters, especially the anterior tibial teeth (Figs. 56-58), and degree of coarseness of the punctures and microsculpture. The Colombian (1 Q ISNB) and Venezuelan (1 Q MNHN) records are most likely mislabelled specimens. In both cases the pronotum is more coarsely and densely punctate, as in the females of the Argentinean population.
26. Ontherus (Planontherus) rectus sp.nov. (Map 12)
ETYMOLOGY. Rectus, (straight) Latin adjective, refers to the aspect of elytral striae.
DESCRIPTION (based on 1 female). Colour. Body dark reddish brown. Pubescence fulvous. Length. 18 mm. Head. Anterior edge broadly arcuate, almost flat, with 2 widely 70 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
separatedteeth, clypeogenal junction finely notched. Clypeus strongly wrinkled throughout. Cly peofrontal suture broadly arcuate, with small and acute conical process behind suture medially. Anterior genal edge much longer than lateral, angle rather blunt, genal surface finely wrinkled on anterior half. Vertex finely and weakly punctate on anterior half.
Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly lobed medially behind head insertion; posterior angles slightly depressed; punctures fine and restricted to anterior angles. Anterior eighth slightly declivous behind head insertion, slightly
more so on median third. Elytra. Elongate, L/W = 1.2; striae fine, finely alutaceous, shallowly impressed throughout their length, punctures as deep as stria, broadly oval to rounded and finely umbilicate, only slightly wider than stria, separated by approximately their diameters on disc. Elytral intervals convex, surface glossy with few minute punctures. Thoracic sterna. Proepistemum with posterolateral portion almost smooth. Prostemum behind procoxa with few setiferous punctures on basal third. Mesostemum moderately impressed and with short longitudinal carina posteriorly, punctation coarse and dense throughout, pubescence fine and short, slightly thicker and longer laterally. Mesepistemum moderately punctate, punctures each with lateral edge ill-defined, surfaceflat and alutaceous. Metastemum with few fine punctures posteriorly, remainder smooth, almost convex; lateral lobes scarcely punctate, punctures on anterior half quickly becoming fine to feeble toward posterior half, surface minutely punctate throughout and completely glossy, pubescence
short and obliquely oriented; median lobe wide, L/mW = 2.4, lateral margins almost straight, parallel on anterior half, surface alutaceous, densely minutely punctate throughout, com pletely bare, junction with mesostemum angulate and carinate throughout, elevated medially. Legs. Anterior tibia quadridentate, ventromedial carina lacking intervening setae on basal three-fourths, apical spur inwardly bent on apical fourth. Posterior legs unmodified. Abdo men. Stemites 2-5 with anterolateral portion moderately impressed and with row of finely setiferous punctures, surface finely alutaceous throughout, densely and minutely punctate laterally, stemite 6 densely and finely punctate, punctures slightly larger laterally, suture between stemites 5-6 slightly furrowed laterally, with few irregular punctures laterally.
Pygidium, W IL = 2.1, punctures fine basally, becoming scarce and minute apically, apical margin clearly delimited throughout its length in both sexes(?). Male. Unknown.
MATERIAL STUDIED. l Q (SMTD).
Holotype Q (SMTD): Huancabamba, N. Peru, 3000m., H. Rolle/ Coll. C. Felsche, Kauf 20, 1918/ Ontherus elongatus, Waterh.(hand written)/ Staatl. Museum fiir, Tierkunde Dres den/ HOLOTYPE, Ontherus rectus, F. Genier, 1992.
REMARKS. Ontherus rectus sp.nov. is described from a single female. The following characters, based on variation within the genus, are sufficient to justify this placement. It is distinct fromthe only other species in the subgenus by the shape, and the completely smooth median lobe of the metastemum. The differently configured anterior of the pronotum, combined with the presumed larger size, also distinguish this allopatric species.
SUBGENUS ONTHERUS S.STR.
Type species: Ontherus sulcator (Fabricius)
DESCRIPTION. Body usually oval, sides usually broadly arcuate, dorsum variably convex. Head convex or with median tubercle, simple clypeofrontal carina, fine clypeofrontal carina tuberculate medially, or bi- or trituberculate clypeofrontal carina. Pronotum convex, usually with distinct, longitudinal median sulcus basally; some species with anterior portion of disc declivous and transversely bulbous behind declivity. Elytra convex; striae variously shaped, eighth stria usually distinguishable from ninth on apical two-thirds; intervals usually finely punctate. Mesostemum with or without short longitudinal carina posteriorly. Mesepistemum A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 71 always carinate parallel to posterior edge. Median lobe of metastemum always angulate medially (except for 0. irinus Balthasar). Anterior tibia with 3-5 teeth in males, 4 in females. Posterior leg usually modified in males. Parameres variable in shape, subtriangular to elongate.
SULCATOR GROUP
DIAGNOSIS. Species included in this group separate from all others in genus by following combination of characters: head tuberculate; median lobe of metastemum wide (L/mW = 1.2); mesometasternal suture angulate medially and pronotum convex to narrowly and slightly declivous behind head insertion. Small and laterally oriented apical tooth of anterior tibia in males unique in genus. Parameres simple, albeit with small emargination on basal margin ventrally.
27. Ontherus (Ontherus) pubens sp.nov. (Figs. 73, 166-168, 284; Map 13)
ETYMOLOGY. Pubens, (hairy) Latin adjective, refers to pilosity of posterior surface of posterior femur.
DESCRIPTION. Colour. Body reddish brown to black. Pubescence fulvous. Length. 9-17 mm. Head. Anterior edge arcuate, feebly to moderately upturned, bisinuate medially, clypeogenal junction straight. Clypeus transversely wrinkled on apical four-fifths in both sexes, remainder coarsely punctate, in some specimens finely granulate, clypeofrontal suture finely carinate, with small conical process medially in both sexes, process larger in females, smaller individuals with small swelling. Anterior genal edge much longer than lateral, genal surface transversely wrinkled on anterior half, posterior half granulate. Vertex coarsely granulopunctate, basal portion with smooth transverse area in both sexes. Pronotum. Transverse,W /L = 1.6; anterior edge narrowly membranous and almost straight behind head insertion. Disc slightly gibbous on anterior sixth in males or slightly declivous medially on anterior fourth, posterior portion transversely swollen in females, anterior portion convex in smaller individuals, basal third to half with shallow and weakly delimited median longitu dinal sulcus; punctures fine, rounded and scattered on disc and lateral swellings, larger, oval and denser along base, anterior and lateral declivities. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, as deep as stria, approximately 1.2 times as wide as stria, separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely and evenly punctate throughout, lacking a lutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion pubescent and finely alutaceous throughout. Prostemum behind procoxa moderately pubescent, setae longer medially along base. Mesosternum shallowly and transversely impressed on disc, punctures coarse and dense throughout, pubescence fine,few longer setae along mesocoxa. Mesepisternum irregularly punctate, surfacefinely alutaceous and strongly carinate parallel to posterior edge. Metasternum with disc finely punctate, with round depression medially in males or finely and longitudinally sulcate in females; lateral lobes with punctures oval, each with anteromedial edge raised and sharp, almost evenly distributed on anterior half, broadly oval to rounded and restricted to lateral portion, surface of inner portion almost smooth on posterior half, surface strongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly leaving glossy area anterior to posterior coxa, pubescence moderately long and obliquely oriented, restricted to anteromedial third; median lobe wide, L/mW = 1.2, lateral margins almost parallel on anterior half, convergent on posterior half, disc with strong arcuate impression in larger males or convex in females and smaller males, surface punctate and alutaceous throughout, punctures scarce and fineon anterior fifth, junction with mesosternum angulate and raised, usually produced into short 72 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. pubens
MAP 13. Known distribution of Ontherus pubens.
carina medially on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth reduced in males, ventromedial carina lacking intervening setae on basal two-fifths in males or basal fourth in females, apical spur straight and obliquely truncate, slightly spatulate apically in males. Posterior femur (Fig. 73) with anterior edge sharp, lacking any trace of sulcus ventrally, at most weakly sulcated ventrally on apical half in males; posterior surface conspicuously pubescent in both sexes (best seen in non-eroded speci mens), posterodorsal edge finely carinate throughout in males or unmodified in females. Abdomen. Sternites 2-5 with several punctures laterally, anterior edge depressed and with row of small irregular punctures, surface finely alutaceous, finely punctate medially, sternite 6 finely punctate throughout, surface glossy, suture between sternites 5-6 narrowly and deeply sulcate. Pygidium, WIL = 2.1 in males or 2.3 in females, punctures moderate in size and dense, in some specimens fine on apical third, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 166-168). Tubus approximately 2.0 times as long as parameres, ventral portion finely sulcate. Parameres in lateral view with apex rounded, ventral portion straight, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 284.
MATERIAL STUDIED. 354 a O", 285 Q Q (AMNH, BCRC, BDGC, BMNH, CASC, CMNC, CNCI, FGIC, HAHC, ICCM, IMLA, ISNB, LACM, MCZC, MLUH, MNHN, MZLU, NMPC, SEMC, SMFD, SMTD, USNM, WBWC, ZFMK, ZMHB, ZMUC, ZSMC)
Holotype a (CMNC): ECUADOR: NAPO, 400m, Jatun Sacha Biol. Station (21 km E. Puerto Napo), 11.VII.1994, F. Genier, 2nd grth. veg., fecestrap/HOLOTYPE, Ontherus pubens, F. Genier, 1994. A slightly teneral specimen was selected to show complete pilosity, aedeagus removed.
Allotype Q (CMNC): Same data as holotype. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 73
Paratypes: ARGENTINA: Cordoba: San Francisco,X.1950 (1 ct ZSMC). BOLIVIA: No locality (1ct ISNB) (4ctct, 1 9 ZMUC); Paradito*,22.IX.1949, L. Pena col. (1 ct CASC). Beni: Cosincho Regn, G. L. Harrington (2ctct, 2 9 9 USNM); Huachi, IX,W. M. Mann (2ctct, 6 9 9 USNM); Rio Mamore, IX.1956, L. Zischka (1 ct ZSMC); San Antonio de Mapiri* (1ct SMFD). La Paz: Asunta*,V.1925, G. L. Harrington (1ct USNM); Rio Bopi, V. 1925,L. G. Harrington (2ctct USNM). Santa Cruz: No locality (2ctct, 6 9 9 ZMHB); Buena Vista,XII.1948 (1 ct CASC),same as before except: IV.1950(1 ct CASC),Steinbach (4ctct ICCM); Region Chapare*, 400m, leg. Zischka (23ctct, 69 9 ZSMC), Umg. Buenavista, 450m, Steinbach coll. (2 9 9 MNHN) (3ct ct, 1 9 ZSMC), 1922, R. C. Robert (1 ct,2 9 9 USNM); Santa Cruz de la Sierra (5ct ct, 1 9 ZMHB), same as before except: 500m, I-IV.1904,J. Steinbach S. V.(5 ct ct, 2 9 9 ZMHB). BRAZIL: Amazonas: No locality (1ct ISNB)(1ct,59 9 MNHN) (1 ct SMTD) (19 ZMHB) (1ct ZMUC); Ega (=T efe), H. W. Bates (19 MNHN); Manacapuru, III.1926, S. M. Klages (1 ct ICCM); Manaus, 8.I.1977, B. C. Ratcliffe (1ct BCRC); Santa Rita (do Weil), Tabatinga-Gebiet, IX.1926, Ehrhardt (1 ct SMFD); Sao Paulo de Oliven�a, Hahnel (3ct ct MNHN), same as before except: II.1923, S. Klages (1 ct ICCM),V.1923, S. Klages (1ct ICCM); Serra dos Porcos, 0°25'N 69°22'W, 2-25.V.1975, Franklin Moore (1 ct BCRC); 2km. E. Tabatinga, l.II.1985, F. Genier (1 ct FGIC); Tefe, Raulin, Thayer Exped. (2ctct MCZC); Villanova, 1880, Hahnel (19 MNHN). Rondonia: Porto Ve llo (=Velho) (1 ct, 69 9 USNM), same as before except: Rio Madeira (3 ct ct USNM),Rio Madeira,Mann and Baker (4ctct, 4 9 9 USNM), Rio Madeira, IX (1 ct USNM); Rio Madeira, Pohl. (1ct, 1 9 USNM). COLOMBIA: No locality (1ct SMTD),same as before except: Frere Apollinaire Marie (1ct ISNB), VIII.1935, W. H. W. Kemp (1 ct USNM); Cagus* (1 ct MCZC); Sarare* (1ct,29 9 NMPC) (19 ZFMK). Amazonas: Leticia, 23.II-l.III.1974, B. C. Ratcliffe (1 et BCRC), same as before except: 700' (210m), 9.VII.1970, H. and A. Howden (1 ct HAHC), Pepper Farm, l.III.1974, Nealis (1 9 HAHC), Los Alpes, 700' (210m), 23.II-2.III.1974, S. Peck, T2D (2ctct HAHC), 700' (210m), 23.II-2.III.1974, Howden and Nealis (1ct,29 9 HAHC). Huila: Gigante, X.1980 (1 ct BCRC); Gigante and Vcty., III-IV.1982, 0. Rojas and Coll'rs (4etct WBWC). Meta: Villavicencio,1920, Frere Apollinaire Marie (6ct ct, 3 9 9 ISNB), same as before except: 28.II.-5.III.1972, S. and J. Peck,f orest,dung traps(2 9 9 HAHC),500m, 29.II.-3.III.1972, S. and J. Peck,f orest, bait traps (1ct, 1 9 HAHC); 16 km E. Villavicencio,2-4.III.1972, S. Peck,forest, dung traps (1 ct HAHC); 30 km E. Villavicencio,1-4.III.1972, S. and J. Peck,dung and forest (2ctct HAHC). ECUADOR: No locality (19 ISNB), same as before except: Baron (5ct ct, 12 9 9 CASC) (1ct, 2 9 9 ISNB) (1ct NMPC) (19 SMFD) (4ctct, 6 9 9 SMTD) (1 9 ZSMC), F. Deville (19 ISNB). Loja: Loja, Abbe Gaujon (11 ctct, 7 9 9 MNHN). Morona-Santiago: Macas (19 ZMHB). Napo: No locality, R. Haensch S. (1ct ZFMK) (2 9 9 ZMHB); Archidona, R. Haensch S. (1 9 NMPC) (1et, 39 9 ZMHB); Coca, R. Haensch S. (19 NMPC)(2 ct ct ZMHB),same as beforeexcept: Napo R.,250m, V.1965, L. Pefia(4ctct, 2 9 9 CNCI); Dureno(on Rio Aguarico)*,150m, 23-30.IX.1977, Luis Pefia (1 ct,1 9 HAHC); Jatun Sacha Biol. Station (21 km E. Puerto Napo),8.VII.1994, F. Genier, virgin rain forest, feces tp. (1ct FGIC), same as before except: 2nd grth. veg. (4ctct, 1 9 FGIC), 9.VII.1994 (8ctct, 79 9 FGIC), ll.VII.1 994, 2nd grth. veg. (4etct FGIC); Limoncocha,700' (210m), l 1.III.1976, J. M. Campbell (60ctct, 50 9 9 CNCI),same as before except: 10-15.III.1975 (1ct, 2 9 9 HAHC), 800' (240m), 15.II.1976 (3ct ct, 79 9 CNCI),250m, 18-24.VI.1976, S. Peck,3 dng. tps. (14-16), nature trail for. (6ctct, 59 9 CMNC) (17 ctct, 12 9 9 HAHC),250m, 21-28.VI.1976, S. Peck,carrion traps 20, forest (1 ct HAHC),13.VII.1977, P. J. Spangler and D. R. Givens,#120 (3ctct BCRC), 16.V I.1977, P. J. Spangler and D. R. Givens, #130 (1 ct,1 9 BCRC), VII.1979, T. Ray, human dung trap (4ctet, 6 9 9 MCZC); 2 km. N. Limoncocha, 22-28.VI.1976, S. Peck, dng. tps. 22-24, virgin forest (2 ctct, 2 9 9 CMNC) (8 ctct, 7 9 9 HAHC); 10 km W. 74 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Puerto Misahualli, 20.II.1986, F. Genier, cow dung (7 O'O', 7 9 9 FGIC); Reventador*, III.1983, leg. L. Huggert (2 9 9 MZLU); Rio Napo*, Pozzi (19 MNHN), same as before except: F. Oh[au]s S. (19 SMFD) (1 9 USNM); Km 7.3 Sarayacu-Loreto Rd, 1200m, 14.VII.1994, F. Genier, cloud forest, feces trap (1 9 FGIC); Tena, 400m, 15-21.II.1986, Franc;oisGenier, human feces trap(28 O'O', 19 9 9 FGIC), same as before except: IV.1923, F. X. Williams (19 SEMC), 14. VII.1994, riparian veg., feces trap (8 O' O', 49 9 FGIC); 5km W. Tena, 500m, 6-9.VI.1976, S. Peck, dng. tps. 28-29, 2nd growth scrub (5 O'O', 1 9 HAHC); 12 km W. S. W. Tena, 600m, 8-1 l.VII.1976, S. Peck, for. dung traps 30-33 (20'0', 29 9 HAHC), same as before except: 6-ll.VII.1976, for. dung traps 30-32 (20'O', 2 9 9 HAHC). Pastaza: Canelos, 23.XII.1905, F. Ohaus S.(1 O' USNM); Curaray, 20.I.1906, F. Ohaus S. (1 a SMFD) (1 a USNM); Llandia (17 km. N. Puyo), lOOOm. 16.VII.1994, F. Genier, remnant rain for., hand coll. (1 9 FGIC). Zamora-Chinchipe: Sabanilla, F. Ohaus S. (30' O', 29 9 SMFD); Zamora, 26,IX.1905, Arsen (10' NMPC). PERU: No locality (1 O', 1 9 BMNH) (19 MLUH). Amazonas: Collected on way to Chachapoyas as Rioja D. San Idelfonso, 7.IX.1936, F. Woytkowski, no. 3757 (1 9 SEMC); Rio Santiago, 29.X.1924 (19 AMNH). Cuzco: Quiroz, Rio Paucartambo, 8.XI.1933 (1 O' LACM), same as before except: 10.X.1933 (1 O' LACM), 14.XI.1933 (19 LACM). Huanuco: Tingo Maria, 2200' (670m), 28.V.1947, J. C. Pallister (1 O' AMNH). Junin: Chanchamayo (19 NMPC), same as before except: Ch. 0. Schunke (2 O'O', 1 9 MNHN); La Merced, Smyth (19 LACM), same as before except: I.1934, E. G. Smyth (2 9 9 LACM); Rio Oxabamba, La Merced, Chanchamayo (1 O' ZMHB); Va lle Chanchamayo, 800m, X.1944, leg. Weyrauch(1 9 IMLA). Loreto: Iquitos, IV.1938, J. M.(40' a USNM); Middle Rio Ucayali(1 O'AMNH); Pebas(2 O' O' MNHN), same as beforeexcept: Dr. Hahnel (1 O' MNHN), fin Xbre and ler Tr 1880 (1 O' MNHN). Madre de Dios: Madre de Dios, 29.IX.1975, J. Schunke L. (1 O' HAHC), same as before except: 500' (150m), 29.IX.1975, J. C. Pallister (1 O' AMNH); Manu National Park, 15-30.VIII.1986, A. Forsyth (3 O'O', 2 9 9 BDGC)(60'a, 5 9 9 HAHC). Pasco: Cacazu, lOOOm, 23.IX.1985, Pedro Hocking (1 O' HAHC); Villa America*, 300m, 8.X.1984, Pedro Hocking(1 O' HAHC). San Martin: Moyobamba, 1800' (550m), 20.XII.1946, J. C. Pallister (20'O', 49 9 AMNH); Rioja, ex Stgr (1 O', 1 9 MNHN); Vicinity of Rioja, 21.XI.1936, F. Woytkowski, no. 3759 (19 SEMC); Soritor (21 km W. Rioja), 12.X.1936, F. Woytkowski, no 3758 (19 SEMC); 8 km E. N. E. Tarapoto, 550m, 9.I.1974, R. T. Schuh, ex cow dung, roadside (1 O' HAHC). Ucayali: Pucallpa, 200m, 26.X.1960, J. Schunke (19 HAHC), same as before except: 5.XI.1960 (2 9 9 HAHC), 7.VIII.1968 (1 a HAHC), 4.IX.1968 (20'a, 1 9 HAHC), 5.IX.1968 (19 HAHC), 18.XII.1968 (19 HAHC), 9.I.1969 (19 HAHC), 26.IV.1974 (1 a HAHC), 24.VI.1974 (19 HAHC), 30.VI.1974 (1 a HAHC); Santa Ana (Rio Urubamba), llOOm, 7.XII.1947, col. W Weyrauch (1 a IMLA). VENEZUELA: Apure: No locality (5 O' O', 8 9 9 NMPC) (1 9 ZFMK), same as before except: F. Geay, 33-96 (3 a O', 4 9 9 MNHN). Tachira: San Cristobal, 2.VII.1982, D. Havranek (1 9 BCRC); 20 km N. E. San Cristobal(1200m), 20-22.V.1974, S. Peck (1 a, 1 9 BDGC); 42 km S. E. San Cristobal, 19.V.1974,S. Peck(20' O', 1 9 HAHC); Santo Domingo, 300m, 11-16.VIII.1983, B. Gill(2 a a, 3 9 9 BDGC).
REMARKS. In addition to the characters in the key and description, the smaller average size and the slight development of the depressions and bulges on the disc of the metastemum of large males separate 0. pubens sp.nov. from the closely related 0. sulcator (Fabricius).
28. Ontherus (Ontherus) sulcator (Fabricius), 1775 comb.nov. (Figs. 12, 74, 169-171, 285; Map 14)
Scarabaeus sulcator Fabricius, 1775: 27 (description) Scarabaeus sulcator Fabricius: Olivier, 1789: 142 (redescription) Copris sulcator Fabricius: Olivier, 1790: 168 (redescription) A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 75
Scarabaeus su!cator Fabricius: Fabricius, 1792: 61 (redescription) Copris sulcator Fabricius: Fabricius, 1801: 53 (redesription) Copris nisus Laporte, 1840: 79 (nee Olivier, 1789: 139) (description) Copris ovalipennis Blanchard, 1845 (1843): 180 (description) Ontherus sulcator (Fabricius): Erichson, 1848b: 761 (key) Ontherus sulcator (Fabricius): Gemminger and Harold, 1869: 1008 (catalog) Copris nisus Laporte: Gemminger and Harold, 1869: 1008 (catalog) Pinotus ovalipennis (Blanchard): Gemminger and Harold, 1869: 1010 (catalog) Copris ovalipennis Blanchard: Harold, 1869a: 59 (comment) Ontherus sulcator (Fabricius): Harold, 1869a: 59 (comment) Ontherus sulcator (Fabricius): Harold, 1869c: 503 (comment) Ontherus sulcator (Fabricius): Burmeister, 1874: 126 (list and comment) Copris nisus Laporte: Burmeister, 1874: 126 (list) Copris ovalipennis Blanchard: Burmeister, 1874: 126 (list) Copris senegalensis Gillet, 1910: 15 (catalog) Ontherus sulcator (Fabricius): Bruch, 1911: 187 (catalog) Copris nisus Laporte: Bruch, 1911: 187 (catalog) Copris ovalipennis Blanchard: Bruch, 1911: 187 (catalog) Ontherus sulcator (Fabricius): Gillet, 191 la: 319 (comment) Copris senegalensis Gillet: Gillet, 191 la: 319 (comment) Ontherus sulcator (Fabricius): Gillet, 191 lb: 58 (catalog) Ontherus nisus (Laporte): Gillet, 191 lb: 58 (catalog) Ontherus ovalipennis (Blanchard): Gillet, 191 lh: 58 (catalog) Copris senegalensis Gillet: Gillet, 191 lh: 58 (catalog) Ontherus sulcator (Fabricius): Bruch, 1915: 541 (catalog) Copris senega!ensis Gillet: Bruch, 1915: 541 (catalog) Ontherus sulcator (Fabricius): Lucas, 1920: 459 (catalog) Ontherus nisus (Laporte): Luederwaldt, 1930: 106 (comment) Ontherus sulcator (Fabricius): Luederwaldt, 1931: 364, 371, 385, 386, 387, 389, 392 (key, redescription and distribution) Ontherus nisus (Laporte): Luederwaldt, 1931: 372, 385, 386, 387, 392, 400, 419 (key, redescription and distribution) Copris ovalipennis Blanchard: Luederwaldt, 1931: 386, 387 (comment and listed as synonym) Ontherus senegalensis (Gillet): Luederwaldt, 1931: 386, 387 (comment and listed as synonym) Ontherus sulcator (Fabricius): Pessoa and Lane, 1941: 456, 458 (key) Ontherus nisus (Laporte): Pessoa and Lane, 1941: 457, 458 (key and redescription and distribution) Ontherus ovalipennis (Blanchard): Pessoa and Lane, 1941: 458 (listed as synonym) Ontherus senegalensis (Gillet): Pessoa and Lane, 1941: 458 (listed as synonym) Ontherus sulcator (Fabricius): Blackwelder, 1944: 206 (list) Ontherus nisus (Laporte): Blackwelder, 1944: 206 (list) Ontherus ovalipennis (Blanchard): Blackwelder, 1944: 206 (list) Ontherus senegalensis (Gillet): Blackwelder, 1944: 206 (list) Ontherus nissus (sic) (Laporte): Lange, 1947: 310 (list) Ontherus sulcator (Fabricius): Martinez, 1959: 71 (list and distribution) Copris nisus Laporte: Martinez, 1959: 71 (listed as synonym) Copris ovalipennis Blanchard: Martinez, 1959: 71 (listed as synonym) Copris senegalensis Gillet: Martinez, 1959: 72 (listed as synonym)
REDESCRIPTION. Colour. Body reddish brown to black. Pubescence fulvous. Length. 9-18 mm. Head. Anterior edge arcuate, feebly to moderately upturned, bisinuate medially, clypeogenal junction straight. Clypeus transversely wrinkled on apical four-fifths in both sexes, remainder coarsely punctate, in some specimens finely granulate; clypeofrontalsuture finely carinate, with small conical process medially in both sexes, process larger in females, smaller individuals with small swelling. Anterior genal edge much longer than lateral, genal surface transversely wrinkled on anterior half, posterior half granulate. Vertex coarsely granulopunctate, basal portion with smooth transverse area in both sexes. Pronotum.
Transverse, W/L = 1.6; anterior edge narrowly membranous and almost straight behind head insertion. Disc with small swelling on anterior sixth in males or slightly declivous on median fourth, posterior portion of declivity transversely swollen in females, anterior portion convex in smaller individuals, basal third to half with shallow and weakly delimited median longitudinal sulcus; punctures fine, rounded and scattered on disc and lateral swellings, 76 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
larger, oval and denser along base, anterior and lateral declivities. Elytra. Subquadrate, L/W
= 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, as deep as stria, approximately 1.2 times as wide as stria, separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely and evenly punctate throughout, lacking alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion pubescent and finely alutaceous throughout. Prosternum behind procoxa moderately pubescent, setae longer medially along base. Mesosternum shallowly and transversely impressed on disc, punctures coarse and dense throughout, pubescence fine, few longer setae along mesocoxa. Mesepisternum irregularly punctate, surface finely alu taceous and strongly carinate parallel to posterior edge. Metasternum with disc finely punctate, with round depression medially in males or finely and longitudinally sulcate in females; lateral lobes with punctures oval, each with anteromedial edge raised and sharp, almost evenly distributed on anterior half, broadly oval to rounded and restricted to lateral portion, surface of inner portion almost smooth on posterior half, surfacestrongly alutaceous along mesocoxal edge, microsculpture gradually weakening posteriorly leaving glossy area anterior to posterior coxa, pubescence moderately long and obliquely oriented, restricted to
anteromedial third; median lobe (Fig. 12) wide, L/m W = 1.2, lateral margins almost parallel on anterior half, convergent on posterior half, disc with strong arcuate impression in larger males or convex in females and smaller males, surface punctate and alutaceous throughout, punctures scarce and fine on anterior fifth, junction with mesosternum angulate and raised, usually produced into short carina medially on mesosternum. Legs. Anterior tibia quadri dentate in both sexes, apical tooth reduced in males, ventromedial carina lacking intervening setae on basal two-fifthsin males or basal fourth in females,apical spur straight and obliquely truncate, slightly spatulate apically in males. Posterior femora with anterior edge sharp, finely sulcated ventrally in males (Fig. 74) or unmodified in females; posterior surface with at most few longer setae along posterodorsal edge in both sexes, posterodorsal edge finely carinate, carina ill-defined on apical fourth in males or unmodified in females. Abdomen. Sternites 2-5 with several punctures laterally, anterior edge depressed and with row of small irregular punctures, surface finely alutaceous, finely punctate medially, sternite 6 finely punctate throughout, surface glossy, suture between sternites 5-6 narrowly and deeply
sulcate. Pygidium, W /L = 2.1 in males or 2.3 in females, punctures moderate in size and dense, in some specimens punctures fine on apical third, surfaceglossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 169-171). Tubus approximately 2.0 times as long as parameres, ventral portion finely sulcate. Parameres in lateral view with apex rounded, ventral portion straight, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 285.
MATERIAL STUDIED. 772a a, 819 9 9 (AMNH, BCRC, BDGC, BMNH, CASC, CMNC, CNCI, CUIC, FGIC, HAHC, ICCM, IMLA, ISNB, IZAV, LACM, MACN, MAMC, MCNZ, MCZC, MLPA, MLUH, MNHN, MZLU, NMPC, OSUC, SMFD, SMTD, TAMU, UNSM, USNM, UZMH, WBWC, WDEC, ZFMK, ZMHB, ZMUC, ZSMC)
TYPE MAT ERIAL Scarabaeus sulcator Fabricius: Type lost.
Copris nisus Laporte: Lectotype a (M V MA): Nisus (Cayenne) (on yellow paper)/ LECTOTYPE Copris nisus Laporte, 1840, Des. F. Genier, 1993. Type Material. studied (2 syntypes, 1 male and 1 female, glued on a cork plate). The internal sac of the aedeagus have been dissected and this specimen has been selected as Lectotype, which is here designated. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 77
• 0. sulcator
MAP 14. Known distribution of Ontherus sulcator.
Copris ovalipennis Blanchard: Lectotype 9 (MNHN): Green disk, 5395, 34/ 396/ Museum Paris, Buenos Ayres. A Corrientes, D'Orbigny 1834/ Copris ovalipennis Bl./ Type/LECT OTYPE, Copris, ovalipennis, Blanchard, 1845, Des. F. Genier, 1991. Type material studied and lectotype here designated.
Copris senegalensis Gillet: Type not studied (sufficient evidence).
NON-TYPE MAT ERIAL Distribution: ARGENTINA: Buenos Aires: Adrogue; Buenos Aires; Isla Martin Garcia; La Plata; Las Conchas (=El Tigre); Palermo; Pilar; San Fernando; San Pedro; 12 km S Suipacha; Temperley; Zelaya. Catamarca: No locality. Chaco: Barranqueras; Colonia Florencia*; Fontana; Presidencia Roque Saenz Pefia; Puerto Bermejo. Cordoba: Cordoba; San Francisco; San Vincente; Unquillo. Corrientes: Bella Vista; Itatf; Ituzaingo; Lambare; San Roque; Santo Tome; Juan E. Torrent. Entre Rios: Concordia; Lago.*; Palacio San Jose; Peranacito*; Pronunciamiento; Santa Elena. Formosa: Formosa. Jujuy: El Carmen; El Pongo; El Quemado; Laguna de Yala; Perico; Reyes; San Pedro. Misiones: Puerto Bemberg; Bompland (=Bomplano); Dos de Mayo; Eldorado; Loreto; Parque Nacional Iguazu; Posa das Misiones*; Puerto Deseado*; Puerto Iguazu; San Antonio; San Ignacio. Salta: Alemania; Coronel Olleros; Embarcacion; General Ballivian; Pocitos; Rio La Sala (P arque Nacional El Rey); Salta; San Lorenzo. Santa Fe: Alcorta; Carcarafia; La Hersilia; 20 km W Ocampo (Las Garzas); Rosario; San Cristobal; Santa Fe. Santiago del Estero: Rio Dulce; Rio Salado*. Tucuman: San Miguel de Tucuman. BOLIVIA: Beni: Rosario (Laguna Rogagua). Santa Cruz: Buena Vista; Region Chapare (400m); Saavedra Research Station; Santa Cruz de la Sierra (500m). BRAZIL: Amazonas: at Km 268 on AM 010*. Espirito Santo: Santa Cruz. Mato Grosso: Chapada dos Guimaraes; Cuiaba; Santo Antonio do Leverger. Mato Grosso do Sul: Corumba. Para: Braganc;a; Horacio*; Juruti. Parana: 78 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
P. N. Iguacu. Rio de Janeiro: Rio de Janeiro. Rio Grande do Sul: Barracas*; Cachoeira do sul; Canoas; Cruz Alta; Ipanema*; Pelotas; Porto Alegre; Santa Maria; Santo Augusto; Sao Jeronimo; Sao Leopoldo. Santa Catarina: Jaragua do Sul; Nova Teutonia (27°11 'S 52°23 'W) (300-500m); Rio das Antas. Sao Paulo: Puerto Epitacio*; Sao Paulo. COLOM BIA: No locality. ECUADOR: No locality. FRENCH GUIANA: Bas Maroni*; Nouveau Chantier*. Cayenne: Belizon; Cayenne; Gourdonville; Les Roches de Kourou; Pariacabo (Riviere de Kourou); Passoura. Saint-Laurent-du-Maroni: Chantier forestier Le Charvin (affluent du Maroni); La Forestiere; Mana; Massif de L'Acarouani (entre le Mana et le Maroni); Saint-Jean; Saint-Laurent-du-Maroni. GU YANA: Bartica; Blairmont; Esse quimbo; Kanuku Mts; Kartabo; St. Edwards. PARAGUAY: Estancia Postillon*. Alto Parana: Canjera*; Hohenau. Caaguazu: Estancia Primera*; Pastoreo. Caazapa: Caazapa, Central: San Lorenzo. Concepcion: Concepcion. Guaira: Independencia; Villarrica. Itapua: Encarnacion. Paraguari: Sapucay. San Pedro: San Estanislao; San Pedro. SURI NAM: Maronijne: Moengo. Wanica: Paramaribo. TRINIDAD: Caroni; Maraval; Port of Spain; Sans-Souci; St. Augustine*; Tunapuna. URUGUAY: Dolcras*; Montevideo; Nueva Palmira; Sal to; Santa Lucia; Soriano; Tacuarembo. VENEZUELA: Bolivar: El Bochinche, Res. Foresta! Imataca (200m)*; 10 km S El Dorado (200m); 33 km S El Dorado (220m); Hato Gruber*; Rio Supamo, 50 km S E El Manteco.
Additional data: Specimens were collected in chaco and subtropical forests, from cow dung in bamboo settlements, in carrion, or by using dung and ultraviolet light traps. Specimens collected in January to December.
REMARKS. Luederwaldt (1930, 1931) treated 0. nisus (Laporte) as a valid species. This is misinterpretation of Laporte's description, which seems to relate to Scarabaeus nisus
Olivier [ = Dichotomius nisus (Olivier)].
APPENDICULATUS GROUP
DIAGNOSIS. Species of this group separate from all others in the genus by following combination of characters: head tuberculate or carinate in 0. carinifrons, 0. irinus, 0. virescens; pronotum convex and usually longitudinally sulcate basally; mesometastemal suture angulate; posterior tibia of males feebly to strongly flattened and its posterodorsal edge produced, or tibia unmodified in males of 0. virescens.
29. Ontherus (Ontherus) appendiculatus (Mannerheim), 1829 comb.nov. (Figs. 60, 61, 68-70, 75, 76, 172<-174, 286; Map 15)
Copris appendiculata Mannerheim, 1829: 43 (description) Copris rotundatus Blanchard, 1845 (1843): 181 (description) Copris polynice Blanchard, 1845 (1843): 181 (description) Ontherus quadratus Erichson, 1848a: 564 (description) Ontherus appendiculatus (Mannerheim): Gemminger and Harold, 1869: 1008 (catalog) Ontherus polynice (Blanchard): Gemminger and Harold, 1869: 1008 (catalog) Ontherus quadratus Erichson: Gemminger and Harold, 1869: 1008 (catalog) Ontherus rotundatus (Blanchard): Gemminger and Harold, 1869: 1008 (catalog) Ontherus polynice (Blanchard): Harold, 1869a: 59 (comment) Ontherus rotundatus (Blanchard): Harold, 1869a: 59 (comment) Ontherus quadratus Erichson: Harold, 1869a: 59 (comment) Ontherus contractus Burmeister, 1874: 127 (description) Copris rotundata Blanchard: Burmeister, 1874: 127 (comment) Ontherus rotundatus (Blanchard): Harold, 1875h: 181 (comment) Ontherus quadratus Erichson: Harold, 1875h: 181 (comment) Ontherus appendiculatus (Mannerheim): Bruch, 1911: 186 (catalog) Ontherus contractus Burmeister: Bruch, 1911: 187 (catalog) Ontherus rotundatus (Blanchard): Bruch, 1911: 187 (catalog) Copris polynice Blanchard: Bruch, 1911: 187 (catalog) A REVISION OF THE NEOTROPJCAL GENUS ONTHERUS ERICHSON 79
Ontherus quadratus Erichson: Bruch, 1911: 187 (catalog) Ontherus appendiculatus (Mannerheim): Gillet, 191 lb: 57 (catalog) Ontherus polynice (Blanchard): Gillet, 191 lh: 57 (catalog) Ontherus quadratus Erichson: Gillet, 191 lb: 57 (catalog) Ontherus rotundatus (Blanchard): Gillet, 1911b: 57 (catalog) Ontherus appendiculatus (Mannerheim): Bruch, 1915: 541 (catalog) Ontherus appendiculatus (Mannerheim): Lucas, 1920: 459 (catalog) Ontherus appendiculatus (Mannerheim): Luederwaldt, 1930: 107 (comment) Ontherus appendiculatus (Mannerheim): Luederwaldt, 1931: 371, 372, 389, 391, 392 (key, redescription and distribution) Ontherus polynice (Blanchard): Luederwaldt, 1931: 389 (listed as synonym) Ontherus quadratus Erichson: Luederwaldt, 1931: 389 (listed as synonym) Ontherus rotundatus (Blanchard): Luederwaldt, 1931: 389 (listed as synonym) Ontherus contractus Burmeister: Luederwaldt, 1931: 389 (listed as synonym) Ontherus appendiculatus (Mannerheim): Pessoa and Lane, 1941: 457, 458 (key, description and distribution) Ontherus polynice (Blanchard): Pessoa and Lane, 1941: 458 (listed as synonym) Ontherus quadratus Erichson: Pessoa and Lane, 1941: 458 (listed as synonym) Ontherus rotundatus (Blanchard): Pessoa and Lane, 1941: 458 (listed as synonym) Ontherus appendiculatus (Mannerheim): Blackwelder, 1944: 206 (list) Ontherus polynice (Blanchard): Blackwelder, 1944: 206 (listed as synonym) Ontherus quadratus Erichson: Blackwelder, 1944: 206 (listed as synonym) Ontherus rotundatus (Blanchard): Blackwelder, 1944: 206 (listed as synonym) Ontherus contractus Burmeister: Blackwelder, 1944: 206 (listed as synonym)
REDESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 7-13 mm. Head. Anterior edge arcuate, moderately upturned, at most slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled through out, in some specimens punctate on posterior fourth. Clypeofrontal suture broadly arcuate and finely carinate, with small conical process medially, with larger portion anterior to suture, process reduced to small swelling in smaller individuals. Anterior genal edge slightly longer than lateral, genal surface wrinkled throughout, in some specimens punctate on small area adjacent to eye. Vertex punctate except on small area posteriorly. Pronotum. Transverse,
W/L = 1.8; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc slightly declivous on anterior fifthbehind head insertion in males or narrowly declivous on most of the width in femalesand smaller males, basal half with longitudinal and sharply delimited median sulcus, in some specimens sulcus with additional coarse punctures laterally or sulcus partially interrupted; punctures usually fine and scattered on disc and lateral swellings, larger and denser along base, anterior and lateral declivities, large and confluent on anterior angles. Elytra. Slightly transverse, L/W = 0. 9. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, slightly deeper than stria, approximately 1.3 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely and evenly punctate, punctures slightly finerand denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion pubescent and finely alutaceous through out. Prosternum behind procoxa pubescent, setae longer on disc. Mesostemum deeply and transversely impressed on disc, punctures coarse and dense throughout, pubescence fineon disc, longer along mesocoxae. Mesepistemum irregularly punctate, surface finely alu taceous, moderately to strongly carinate parallel to posterior edge. Metastemum with disc finely punctate,finely and shallowly sulcate, sulcus wider posteriorly in larger males; lateral lobes on anterior half with punctures oval, each with anteromedial edge raised and sharply carinate, except along mesocoxae, posterior half with punctures rounded, scattered, surface strongly alutaceous along mesocoxal edges, microsculpture gradually weakening posteri orly, leaving surface on posterior half completely glossy between punctures, pubescence moderately long and obliquely oriented; median lobe (Figs. 68-70) wide, L/mW = 1.5, lateral margins almost parallel on anterior two-thirds, slightly convergent on posterior third, gradually widening anteriorly, anterior angles of lateral margins subquadrate to rounded 80 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
internally, in some specimens (Fig. 70) lateral margins narrow throughout and anterior angles of lateral margins angulate internally, surface punctate and slightly alutaceous, junction with mesosternum strongly angulate medially, raised throughout its width, slightly produced into short longitudinal carina on mesosternum.Legs. Anterior tibia quadridentate in both sexes, apical tooth (Figs. 60, 61) slightly projecting anteriorly in males, ventromedial carina lacking intervening setae on basal fourth in both sexes, apical spur straight and rounded apically. Posterior femur (Figs. 75, 76) triangularly produced on posterodorsal edge apically, posteroventral surface sulcate, sulcus extending anteriorly at apex and apicoventral surface with shallow to moderately deep impression in larger males or simple in females and smaller males. Posterior tibia (Figs. 75, 76) with internal edge usually triangularly produced on basal half in males or simple in females and smaller males. Abdomen. Sternites 2-5 with few punctures laterally, anterior edge with complete row of setiferous punctures, surface glossy, sternite 6 narrow, finely punctate throughout, suture between sternites 5-6 narrowly and
deeply sulcate. Pygidium, WIL = 2.1 in males or 2.3 in females, punctures moderately coarse and dense, evenly distributed, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 172-174). Tubus approximately 2.0 times as long as parameres, ventral portion unmodified. Parameres in lateral view with apex rounded, ventral portion slightly concave, deeply notched basally, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 286.
MATERIAL STUDIED. 566 a a, 647 Q Q (AMNH, CUIC, BCRC, BDGC, CASC, CNCI, DEIC, FGIC, HAHC, ICCM, IMLA, ISNB, IZAV, LACM, MACN, MCNZ, MCZC, MLPA, MLUH, MNHN, MZLU, NMPC, PAIC, SMFD, SMTD, TAMU, USNM, UZMH, WBWC, WDEC, ZFMK, ZMHB, ZMUC, ZSMC).
TYPE MATERIAL Copris appendiculata Mannerheim: Holotype O' (by monotypy)(ZMAS): Brasilia, 4835 - !.(handwritten on green paper)/ HOLOTYPE, Copris, appendiculata, Mannerheim, 1829, vidit F. Genier, 1991. Holotype studied.
Copris polynice Blanchard: Lectotype O' (MNHN): Green disk, 6424 34/ Museum Paris, Santa Cruz de la Siera, D'Orbigny 1834/ 1431/ Copris polynice BL/ Type/ LECTOTYPE, Copris, polynice, Blanchard, 1845, Des. F.Genier, 1991. Type material studied and lectotype here designated.
Copris rotundatus Blanchard: Holotype Q (MNHN): Green disk, 5759 34/ 754/ Museum Paris, Province de Corrientes, D'Orbigny 1834/ Copris rotundatus BL/ Type/ HOLOTYPE, Copris, rotundatus, Blanchard, 1845, vidit F. Genier, 1991. Holotype studied.
Ontherus quadratus Erichson: Lectotype O' (ZMHB): Gujana Schomb (hand written)/ Hist.-coll., nr. 26632/ Zool. Mus., Berlin/ SYNTYPE, Copris, quadrata Erich/ Ontherus Manh., appendiculatus, det. J. Huijbregts 1984/ LECTOTYPE, Copris quadrata, Erichson, 1848, Des. F. Genier, 1992. Type material studied, lectotype and paralectotype here designated. Paralectotypes: same data as lectotype (1 O', 1 Q ZMHB).
Ontherus contractus Burmeister: Lectotype Q (MACN): Tucuman/ 103/ contractus, Nob., Rep. Arg./ Syntypus, Ontherus contractus, Burmeister, 1874/ LECTOTYPE, Ontherus, contractus, Burmeister, 1874, Des. F. Genier, 1992. Type material studied, lectotype and paralectotype here designated. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 81
• 0. appendiculatus
MAP 15. Known distribution of Ontherus appendiculatus.
Paralectotype: Same data as lectotype (1 9 MACN).
NON-TYPE MATERIAL Distribution: ARGENTINA: El Quenada*; Gran Chaco. Buenos Aires: San Ignacio. Catamarca: El Alto; Icafio; La Vifia;Recreo. Cordoba: Rio Cuarto. Corrientes: Ituzaingo, Isla Apipe Grande. Jujuy: El Quemado; lkm. N Pampa Blanca (700m). Misiones: Puerto Iguazu; San Ignacio. Salta: Coronel Moldes; 15km S El Carril, Vifiaco; General Ballivian; Metan; Rosario de Lerma (1325m); Salta; Tabillas*; Tartagal. Santiago del Estero: Santiago del Estero. Tucuman: Alpachiri; Burruyacu, Chilcas; Concepcion; La Soledad*; Los Vazques; San Pedro; Tapia (600m); Trancas. BOLIVIA: Chaco de Bolivia*; Rio Japacani*; San Antonio de Mapiri*. Beni: Cavinas; Ivon; Laguna Rogagua, Rosario; Reyes; Riberalta. Chuquisaca: Machareti; Tiguipa. La Paz: Caranavi. Santa Cruz: Buena Vista; Lagunillas; Robore (300m); Saaverde Res. Sta.; San Jose de Chiquitos; Santa Cruz de la Sierra (500m); Tatarenda; Rancho Yoay; Sara (= Provincia Gutierrez). BRAZIL: Acre: Rio Branco. Alagoas: Macei6. Amazonas: Macura; Manaus. Bahia: Lacerda*; San Antonio da Barra*; Terra Nova. Ceara: Fortaleza. Goias: Anapolis; Goiania; Jataf; Viannopolis. Mato Grosso: 20mi. W Araguaia (=Araguaiana); Carumba (=Carumbe); Esta9ao Joaquim Murtinho; Santo Ant6nio do Leverger. Mato Grosso do Sul: Campo Grande. Minas Gerais: Aguas Vermelhas; Diamantina; Lassance; Montes Claros; Sete Lagoas; Uberaba. Para: Bragan9a; Santarem; Santo Ant6nio do Taua; Soure. Paraiba: Independencia. Pernambuco: No locality. Rio de Janeiro: Nova Friburgo; Petr6polis. Rio Grande do Norte: Ceara-Mirim; Natal. Rio Grande do Sul: Santa Maria; Sao Jeronimo; Sao Leopoldo. Santa Catarina: Jaragua do Sul. Sao Paulo: Batatais; Fazenda Barreiro Rico; Fazenda Campininas, Mogi Guacu; Piracicaba; Puerto Epitacio*; Ribeirao Preto; Rio Claro; Sao Paulo; Ubirama (= Lencois Paulista). COLOMBIA: Casanare: Orocue. Cundinamarca: Medina. Distrito Especial: Bogota. Meta: Carimagua (300m); 30km E San Martin; Villavicencio. Tolima; Mariquita. ECUADOR: Napo: Napo river*. FRENCH GUIANA: Cayenne: Cayenne; Crique Gourdonville. GU YANA: Essequibo. PARAGUAY: Cabafias*; Trinidad*. Amambay: 82 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Parque Nacional Cerro Cora. Boqueron: Filadelfia; Mariscal Estigarribia. Caaguazu: Estancia Primera*; Pastoreo. Caazapa: Caazapa. Central: Asuncion; Capiata;San Lorenzo; Santa Clara. Concepcion: Horqueta. Cordillera: Caacupe; San Bernardino. Guaira: Inde pendencia; Villarrica. Paraguari: Parque Nacional Ybycui. San Pedro: Carumbe*; San Pedro. URUGUAY: Salto. VENEZUELA: Anzoategui: El Tigre. Apure: Hato El Frio (lOOm)*; San Fernando. Aragua: El Limon (450m); Pie del Cerro (820-1120m). Bolivar: Ciudad Bolivar; 48km ES E Ciudad Bolivar, RanchoSanta Rita; 35km. S W Puerto Ordaz; Roraima (1800m). Carabobo: Valencia. Distrito Federal: Caracas; Antimano. Guarico: Cazorla;La Cruz-Ruviera*. Miranda: Caucagua. Monagas: Josepin (=La Josefina?); 42km. S E Maturfn. Portuguesa: Guanare (400m); Estacion Experimental San Nicolas, 56km. Guanare (l 80m). Tachira: Santo Domingo (300m). Yaracuy: Boquer6n; San Felipe; Yaritagua.
Additional data: Some specimens were collected fromcattle dung and human feces in forests. Specimens were also collected using malaise/flight intercept traps set in sandy woodlands, meat traps, and ultraviolet or incandescent light traps. Specimens collected in January to December.
REMARKS. In the material studied variation was present in the thickness of the coxal margin of the median lobe of the mesosternum. Some individuals have a narrow margin anteriorly (Fig. 70) and the internaledge of the margin is angulate [as in type(s) of C. polynice and 0. contractus]. In other individuals the margin is much wider (Fig. 68) and the internal edge is rounded [as in type(s) of C. appendiculata, C. rotundatus and 0. quadratus]. Few individuals have the intermediate condition (Fig. 69). One specimen had an angulate inner edge on one side and the rounded edge on the other side. Variation was also encountered in male sexual dimorphism. In some specimens the anterior tibia has the apical and penultimate teeth relatively closely set (Fig. 60), whereas in others the teeth are more widely separated (Fig. 61). In addition, the posteroventral sulcus is extended anteriorly in some cases (Fig. 75), and the anterior edge of the sulcus is straight in others (Fig. 76). No clear correlations could be made between these variations and geographic distribution. The aedeagus and sclerite of the internal sac are identical over the entire range and, therefore, I consider these variations intraspecific.
30. Ontherus (Ontherus) ulcopygus sp.nov. (Figs. 2, 3, 175-177, 287; Map 16)
ET YMOLOGY. Ulcopygus, (ulcerate+ buttock) Greek adjective, refers to pygidial margin aspect.
DESCRIPTION. Colour. Body dark brown to black. Pubescence fulvous. Length. 10-15 mm. Head. Anterior edge arcuate, moderately upturned, at most slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled throughout, in some specimens punctate on posterior fourth. Clypeofrontal suture broadly arcuate, finely and sharply carinate, with small conical process medially, with larger portion posterior to suture, process reduced to small, but acute, tubercle in smaller individuals. Anterior genal edge slightly longer than lateral, genal surface wrinkled on anterior half, coarsely punctate on posterior half. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc declivous on anterior fifth, declivity approximately fourth total width of pronotum in males or fifth in females, convex in smaller individuals, basal half with longitudinal and moderately to sharply delimited median sulcus, in some specimens sulcus partially interrupted; punctures usually fine and scattered on disc and lateral swellings, larger and denser along base on A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 83 median fifth,anterior and lateral declivities, large and confluent on anterior angles. Elytra.
Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, deeper than stria, approximately 1.3 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely and evenly punctate, punctures slightly finer and denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion pubescent and finely alutaceous throughout. Prostemum behind procoxa pubescent, setae longer on posterior two-thirds. Mesosternum moderately transversely impressed on disc, punctures coarse and dense throughout, pubescence fine on disc, longer along meso coxae. Mesepisternum irregularly punctate, surface finely alutaceous, moderately to strongly carinate parallel to posterior edge. Metasternum with disc finely and scarcely punctate, with longitudinal impression on posterior half, impression much wider in larger males; lateral lobes on anterior half with punctures oval, each with anteromedial edge raised and sharply carinate, except along mesocoxae, posterior half with punctures rounded, scattered, surface strongly alutaceous along mesocoxal edges, surface between punctures finely alutaceous throughout, pubescence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 1.8, lateral margins almost parallel on anterior two-thirds, slightly convergent on posterior third, anterior angles of lateral margins angulate internally, surface punctate, each puncture with anterior edge finely carinate on anterior third, surface between puncture slightly alutaceous, junction with mesosternum strongly angulate medially, raised throughout its width, slightly produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth slightly projecting anteriorly in males, ventromedial carina lacking intervening setae on basal fourth in both sexes, apical spur straight, slightly tapered from base to apex in males or parallel-sided and inwardly bent on apical fifthin females. Posterior femur triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on same plane, forming large subrectangular depression in larger males, in smaller males both surfaces partially to completely separated by pos teroventral edge, apicoventral portion of depression never extending anteriorly or femur unmodified in females. Posterior tibia with internal edge feebly to moderately expanded on basal third, posterior portion of expansion feebly to moderately toothed posteriorly in males or simple in females. Abdomen (Figs. 2, 3). Stemites 2-5 with few weakly impressed punctures laterally, anterior edge with complete row of setiferous punctures, surface finely alutaceous, stemite 6 finely punctate throughout, suture between stemites 5-6 narrowly sulcate laterally. Pygidium, W/L = 1.9 in males or 2.1 in females, punctures moderately coarse and dense, scattered along midline, surface glossy, apical margin ill-defined in both sexes. Male genitalia (Figs. 175-177). Tubus approximately 2.8 times as long as parameres, ventral portion almost flat on apical half. Parameres in lateral view subtriangular, with apex rounded, ventral portion slightly concave, deeply and widely notched basally, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 287.
MATERIAL STUDIED. 65 a a, 78 Q Q (AMNH, BMNH, CASC, CNCI, DEIC, ICCM, ISNB, LACM, MLUH, MNHN, NMPC, PAIC, SMFD, SMTD, USNM, ZFMK, ZMHB, ZMUC). Holotype CJ' (CNCI): BRAZIL, Distrito Federal, Estacao Florestal, Cabeca do Veado, 1100 m, 27 oct. 1971, E.G., I. and E.A. Munroe/ HOLOTYPE, Ontherus ulcopygus, F. Genier, 1992. Dissected. Allotype Q (CNCI): Same data as holotype.
Paratypes: No locality ( 1 a, 1 Q MLUH) (3 a a MNHN). BRAZIL: No locality ( 1 Q CASC) (1 a MLUH) (2 Q Q ZMUC) (1 a MNHN); Slid-Brasil (1 a, 1 Q MNHN) (1 Q NMPC) (1 CJ', 1 Q SMTD); Cachoeira* (1 Q MLUH). Distrito Federal: Estacao 84 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. ulcopygus
MAP 16. Known distribution of Ontherus ulcopygus.
Florestal, Cabeca do Veado, llOOm, 23-27.X.1971, E.G., I. and E. A. Munroe (2aa, 2Q Q CNCI), same as before except: 18-19.X.1971 (1a CNCI),27.X.1971 (1a, 1 Q CNCI), 29-30.XI.1971 (1a, 1 Q CNCI); Parque Nacional*, l lOOm, 3.III.1970, J. M. and B. A. Campbell (2Q Q CNCI). Goias: Jatahi (=Jataf), 1898, Ch. Pujol (1 a MNHN); 20km N. Sao Joao da Alienc;a,18.IV.1956, F. S. Truxal (2a a, 1Q LACM). Mato Grosso: Chapada (= Chapada dos Guimaraes) (1Q USNM), same as before except: Acc. No. 2966 (4a a, 6Q Q ICCM), Acc. No. 2966, I (4a a, 3 Q Q ICCM), Acc. No. 2966,III (1a,1Q ICCM), Acc. No. 2966,IV (1a ICCM),IX (2a a USNM), Acc. No. 2966, IX (9 a a, 17Q Q ICCM), X (1a USNM), Acc. No. 2066, X (1 a, 4Q Q ICCM), XI (1 Q USNM), Acc. No. 2966, XI (3a a, 2 Q Q ICCM), Acc. No. 2966, XII (4a a, 6 Q Q ICCM). Mato Grosso do Sul: CampoGrande, X.1947, A. Maller Coll. (1a AMNH). Minas Gerais: No locality, Reinhardt (2a a, 1Q ZMUC); Aguas Vermelhas, XII.1983, Coll. M. Alvarenga (1Q ICCM); Lagoa Santa (1 Q MLUH), same as before except: Reinhardt (3aa, 2Q Q ZMUC); S-Joao d'el Rey (=Sao Joao del Rei),Coll. Van Volxem(1 Q ISNB); Sete Lagoas,Reinhardt (1 Q ZMUC), same as before except: 22.XII.1898, F. Ohaus S. (1Q SMFD) (1Q USNM); Uberaba (1a ISNB) (1Q PAIC). Parana: No locality (1a BMNH). Rio Grande do Sul: No locality (1a ZMHB). Santa Catarina: Ibirama, 22.X.1947, Boje Benzon (1a, 1 Q ZMUC). Sao Paulo: Campinas, Alwine Braatz V. (2a a, 2Q Q ZMHB); Franca, IX.1910, E. Gargi leg. (1Q USNM), same as before except: X.1910 (1Q USNM); Mato d. Gov.*, 24.1.[19]27, F. Ohaus (1a USNM); Rio Claro, III.1949, P. Pereira (1 a ISNB); Salta Grande de Paranapenema, 21.X.1908, Haseman (1Q ICCM); Sao Paulo (1 a DEIC) (2 Q Q ZFMK); Ypiranga(=Ipiranga),F. Ohaus S. (1a,1Q SMFD) (1 a USNM),same as before except: 3.XI.[19]26 (1a SMFD), 17.XI.[19]26 (1 Q USNM). PERU: No locality (1Q NMPC).
REMARKS. The Peruvian record is based on a single, old specimen. In the absence of recently collected material, I consider this record dubious. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 85
31. Ontherus (Ontherus) podiceps Harold, 1868 comb.nov. (Figs. 3, 4, 178-180, 288; Map 17)
Ontherus podiceps Harold, 1868a: 82 (description) Ontherus podiceps Harold: Gemminger and Harold, 1869: 1008 (catalog) Ontherus podiceps Harold: Gillet, 191 lb: 58 (catalog) Ontherus podiceps Harold: Luederwaldt, 1931: 372, 392 (key, distribution and redescription) Ontherus podiceps Harold: Pessoa and Lane, 1941: 456 (key) Ontherus podiceps Harold: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark brown to black. Pubescence fulvous. Length. 11-15 mm. Head. Anterior edge arcuate, moderately upturned,slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior two-thirds in females, remainder of surface coarsely punctate, disc with usually distinct transverse swelling on posterior two-thirds. Clypeofrontal suture broadly arcuate, finely carinate in males or finely and sharply carinate in females, with small conical process medially, with larger portion posterior to suture, process reduced to small, but acute, tubercle in smaller individuals. Anterior genal edge subequal in length to lateral, genal surface moderately to coarsely punctate throughout. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc slightly declivous on anterior fifth, declivity approximately one-fifth the total width of pronotum in males or one-sixth in females, convex in smaller individuals, basal half with longitudinal and sharply delimited median sulcus; punctures fine and scattered on disc and lateral swellings, larger and denser along base on median fourth, anterior and lateral declivities, large and almost confluent on anterior angles. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, deeper than stria, approximately 1.4 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely and evenly punctate, punctures slightly larger and denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion pubescent and finely alutaceous throughout. Prosternum behind procoxa pubescent, setae longer on posterior two-thirds. Mesosternum deeply and trans versely impressed on disc, punctures coarse and dense throughout, coarsely microsculptured, pubescence fine on disc, much longer and denser along mesocoxae. Mesepisternum irregu larly punctate, surface finely alutaceous, moderately to strongly carinate parallel to posterior edge. Metasternum with disc finely and rather densely punctate, with longitudinal impres sion on posterior two-thirds, impression much wider in males; lateral lobes on anterior half with punctures oval, lateral ones each with anteromedial edge finely carinate, on ante riormost portion punctures confluent and forming indistinct to distinct wrinkles, posterior half with punctures rounded, scattered, surface moderately alutaceous along mesocoxal edges, surface between punctures on posterior half glossy, pubescence moderately long and obliquely oriented; median lobe moderately wide, L/mW = 2.3, lateral margins slightly convergent on anterior two-thirds, convergent on posterior third, anterior angles of lateral margins angulate internally, surface finely punctate, surface between punctures slightly alutaceous, junction with mesosternum angulate medially, raised throughout its width, slightly produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth slightly projecting anteriorly in males, ventromedial carina lacking intervening setae on basal fourth in both sexes, apical spur straight, slightly tapering from base to apex in males or parallel-sided and inwardly bent on apical fifth in females. Posterior femur triangularly produced on posterodorsal edge apically, pos teroventral sulcus and posterior surface fused and almost on same plane, forming large subrectangular depression in larger males, in smaller males both surfaces partially to completely separated by posteroventral edge, apicoventral portion of depression never 86 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. podiceps
MAP 17. Known distribution of Ontherus podiceps.
extending anteriorly or femur unmodified in females. Posterior tibia with internal edge slightly to moderately expanded on basal third, posterior portion of expansion finely toothed posteriorly in males or simple in females. Abdomen (Figs. 4, 5). Sternites 2-5 with few coarse punctures laterally, anterior edge with complete row of setiferouspunctures, surface glossy, sternite 6 finely punctate throughout, suture between sternites 5-6 narrowly and rather deeply sulcate throughout, sulcus slightly wider laterally. Pygidium, W/L = 2.0 in males or 2.1 in females, punctures moderately coarse and dense, surface glossy, apical margin partially to completely ill-defined in both sexes. Male genitalia (Figs. 178-180). Tubus approximately 2.3 times as long as parameres, ventral portion moderately produced on apical fourth. Parameres in lateral view subtriangular, with apex rounded, ventral portion slightly concave, deeply and widely notched basally, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 288.
MATERIAL STUDIED. 28 a a, 13 Q Q (CASC, ISNB, MLUH, MNHN, NMPC, SMFD, SMTD, USNM, ZMAS, ZMHB, ZMUC).
TYPE MATERIAL Holotype CJ' (MNHN): Brasilia/ podiceps, t. Harold/ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthiir, 1952/ HOLOTYPE, Ontherus podiceps, Harold, 1868, vidit F. Genier, 1991. Holotype studied.
NON-TYPE MATERIAL Distribution: BRAZIL: Bahia: No locality. Espirito Santo: No locality; Timbui. Minas Gerais: Sete Lagoas. Rio de Janeiro: Nova Friburgo. Sao Paulo: No locality. COLOM BIA: No locality. FRENCH GUIANA: Cayenne.
Additional data: Collecting month: December.
REMARKS. The Colombian and French Guianan records need to be confirmed. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 87
32. Ontherus ( Ontherus) azteca Harold, 1869 comb.nov. (Figs. 77, 181-183, 289; Map 18)
Ontherus a:teca Harold, l 869c: 503 (description) Ontherus azteca Harold: Bates, 1887: 50 (distribution and redescription) Ontherus azteca Harold: Gillet, 191 lh: 57 (catalog) Ontherus villosus Luederwaldt, 1930: 107 (description and distribution) new synonymy. Ontherus villosus Luederwaldt: Luederwaldt, 1931: 371, 372, 391 (key, distribution and redescription) Ontherus azteca Harold: Luederwaldt, 1931: 399 (key) Ontherus villosus Luederwaldt: Pessoa and Lane, 1941: 457 (key) Ontherus azteca Harold: Blackwelder, 1944: 206 (list) Ontherus vi!losus Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus villosus Luederwaldt: Lange, 1947: 310 (list) Ontherus strius Howden and Young, 1981: 122 (key, description and distribution) new synonymy. Ontherus strius Howden and Young: Gill in Hanski and Carnbefort,1991: 412 (biology)
REDESCRIPTION. Colour. Body reddish brown to black. Pubescence fulvous. Length. 11-17 mm. Head. Anterior edge arcuate, moderately upturned, slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior two-thirds, surface punctate on posterior third, with feeble to moderate transverse swelling on disc. Clypeofron tal suture usually arcuate and finely carinate, angulate and weakly carinate in some individu als, with small and acute conical process medially, with larger portion posterior to suture. Anterior genal edge slightly shorter than lateral, genal surface with punctures almost confluent throughout. Vertex punctate except on small area posteriorly. Pronotum. Trans verse, W IL = 1.8; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc slightly declivous on anterior fifth behind head insertion in males or almost convex in females and smaller males, basal half with longitudinal and vaguely delimited median sulcus, punctures of sulcus unevenly distributed; punctures moderate in size on disc and lateral swellings, slightly larger and denser along base, anterior and lateral declivities,
confluent on anterior angles in some individuals. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, deeper than stria, approximately 1.3 times as wide as stria and separated by 2-4 diameters on disc and apical declivity, conspicuously encroaching intervals. Intervals convex, surface finely and evenly punctate, punctures denser on apical declivity, almost completely glossy in some individuals to moderately alutaceous in some others. Thoracic sterna. Proepistemum with posterolateral portion densely pubescent and finely alutaceous throughout. Prosternum behind procoxa finely pubescent, setae apposed, usually with brush of longer setae medially on posterior edge, in some individuals (from Ecuador, Peru and Bolivia) prosternum minutely pubescent throughout. Mesosternum slightly and transversely impressed on disc, punctures moderate in size, coarse and dense throughout, pubescence fine and short throughout. Mesepisternum regularly punctate, punctures rounded, umbilicate and of different sizes, surface finely alutaceous, strongly and sharply carinate parallel to posterior edge. Metasternum with disc finelypunctate, widely and rather deeply sulcate; lateral lobes on anterior half with punctures rounded and umbilicate, outermost punctures each with posterolateral edge ill-defined, posterior half with punctures rounded, scattered, surface strongly alutaceous on anterior half, microsculpture gradually weakening
posterioriy, pubescence long and obliquely oriented; median lobe wide, L/mW = 1.5, lateral margins almost parallel on anterior half, slightly convergent on posterior half, lateral margins narrow throughout and anterior angles of lateral margins angulate internally, surface punc tate, anteriormost punctures larger and setiferous, surface between punctures strongly alutaceous anteriorly, junction with mesosternum angulate medially, finely carinate through out, not produced into short carina longitudinally on mesostemum. Legs. Anterior tibia quadridentate in both sexes, apical tooth projecting anteriorly and notched at apex (Mexican population) or with small to wide lobe externally in males or unmodified in females, 88 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. azteca * 0. cambeforti
MAP 18. Known distribution of Ontherus azteca and 0. camheforti. ventromedial carina lacking intervening setae on basal third in both sexes, apical spur spiniform and slightly outwardly bent. Posterior coxae finely serrate along posterior edge. Posterior femur (Fig. 77) elongate, triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on same plane, forming largely depressed area, surface of depression strongly alutaceous in larger males, apical edge of depression sharply delimited, femur simple in females. Posterior tibia (Fig. 77) with internal edge carinate to lobate on basal half in males or simple in females and smaller males. Abdomen. Sternites 2-5 punctate laterally, anterior edge with row of setiferous punctures laterally, surface alutaceous, sternite 6 finely punctate laterally, minutely punctate medially, suture between sternites 5-6 deeply and sharply sulcate laterally. Pygidium, WIL = 2.1 in males or 2.2 in females,punctures moderately coarse and dense, evenly distributed, surface glossy, apical margin clearly delimited in males or ill-defined in females, basal portion usually deeply sulcate along margin medially. Male genitalia (Figs. 181-183). Tubus approximately 2.5 times as long as parameres, flattened apically. Parameres in lateral view elongate and with apex rounded, ventral portion slightly concave, deeply notched basally, posteroventral portion long. Sigmoid sclerite of internal sac as in Figure 289.
MATERIAL STUDIED. 45 a a, 23 Q Q (AMNH, BDGC, BMNH, CASC, FGIC, HAHC, ICCM, IMLA, INBC, ISNB, MNHN, MZSP, SMFD, SMTD, USNM, WDEC, ZMHB).
TYPE MAT ERIAL Ontherus azteca Harold: Holotype Q (BMNH): (Mexico) Oaxaca/ Type/ Ontherus azteca, Harold,® QI 731/ Mexico, Salle Coll./ Ontherus aztecus, Harold, type, apnd (sic) Salle./ B.C.A., p50 sp l/ HOLOTYPE, Ontherus azteca, Harold, 1869, vidit F. Genier, 1991. Holotype studied.
Ontherus villosus Luederwaldt: Lectotype a (MZSP): Sao Paulo,Ypiranga, XI.[19]06,Luederw. leg./Ontherus a, villosus Luederw., Luerderw. det. 29/ 5.886./ blue disk/ COTYPO (pink paper)/ 16720/ A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 89
LECTOTYPE, Ontherus villosus, Luederwaldt, 1930, Des. F. Genier, 1992. Type material studied and lectotype here designated.
Ontherus strius Howden and Young: Holotype ct (USNM): (Panama) Canal Zone, B.C.I., 17.IX.1975, on ground, Col. O.P. Young, 0900 hrs/ Type Number, 104484, U.S.N.M./ Holotype, Ontherus strius, H. Howden and, 0. Young/ Ontherus azteca, Harold, 1869, Det. F. Genier, 1991. Holotype studied.
NON-TYPE MATERIAL Distribution: BOLIVIA: Cochabamba: No locality. Santa Cruz: Santa Cruz. BRAZIL: Bahia: Entre Rios. Espirito Santo: Timbui. Mato Grosso: Chapada dos Guimaraes. Rio de Janeiro: Tijuca. Santa Catarina: Nova Teutonia, 27°11' S 52°23' W; Rio Vermelho. Sao Paulo: Anhembi (Fazenda Barreiro Rico); Bocaina; Ipiranga; Itu (Fazenda Pau D' Alho); Sao Paulo. COLOMBIA: Meta: Villavicencio. COSTA RICA: Guanacaste: Estancia Pitilla (9km S. Santa Cecilia, P. N. Guanacaste) (700m). Limon: Finca de E. Rojas, Sector Cerro Cocori (150m); 6km N Guacimo. Puntarenas: El Rodeo* (Ciudad Colon). ECUA DOR: Napo: Limoncocha (250m); 2km N Limoncocha (250m); Km 7 .3 Sarayacu-Loreto Rd. (1200m). Pastaza: 25km N NEPuyo (lOOOm); 22km SE Puyo (900m). MEXICO: J alapa*. Chiapas: Parque Laguna Belgica*. Jalisco: l .4km NW Barra de Navidad. Nayarit: El Cora. Quintana Roo: 24km W Chetumal. PANAMA: Canal Zone: Barro Colorado Island. PARAGUAY: Caaguazu: Paso Yobai *. Caazapa: Caazapa. Paraguari: Ybicui. PERU: Cuzco: Rio Tono. Junin: Chanchamayo; San Francisco de Satipo.
Additional data: Some specimens were collected using a dung trap baited with human feces. Some traps were set in virgin forests along a nature trail. Two specimens were collected using a feces trap set in a cloud forest. One specimen was collected in a trap baited with decaying meat set in an unspecified habitat. Specimens collected in January to April, June to December.
REMARKS. The type material of 0. azteca Harold, 0. villosus Luederwaldt and 0. strius Howden and Young was studied and no differences were found to support the validity of the two latter species. Therefore, 0. villosus Luederwaldt and 0. strius Howden and Young are junior synonyms of 0. azteca Harold. In the material studied one female specimen (MNHN) collected from Bragam:;a (Brazil: Para) was found. This specimen was compared with females of 0. azteca and 0. cambef orti, but could not be associated with certainty to either of these species.
33. Ontherus (Ontherus) cambefortisp.nov. (Figs. 78, 184-186, 290; Map 18)
ETYMOLOGY. Cambeforti, a patronym in gratitude to Yves Cambefort, collector of holotype of this species.
DESCRIPTION. Colour. Body dark reddish brown. Pubescence fulvous. Length. 12-16 mm. Head. Anterior edge arcuate, moderately upturned, slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half, surface with punctures confluent on posterior half, with feeble to moderate transverse swelling on disc. Clypeofrontal suture arcuate, weakly carinate, with small and acute conical process medially. Anterior genal edge subequal in length to lateral, genal surface with punctures confluent throughout. Vertex punctate except on a small area posteriorly. Prono tum. Transverse, W IL = 1. 7; anterior edge narrowly membranous and almost straight behind head insertion. Disc slightly declivous on anterior fifth behind head insertion in larger individuals or almost convex in smaller individuals, basal half with longitudinal, ill-defined median sulcus, punctures of sulcus scarce and shallow; punctures moderate in size on disc 90 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 and lateral swellings, slightly larger and denser along base, anterior and lateral declivities, not confluent on anterior angles. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, deeper than stria, approxi mately 1.2 times as wide as stria and separated by 2-4 diameters on disc and apical declivity, slightly encroaching intervals. Intervals convex, surface finely and evenly punctate, punc tures denser on apical declivity, surface between punctures completely glossy. Thoracic sterna. Proepisternum with posterolateral portion densely pubescent and finely alutaceous throughout. Prosternum behind procoxa finely pubescent, setae apposed, with brush of longer setae medially on posterior edge. Mesosternum slightly and transversely impressed on disc, punctures moderate in size, coarse and dense throughout, pubescence fine and short, few longer setae along mesocoxae. Mesepisternum regularly punctate, punctures rounded, umbilicate and of different sizes, surface finely alutaceous, strongly and sharply carinate parallel to posterior edge. Metasternum with disc finely punctate, widely and rather deeply sulcate; lateral lobes on anterior half with punctures rounded and umbilicate, outermost punctures each with posterolateral edge ill-defined, posterior half with punctures rounded, scattered, surface strongly alutaceous on anterior half, microsculpture gradually weakening posteriorly leaving posterior third completely glossy, pubescence long and obliquely oriented; median lobe wide, L/mW = 1.6, lateral margins almost parallel on anterior half, slightly convergent on posterior half, lateral margins narrow throughout and anterior angles of lateral margins angulate internally,surface finely punctate, anteriormost punctures slightly larger and finely setiferous, surface between punctures finely alutaceous anteriorly, junction with mesosternum angulate medially, finely carinate throughout, not produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate, apical tooth projecting anteriorly, moderately to widely lobed externally in males or unmodified in females,ventromedial carina lacking intervening setae on basal third in males or basal fourth in females, apical spur spiniform and slightly outwardly bent. Posterior coxae finely serrate along posterior edge. Posterior femur (Fig. 78) triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on same plane, forming largely depressed area, surface of depression punctate, surface between punctures glossy, finely alutaceous along posterior edge in larger males, apical edge of depression sharply delimited, femur simple in females. Posterior tibia (Fig. 78) with internal edge at most slightly lobate on basal half in males, simple in females and smaller males. Abdomen. Sternites 2-5 punctate laterally, anterior edge with row of setiferous punctures laterally, surface glossy, sternite 6 finely punctate laterally, minutely punctate medially, suture between sternites 5-6 deeply and sharply sulcate laterally. Pygidium, W/L = 2.3 in males or 2.4 in females, punctures coarse and dense, evenly distributed, surface glossy, apical margin clearly delimited in males, basal portion slightly sulcate along margin medially. Male genitalia (Figs. 184-186). Tubus approximately 2.0 times as long as parameres, convex apically. Parameres in lateral view elongate and with apex rounded, ventral portion slightly concave, deeply notched basally, posteroventral portion long. Sigmoid sclerite of internal sac as in Figure 290.
MATERIAL STUDIED. 3 O' O', 1 9 (ICCM, MNHN).
Holotype O' (MNHN): Guyane (French Guiana), Oyapok, 2°47N 52°29W, 9(IX).[19]72, Y. Cambefort/ HOLOTYPE, Ontherus cambeforti, F. Genier, 1992. Dissected.
Allotype 9 (MNHN): MUSEUM PARIS, GUYANE FRAN<;., St-LAURENT, (Maroni), Audouit 1862./ 331, 62 (recto of green disc)/ ALLOTYPE, Ontherus cambeforti, F. Genier, 1992. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 91
Paratypes: GUYANA: No locality, 12.VII.1925, near camp, human faeces (1 O' ICCM). FRENCH GUIANA: Saint-Laurent-du-Maroni: St-Laurent, Melinon, 1862 (1 O' MNHN).
34. Ontherus (Ontherus) grandis Luederwaldt, 1931 comb.nov. (Figs. 187-189, 291; Map 19)
Ontherus grandis Luederwaldt, 1931: 419 (description) Ontherus grandis Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus grandis Luederwaldt: Martinez, 1959: 71 (catalog)
REDESCRIPTION. Colour. Body dark brown to black. Pubescence reddish. Length. 13-21 mm. Head. Anterior edge arcuate, moderately upturned, narrowly emarginate me dially in males or slightly bisinuate in females, clypeogenal junctionfinely notched. Clypeus transversely wrinkled throughout, in some specimens scarcely to moderately punctate on posterior fifth. Clypeofrontal suture subangular and finely carinate, with large conical process medially with larger portion posterior to suture, acute in males or transverse in females, process reduced to small swelling in smaller individuals. Anterior genal edge slightly longer than lateral, genal surface wrinkled on anterior half, punctate on remainder of surface. Vertex punctate on anterior on-half, surface smooth and convex on posterior half.
Pronotum. Transverse, W!L = 1.7; anterior edge widely membranous and emarginate internally to eyes behind head insertion. Disc strongly declivous on anterior fifthbehind head insertion, upper edge of declivity straight and bulging on median third in males or slightly arcuate and slightly bulbous in females, disc convex in smaller individuals, basal half with longitudinal and weakly delimited median sulcus; punctures usually fine and slightly more scattered on posterior half of disc and lateral swellings, larger and denser along base on median tenth, lateral declivities and anterior half of disc, large and almost confluent on anterior angles. Elytra. Subquadrate, L/W = 1.0. Striae fine, finely alutaceous, weakly impressed on disc, distinctly deeper basally and on apical declivity, punctures ill-defined, slightly deeper than stria, approximately 1.0 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals almost flat on disc, slightly convex on basal fourth, surface finely and evenly punctate, punctures slightly larger and denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion pubescent and finely alutaceous throughout. Prosternum behind procoxa pubescent, setae subequal in length. Mesosternum deeply and transversely impressed on disc, punctures moderate in size and dense throughout, pubescence fine on disc, much longer and denser along mesocoxae. Mesepisternum rather finely and irregularly punctate, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metastemum with disc finely punctate and glabrous, widely depressed on basal two-thirds, depression wider anteriorly in males, depression reduced to longitudinal sulcus in smaller individuals; lateral lobes setiferously punctate throughout, punctures moderate in size, rounded, setae obliquely oriented to erect and long, surface between punctures strongly alutaceous throughout; median lobe wide, L/mW = 1.7, lateral margins almost parallel on anterior two-thirds, slightly convergent on posterior third, anterior angles of lateral margins angulate internally, surface setiferously punctate on anterior half, setae similar to those on lateral lobes, posterior half finely punctate along mesocoxae, surface between punctures slightly alutaceous throughout, junction with mesosternum strongly angulate medially, sharply carinate throughout, slightly produced into short carina longitu dinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, teeth similar in both sexes, slightly more slender in males, ventromedial carina lacking intervening setae on basal fourth in both sexes, apical spur spiniform and inwardly bent on apical third in males or parallel-sided and inwardly bent on apical third in females. Posterior femur triangularly and acutely produced on posterodorsal edge apically, posteroventral surface sulcate, sulcus 92 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 weakly delimited anteriorly and not extending anteriorly at apex or simple in females and smaller males. Posterior tibia unmodified. Abdomen. Sternites 2-5 with few coarse and rather small punctures laterally, anterior edge with complete row of setiferous punctures, surface alutaceous throughout, sternite 6 punctate throughout, suture between sternites 5- 6 weakly sulcate laterally. Pygidium, WIL = 2.0 in males or 2.2 in females, punctures fine and moderately dense, evenly distributed, surface glossy, apical margin weakly delimited in both sexes. Male genitalia (Figs. 187-189). Tubus approximately 2.3 times as long as parameres, ventral portion sharply carinate apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly concave, slightly notched basally, dorsal portion concave. Sigmoid sclerite of internal sac as in Figure 291.
MATERIAL STUDIED. 16a a, 23 9 9 (BDGC, CMNC, HAHC, MLPA, MNHN).
TYPE MATERIAL Holotype 9 (MNHN): Argentina, Prov. Tucuman, Lagunita 3000m., C.A. Baer 1-1903/ Museum Paris, 1936, coll A. Boucomont/ Typus/ Ontherus a, grandis n. sp., Typus, Luederwaldt det. 1931/ HOLOTYPE, Ontherus grandis, Luederwaldt, 1931, vidit F. Genier, 1991. Holotype studied.
NON- TYPE MATERIAL Distribution: ARGENTINA: Salta: El Rey National Park (900m); El Rey National Park, Pozo Verde Trail km 5 (950m); El Rey National Park, Pozo Verde Trail km 7 (I OOOm); El Rey National Park, Rio LaSala (900m); El Ucumar, 20km N La Caldera (780m); El Ucumar, 22km N La Caldera (1550m). Tucuman: No locality; Ruta 307 (1300m).
Additional data: Some specimens were collected using a dung trap set in chaco forest. Specimens collected in February, March, December.
35. Ontherus (Ontherus) carinifrons Luederwaldt, 1930 comb.nov. (Figs. 190-192, 292; Map 19)
Ontherus carinifrons Luederwaldt, 1930: 105 (description) Ontherus carinifrons Luederwaldt: Luederwaldt, 1931: 370, 378, 391 (key, distribution and redescription) Ontherus carinifrons Luederwaldt: Balthasar, 1938: 223 (comment) Ontherus carinifrons Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus carinifrons Luederwaldt: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark brown to black. Pubescence fulvous. Length. 10-13 mm. Head. Anterior edge arcuate, moderately upturned, lacking emargination medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior three-fourths in females, remainder of surface punctate. Clypeofrontal suture broadly arcuate, strongly carinate, carina slightly angulate, wider and higher medially. Anterior genal edge subequal in length to lateral, genal surface moderately to coarsely punctate throughout, anteriormost portion slightly wrinkled in some individu als. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, WIL= 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex, larger individuals with barely distinct swelling on anterior fifth medially, basal half with longitudinal and rather sharply delimited median sulcus; punctures fine and scattered on disc and lateral swellings, larger and denser along base medially, anterior and lateral declivities, large and almost confluent on anterior angles.
Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, slightly deeper than stria, approximately 1.3 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals slightly convex, surface finely, sharply and evenly punctate, punctures slightly larger and much denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 93
• 0. grandis
• 0. carinifrons
+ 0. virescens
* 0. irinus
MAP 19. Known distribution of Ontherus grandis, 0. carinifrons, 0. virescens, and 0. irinus.
Proepistemum with posterolateral portion pubescent and finely alutaceous throughout. Prosternum behind procoxa pubescent, setae longer on posterior half. Mesosternum deeply and transversely impressed on disc, punctures coarse and dense throughout, coarsely microsculptured, pubescence fine on disc, much longer and denser along mesocoxae. Mesepisternum irregularly punctate, surface finely alutaceous, strongly carinate parallel to posterior edge. Metastemum with disc finely and rather densely punctate, with longitudinal impression on posterior two-thirds, impression wider and deeper in larger males; lateral lobes on anterior half with punctures oval, lateral ones each with anteromedial edge finely carinate, on anteriormost portion punctures confluent and forming indistinct to distinct wrinkles, posterior half with punctures rounded, scattered, surface moderately alutaceous along mesocoxal edges, surface between punctures on posterior half slightly alutaceous, pubes cence obliquely oriented, moderately long along mesocoxal edge, decreasing in length toward lateral edge; median lobe moderately wide, L/mW = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins angulate internally, coarsely and setiferously punctate on anterior half and along mesocoxae, setae conspicuous, finely punctate on posterior half, surface between punctures slightly alutaceous, junction with mesosternum angulate medially, raised throughout its width, finely produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth slightly projecting anteriorly in males, ventromedial carina lacking intervening setae on basal third in both sexes, apical spur straight, parallel-sided in males or slightly spatulate apically in females. Posterior femur triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on the same plane, forming large subrectangular depression in larger males, in smaller males both surfaces partially to completely separated by posteroventral edge, apicoventral portion of depression never extending anteriorly and with edge ill-defined to moderately sharp, femur unmodified in females. Posterior tibia with internal edge carinate to moderately lobate on basal third, posterior portion of 94 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 expansion unmodifiedto finely toothedposteriorly in males or simple in females.Abdom en. Sternites 2-5 with few coarse punctures laterally, anterior edge with complete row of complete to fused and irregular setiferous punctures, surface glossy, sternite 6 finely punctate throughout, suture between sternites 5-6 narrowly and rather deeply sulcate throughout, sulcus wider laterally. Pygidium, W /L = 2.1 in males or 2.2 in females, punctures moderately coarse and dense, surface glossy, apical margin clearly delimited in males or ill-definedin females. Male genitalia (Figs. 190-192). Tubus approximately 2.0 times as long as parameres, ventral portion slightly to moderately depressed on apical fourth. Parameres in lateral view subtriangular, with apex rounded, ventral portion slightly concave, deeply and widely notched basally, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 292.
MATERIAL STUDIED. 29a a, 23 Q Q (ICCM, ISNB, MNHN, NMPC, PAIC, SMFD, USNM, ZMHB).
TYPE MATERIAL Lectotype O" (MZSP): O"I (Brazil) Amazonas, Manaos, VII.[19]27, Zink.an/ blue disk/ Cotypus/ Ontherus O", carinifrons, Luederw., Luederw. det. 29./ Ontherus carinifrons Luederw/ 16364/ LEC TOTYPE, Ontherus carinifrons, Luederwaldt, 1930, Des. F. Genier, 1992. Type material studied, lectotype and paralectotypes here designated.
Paralectotypes: Same data as lectotype ( 1 O", 2 Q Q MZSP).
NON-TYPE MATERIAL Distribution: BRAZIL: Amazonas: Manaus; Tefe. Para: Barcarena; Bragarn.;a; Obidos; Santarem; Taperinha. PERU: No locality. Cuzco: Rio Tono.
Additional data: Specimens collected in February, April, May, July, September.
36. Ontherus (Ontherus) virescens (Lucas), 1857 comb.nov. (Figs. 193-195, 293; Map 19)
Choeridium virescens Lucas, 1857: 103 (description) Canthidium virescens (Lucas): Gemminger and Harold, 1869: 1006 (catalog) Ontherus virescens (Lucas): Gillet, 191lh: 58 (catalog) Ontherus virescens (Lucas): Luederwaldt, 1930: 106, 107 (comment) Ontherus virescens (Lucas): Luederwaldt, 1931: 371, 381, 382, 383 (key, distribution and redescription) Ontherus virescens (Lucas): Pessoa and Lane, 1941: 456 (key) Ontherus virescens (Lucas): Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark brown, with strong metallic green sheen on pronotum and elytra, sheen weaker on head, ventral surface and legs. Pubescence fulvous. Length. 10-11 mm. Head. Anterior edge arcuate, moderately upturned, slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third in males or on anterior three-fourths in females, remainder of surface punctate. Clypeofrontal suture broadly and regularly arcuate, strongly carinate, slightly higher medially. Anterior genal edge subequal in length to lateral, genal surface moderately to coarsely punctate throughout in males or punctures confluent in females. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion, distinctly wider over short distance medially. Disc convex, basal half with shallow and vaguely delimited longitudinal median sulcus; punctures fine on disc and lateral swellings, large and denser along base medially, anterior and lateral declivities, large and almost confluent on anterior angles. Elytra. Slightly wider than large, L/W = 0.9. Striae moderately wide and impressed on disc and apical declivity, punctures rounded, slightly deeper than stria, approximately 1.6 times as wide as stria and separated by 2-3 diameters on disc and A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 95 apical declivity, conspicuously encroaching intervals. Intervals moderately convex, surface finely, sharply, and evenly punctate, punctures slightly larger and denser on apical declivity, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepister num with posterolateral portion pubescent and finely alutaceousthroughout. Prosternum behind procoxa pubescent, setae scattered, slightly longer on disc. Mesosternum mod erately and transversely impressed on disc, punctures coarse and dense throughout, pubescence fine on disc, slightly longer and denser along mesocoxae. Mesepisternum irregularly punctate, surface finely alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely punctate, with longitudinal impression on anterior three-fourths, extending on median lobe, impression wider and slightly deeper in larger males; lateral lobes on anterior half with punctures oval, each with anteromedial edge finely carinate, on anteriormost portion punctures almost confluent, posterior half with punctures rounded, scattered, surface strongly alutaceous along mesocoxal edges, sur face between punctures on posterior half slightly alutaceous, pubescence obliquely oriented, moderately long along mesocoxal edge, decreasing in length toward lateral edge; median lobe rather wide, L/m W = 1.8, lateral margins parallel on anterior half, convergent on posterior half, anterior angles of lateral margins angulate internally, coarsely and setiferously punctate along mesocoxae and along anterior edge, setae conspicuous along mesocoxal edges, decreasing in length and thickness toward anterior edge, finely punctate medially, surface between punctures alutaceous, junction with mesosternum slightly angulate medially, raised throughout its width, not produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, teeth slender, apical tooth obliquely projecting in males or unmodified in females, ventromedial carina lacking intervening setae on basal fourth in both sexes, apical spur straight and slightly spatulate in males or parallel-sided and inwardly bent apically in females. Posterior femur unmodified in both sexes. Posterior tibia simple in both sexes. Tarsal claws modified on all legs, J-shaped, as long as penultimate tarsal segment which is enlarged apically and laterally compressed, spiniformly produced ventrally at apex. Abdomen. Sternites 2-5 with few coarse punctures laterally, anterior edge with com plete row of entire to fused and irregular setiferous punctures, surface slightly alu taceous, sternite 6 scarcely and finely punctate throughout, suture between sternites 5-6 narrowly and rather deeply sulcate throughout, sulcus wider laterally. Pygidium, W /L
= 2.2 in males or 2.0 in females, punctures coarse, moderately large and dense, surface glossy, apical margin clearly delimited in males or ill-defined in females. Male genitalia (Figs. 193-195). Tubus approximately 2.3 times as long as parameres, ventral portion unmodified. Parameres in lateral view subtriangular, with apex rounded, ventral portion slightly concave, deeply and widely notched basally, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 293.
MATERIAL STUDIED. 20' O', 2 9 9 (ICCM, MNHN, UZMH).
TYPE MATERIAL Holotype 9 (by monotypy)(MNHN): Museum Paris, Goyaz a Cuyaba, De Castelnau 6-47I green disk, 6-47I Choeridium virescens Luc./ Type/ Ontherus virescens Luc., P. Pereira det. 957I HOLOTYPE, Choeridium, virescens, Lucas, 1857, vidit F. Genier, 1991. MNHN. Holotype studied.
NON- TYPE MATERIAL Distribution: BRAZIL: Mato Grosso: Cuiaba (Cuyaba). Rio de Janeiro: Rio de Janeiro
Additional data: Specimens collected in April. 96 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
REMARKS. This peculiar species has greatly modifiedtarsi. These tarsi are similar to those of the Australian Onthophagus that are phoretic on marsupial fur. This adaptation suggests a similar ecological niche for 0. virescens.
37. Ontherus (Ontherus) irinus Balthasar, 1938 comb.nov. (Figs. 196-198, 294; Map 19)
Ontherus irinus Balthasar, 1938: 222 (description) Ontherus irinus Balthasar: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark brown, pronotum and elytra with strong iridescent sheen. Pubescence fulvous. Length. 10-13 mm. Head. Anterior edge arcuate, moderately upturned, lacking emargination medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior three-fourths in females, remainder of surface punctate. Clypeofrontal suture broadly arcuate, strongly carinate, carina slightly angulate medially, slightly wider and higher medially. Anterior genal edge slightly longer than lateral, genal surface shallowly punctate throughout, anteriormost portion slightly wrinkled. Vertex rather coarsely and densely punctate except on small area
posteriorly. Pronotum. Transverse, W/L = 1.5; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex, larger individuals with small transverse swelling on anterior fifthmedially, basal half with longitudinal and weakly delimited median sulcus; punctures ill-defined on disc and lateral swellings in males or moderate in size on disc and lateral swellings in females, larger and denser along base medially, anterior and lateral declivities, rounded and coarse on anterior angles. Elytra. Slightly transverse,
L/W = 0.9. Striae rather fine, deeper on apical declivity, punctures oval, deeper than stria, each with small tubercle, approximately 2.5 times as wide as stria and separated by 3-5 diameters on disc and apical declivity. Intervals convex, surface minutely and evenly punctate on disc, punctures fine and denser on apical declivity, lacking any traces of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion pubescent and finely alutaceous throughout. Prostemum behind procoxa pubescent, setae subequal in length throughout. Mesosternum shallowly and transversely impressed on disc, punctures coarse and dense throughout, uneven in size, pubescence fine on disc, few stout setae along posterior edge. Mesepistemum coarsely and irregularly punctate, surface alu taceous, bluntly carinate parallel to posterior edge. Metasternum with disc finely and rather densely punctate, with longitudinal impression on posterior half, impression wider and deeper in males; lateral lobes on anterior half with punctures oval, lateral ones each with anteromedial edge finely carinate, posterior half with punctures rounded, scattered, leaving large smooth area internally, surface moderately alutaceous along mesocoxal edges, surface between punctures on posterior half glossy, pubescence obliquely oriented, moderately long along mesocoxal edge, decreasing in length toward lateral edge, minute on posterior half;
median lobe moderately wide, L/mW = 2.0, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins angulate inter nally, finely punctate throughout, surface between punctures alutaceous, junction with mesosternum slightly angulate medially, weakly carinate throughout its width, not produced into short carina longitudinally on mesostemum. Legs. Anterior tibia quadridentate in both sexes, apical tooth slightly projecting anteriorly in males, ventromedial carina lacking intervening setae on basal two-thirds in males or half in females, apical spur straight, slightly spatulate apically. Posterior femur triangularly produced on posterodorsal edge apically, apex of ventroposterior sulcus not extending anteriorly in males or femur unmodified in females.Posterior tibia with internaledge finely carinate to moderately lo bate on basal third, posterior portion of expansion toothed posteriorly in males or simple in females. Abdomen. Sternites 2-5 with several coarse punctures laterally, anterior edge with few ill-defined punctures, surface alutaceous laterally, stemite 6 and median portion of sternite 5 finely A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 97
punctate, suture between sternites 5-6 narrowly and finely sulcate laterally. Pygidium, W/L = 2.2 in males and females, punctures coarse and dense, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 196-198). Tubus approximately 2.4 times as long as parameres, ventral portion depressed on apical fourth. Parameres in lateral view elongate, with apex rounded, ventral portion slightly concave, slightly emargi nate basally, ventral edge of right paramere widely and triangularly membranous, dorsal portion almost straight. Sigmoid sclerite of internal sac as in Figure 294.
MATERIAL STUDIED. 1 a, 1 Q (BMNH, NMPC).
TYPE MATERIAL Holotype a (by monotypy)(NMPC): Theophilo Oltoni ( =Teofilo Otoni), Minas Geraes/ TYPUS/ Ontherus irinus n. sp., Dr. V. Balthasar det./ irinus m./ Mus. Nat. Pragae, inv. 65457./HOLOTYPE, Ontherus irinus, Balthasar, 1938, viditF. Genier, 1991. Holotype studied.
NON-TYPE MATERIAL Distribution: BRAZIL: Rio de Janeiro: Rio., C. Darwin, 87-42. 415 (1 Q BMNH).
REMARKS. Ontherus irinus is intermediate between those of the Appendiculatus and Brevipennis groups. Externally it shares many characters with 0. carinicollis and 0. vires cens, but the aedeagus has the membranous projection, which characterizes species in the following group.
BREVIPENNIS GROUP
DIAGNOSIS. The following combination of characters will distinguish males and females of the following species: mesometasternal suture angulate; dorsum with general glossy appearance; clypeofrontal suture strongly carinate, never tuberculate medially; pronotum strongly convex. In males, parameres (in lateral view) (Figs. 199, 202, 205, 208, 211, 214, 217, 220, 223, 226) denticulate subapically, possessing strongly enlarged and membranous ventral projection (Figs. 201, 204, 207, 210, 213, 216, 219, 222, 225, 228). Modifications in parameres unique to this species group, with exception of 0. irinus (see remarks under that species).
38. Ontherus (Ontherus) raptor sp.nov. (Figs. 79, 199-201, 295; Map 20)
ETYMOLOGY. Raptor, (plunderer) Latin adjective, refers to shape of the anterior tibiae.
DESCRIPTION. Colour. Body reddish brown. Pubescence fulvous. Length. 15 mm. Head. Anterior edge arcuate, moderately upturned, bisinuate medially, clypeogenal junction almost straight. Clypeus with weak and transverse wrinkles on anterior third in males, remainder of surface almost smooth. Clypeofrontal suture almost semicircular, strongly carinate anteriorly, carina gradually decreasing in height posteriorly, carina slightly lower on median third. Anterior genal edge slightly shorter than lateral, genal surface almost smooth throughout. Vertex finely and shallowly punctate on anterior two-thirds, posterior third smooth. Pronotum. Transverse, W/L = 1.3; anterior edge narrowly membra nous and slightly arcuate behind head insertion. Disc simple and strongly convex, basal half lacking longitudinal median sulcus; punctures fine, dense and shallow on anterior angles, remainder of surface smooth, except single row of small umbilicate punctures along posterior margin. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed, slightly deeper on apical declivity, punctures rounded, deeper than stria and distinctly encroaching on intervals, approximately 2.0 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals convex, surface completely glossy, 98 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
* 0. raptor
• 0. planus
• 0. rectangulidens
MAP 20. Known distribution of Ontherus raptor, 0. planus, and 0. rectangulidens.
lacking any traces of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent laterally and finely alutaceous throughout. Prosternum behind procoxa pubescent, setae subequal in length throughout. Mesosternum shallowly and transversely impressed on disc, punctures coarse and dense, uneven in size, lacking on posterior fourth, pubescence fine on disc, few stout setae along coxal edge. Mesepisternum shallowly and rather scarcely punctate, surface finely alutaceous, cari nate parallel to posterior edge. Metasternum with disc smooth, with strong longitudinal impression on posterior two-thirds; lateral lobes with punctures rather scarce, oval, each with anteromedial edge finely and sharply carinate, surface between punctures alutaceous on anterior half, surface between punctures glossy on posterior half, pubescence obliquely oriented and restricted to anterior half, moderately long along mesocoxal edge, decreasing in length toward lateral edge; median lobe moderately wide, L/mW = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded internally, surface lacking punctures, finely alutaceous throughout, junction with mesosternumstrongly angulate medially, sharply carinate throughout its width, produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadriden tate, apical tooth projecting anteriorly and far apart from penultimate tooth, ventromedial carina lacking intervening setae on basal third, apical spur spiniform. Posterior femur (Fig. 79) triangularly produced on posterodorsal edge apically, apex of ventroposterior sulcus extending anteriorly, sharply delimited apically. Posterior tibia (Fig. 79) with internal edge finely carinate to moderately lobate on basal third, posterior portion of expansion acutely toothed posteriorly, tarsal claws short, approximately as long as ventroapical projec tion of last tarsal segment. Abdomen. Sternites 2-5 with few fine punctures laterally, anterior edge with irregular row of umbilicate punctures laterally, surface finely alutaceous laterally, sternite 6 completely glossy, suture between sternites 5-6 sulcate laterally.
Pygidium, WIL = 2.0, deeply sulcated along basal margin, punctures minute, surface between punctures glossy, apical margin clearly delimited. Male genitalia (Figs. 199-201). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 99
Tubus approximately 2.2 times as long as parameres, ventral portion depressed on apical fifth.Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into short tooth just before apex, ventral edge of left paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 295.
REMARKS. Ontherus raptor is described based on a single specimen. However, this specimen is a well-developed male, which features the best diagnostic characters. The unique specimen was collected in a threatened portion of the rain forest in northernBolivia.
MATERIAL STUDIED. 1 O' (ZSMC). Holotype O' (ZSMC): Sarampiuni, San Carlos, lOOOm. 2.9./ BOLIVIA 1950, leg. W. Forster/ Zool. Staatsslg., Miinchen/HOLOTYPE, Ontherus raptor, F. Genier, 1992. Dissected.
39. Ontherus (Ontherus) planus sp.nov. (Figs. 80, 202-204, 296; Map 20)
ETYMOLOGY. Planus, (flat, level) Latin adjective, refers to ventral aspect of anterior femora in males.
DESCRIPTION. Colour. Body black. Pubescence fulvous. Length. 12-17 mm. Head. Anterior edge arcuate, moderately upturned, slightly bisinuate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior three-fourths in females, remainder of surface finely to moderately punctate. Clypeofrontal suture subtrapezoidal, strongly carinate, carina almost even in height and thickness anteri orly, much lower but distinct laterally. Anterior genal edge much shorter than lateral, genal surface moderately to strongly wrinkled on anterior half, remainder of surface shallowly punctate. Vertex rather coarsely and densely punctate except for small area posteriorly.
Pronotum. Transverse, W/L = 1.5; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin, in most individuals, shortly enlarged medially. Disc strongly convex, basal half longitudinally depressed medially; punctures minute on disc, fine on anterior and lateral declivities, with row of umbilicate punctures along base. Elytra.
Subquadrate, L/W = 1.0. Striae moderately wide, similarly impressed on disc and apical declivity, punctures rounded, deeper than stria, encroaching on intervals, approximately 2.0 times as wide as stria and separated by 3-5 diameters on disc and apical declivity. Intervals convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and finely alutaceous through out. Prosternum behind procoxa scarcely pubescent, setae mostly restricted to base and lateral portion, subequal in length throughout. Mesosternum shallowly and transversely impressed on disc, puncture coarse and dense anteriorly, lacking on posterior third, uneven in size, pubescence fine on anterior third, stouter on median third and lacking posteriorly. Mesepisternum shallowly and rather scarcely punctate, surface alutaceous, carinate parallel to posterior edge. Metasternum with disc finely punctate and alutaceous throughout, with deep longitudinal impression, impression wider and slightly deeper in males; lateral lobes on anterior half with punctures oval, each puncture with anteromedial edge sharply carinate, posterior half with punctures broadly oval to rounded and mostly restricted to metacoxal edge, leaving large smooth area internally, surface between punctures alutaceous, except for small glossy area along metacoxal edge internally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, decreasing in length toward lateral edge, lacking on posterior half; median lobe moderately wide, L/m W = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded internally, surface finely punctate throughout, surface between punctures strongly 100 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
alutaceous, junction with mesosternumstrongly angulate medially, sharply carinate through out its width, produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth projecting anteriorly in males, ventromedial carina lacking intervening setae on basal third in both sexes, apical spur straight, obliquely truncate and slightly spatulate apically. Posterior femur (Fig. 80) triangularly produced on postero dorsal edge apically, apex of ventroposterior sulcus extending anteriorly and sharply delim ited throughout in males or femur unmodified in females. Posterior tibia (Fig. 80) with internal edge finely carinate on basal third, posterior portion of carina toothed posteriorly in males or simple in females. Abdomen. Sternites 2-5 with few fine punctures laterally, anterior edge with row of moderately umbilicate punctures laterally, surface finely alutaceous, sternite 6 glossy, suture between sternites 5-6 narrowly and deeply sulcate laterally. Pygidium, WIL = 1.9 in males or 2.1 in females, deeply sulcate along basal margin, punctures minute, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 202- 204). Tubus approximately 2.2 times as long as parameres, ventral portion emarginate apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 296.
MATERIAL STUDIED. 5 a a, 4 9 9 (CASC, CMNC, USNM). Holotype a (CMNC): El Palmar, (YUNGAS)-BOLOV IA, SEPT-OCT. 1948/ H. and A. HOWDEN COLLECTION, ex. A. Martinez Coll./ HOLOTYPE, Ontherus planus, F. Genier, 1992. Dissected.
Allotype 9 (CASC): Buena Vista, Bolivia, XII-[19]48/ E. R. Leach, Collection/ Collection of the CALIFORNIA ACADEMY OF SCIENCES, san Francisco, Calif./ ALLOTYPE, Ontherus planus, F. Genier, 1992. Dissected.
Paratypes: BOLIVIA: Santa Cruz: Buena Vista, XII.[19]48, E. R. Leach Collection (20' O', 3 9 9 CASC), same as before except: Ichijo (=Ichilo?), XI-XIl.[19]48, L. Pefia (2 a a USNM).
40. Ontherus (Ontherus) lichyiMartinez, 1947 comb.nov. (Figs. 81, 205-207, 297; Map 21)
Ontherus lichyi Martinez, 1947: 45 (description) Ontherus mirandai Pereira, 1955: 461 (description) Ontherus lichyi Martinez: Roze, 1955: 44 (list) Ontherus lichyi Martinez: Pereira and Martinez, 1960: 49 (comment) Ontherus mirandai Pereira: Pereira and Martinez, 1960: 49 (comment)
REDESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 12-17 mm. Head. Anterior edge arcuate, moderately upturned, slightly bisinuate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior three-fourths in females, remainder of surface finely to moderately punctate. Clypeofrontal suture subquadrate, subtrapezoidal in smaller individuals, strongly carinate, carina tuberculate anterolaterally, almost even in height and thickness anteriorly, much lower but distinct laterally. Anterior genal edge much shorter than lateral, genal surface moderately to strongly wrinkled on anterior half, remainder of surface finely and shallowly punctate. Vertex rather coarsely and densely punctate in some individuals to minutely punctate, except for small area posteriorly. Pronotum. Transverse, WIL = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin, in most individuals, shortly enlarged medially. Disc strongly convex, basal half longitudinally and shallowly depressed medially; punctures minute on disc, fine on anterior and lateral A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 101
declivities, with few umbilicate punctures along base. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide, similarly impressed on disc and apical declivity, punctures rounded, slightly deeper than stria on disc, deep on apical declivity, slightly encroaching on intervals, approximately 2.0 times as wide as stria and separated by 3-5 diameters on disc and apical declivity. Intervals convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion moderately pubescent and finely alutaceous throughout. Prostemum behind procoxa densely pubescent on basal two-thirds, setae subequal in length and thickness throughout. Mesosternum shallowly and transversely impressed on disc, punctures irregular and dense anteriorly, lacking on posterior fourth, pubescence fine on anterior third, stouter on median third and lacking posteriorly. Mesepistemum shallowly and rather scarcely punctate, surface alutaceous, carinate parallel to posterior edge. Metastemum with disc minutely punctate and moderately alutaceous throughout, with deep longitudinal impression on posterior two-thirds, impression wider and slightly deeper in males; lateral lobes on anterior half with punctures oval, anteriormost punctures each with anteromedial edge sharply carinate, posterior half with punctures broadly oval to rounded and mostly restricted to metacoxal edge, leaving scarcely punctate area internally, surface between punctures alutaceous, except for small glossy area along metacoxal edge internally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior half; median lobe moderately wide, Lim W = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded inter nally, surface lacking punctures and strongly alutaceous, junction with mesosternum strongly angulate medially, sharply carinate throughout its width, produced into short carina longitudinally on mesosternum. Legs. Anterior tibia appearing tridentate, with apical (fourth) tooth vestigial and adjacent to apical spur in males or quadridentate in females, ventromedial carina lacking intervening setae on basal half in males or third in females, apical spur straight and acute in males or obliquely truncate and slightly spatulate apically in females. Posterior femur (Fig. 81) triangularly produced on posterodorsal edge apically, outer edge of projection rounded and sharp, ventroposterior sulcus sharply delimited throughout, extending anteriorly at apex in males or femurunmodified in females.Posterior tibia (Fig. 81) unmodified in both sexes. Abdomen. Stemites 2-5 with occasional fine punctures laterally, anterior edge with row of coarse umbilicate punctures laterally, surface finely alutaceous, sternite 6 simple and finely alutaceous, suture between stemites 5-6 deeply sulcate laterally. Pygidium, W/L = 2.0 in males or 2.2 in females, deeply sulcate along basal margin, sulcus wider medially, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 205-207). Tubus approximately 2.0 times as long as parameres, ventral portion sulcated on apical fourth. Parameres in lateral view elongate, with apex acute, ventral portion slightly concave, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 297.
MATERIAL ST UDIED. 5 a a, 8 9 9 (HAHC, ISNB, MNHN, MZSP, NMPC, SMTD, USNM, ZMUC).
TYPE MATERIAL Ontherus lichyi Martinez: Holotype a (AMIC): Venezuela, Estado de Carabobo, Valle del Rio Borburata, VI-V II.1941, Sr. Rene Lichy Leg. Holotype not studied (illustration provided in description sufficient).
Ontherus mirandai Pereira: 102 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. lichyi
+ 0. edentulus
MAP 21. Known distribution of Ontherus lichyi and 0. edentulus.
Holotype 9 (MZSP): (Brazil) Amazonas, Rio Negro, 1915/P. Pereira/ TIPO/Ontherus mirandai, sp. n., P. Pereira det. 948/ 16330/ Ontherus lichyi, Martinez, 1947, Det. F. Genier, 1992. Holotype studied.
NON-TYPE MATERIAL
Distribution: BRAZIL: Rio de Janeiro: Rio de Janeiro. Santa Catarina: Joinville. - COLOMBIA: Magdalena: Campana, 25km S Santa Marta (914m); Santa Marta Mts. Meta: Villavicencio. Norte de Santander: Ocafia. FRENCH GUIANA: No locality. VENEZUELA: Distrito Federal: Caracas.
Additional data: Specimens collected in January, April, May, July.
REMARKS. The holotype of 0. mirandai is an abraded female. I have decided to leave 0. mirandai in synonymy with 0. lichyi because this specimen falls within the known distribution range of 0. lichyi. Some variation occurs in the shape of the posterior femur in males. At this point, I will tentatively consider these variations as intraspecific. However, the status of 0. lichyi will have to be reconsidered when more material becomes available.
41. Ontherus (Ontherus) edentulus sp.nov. (Figs. 82, 208-210, 298; Map 21)
ETYMOLOGY. Edentulus, (without+ tooth) Latin adjective, refers to absence of fourth protibial tooth in males.
DESCRIPTION. Colour. Body reddish brown to black. Pubescence fulvous. Length. 13-18 mm. Head. Anterior edge arcuate, moderately upturned, slightly bisinuate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or on anterior three-fourths in females, remainder of surface punctate. Clypeofrontal suture A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 103 subquadrate, subtrapezoidal in smaller individuals, strongly carinate, carina tuberculate anterolaterally, slightly higher and sinuate medially, much lower but distinct laterally. Anterior genal edge much shorter than lateral, genal surfacemoderately to strongly wrinkled on anterior half, remainder of surface finely and shallowly punctate. Vertex finely punctate in some individuals to minutely punctate, except for small area posteriorly. Pronotum.
Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin of most individuals shortly enlarged medially. Disc strongly convex, basal half longitudinally and shallowly depressed medially; punctures minute on disc, fine on anterior and lateral declivities, with few umbilicate punctures along base. Elytra.
Subquadrate, L/W = 1.0. Striae moderately wide, similarly impressed on disc and apical declivity, punctures rounded, slightly deeper than stria on disc, deeper on apical declivity, slightly encroaching on intervals, approximately 2.0 times as wide as stria and separated by 3-5 diameters on disc and apical declivity. Intervals convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent and finelyalutaceous throughout. Prosternum behindprocoxa densely pubescent and with setae subequal in length and thickness throughout. Mesosternum shallowly and transversely impressed on disc, punctures irregular and dense anteriorly, lacking on posterior fourth, pubescence fine on anterior third, stouter on median third and lacking posteriorly. Mesepisternum shallowly and rather scarcely punctate, surface alutaceous, carinate parallel to posterior edge. Metasternum with disc minutely punctate and moderately alutaceous throughout, with deep longitudinal impression on posterior two thirds, impression wider and slightly deeper in males; lateral lobes on anterior half with punctures oval, anteriormost punctures each with anteromedial edge sharply carinate, posterior half with punctures broadly oval to rounded and mostly restricted to metacoxal edge, leaving scarcely punctate area internally, surfacebetween punctures alutaceous, except for small glossy area along metacoxal edge internally,pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior half; median lobe moderately wide, L/mW = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded internally, surface lacking punctures and strongly alutaceous, junction with mesostemum strongly angulate medially, sharply carinatethroughout its width, produced into short carina longitudinally on mesostemum. Legs. Anterior tibia tridentate in males or quadridentate in females, ventromedial carina lacking intervening setae on basal half in males or third in females, apical spur straight and parallel-sided in males or obliquely truncate and slightly spatulate apically in females. Posterior femur (Fig. 82) triangularly produced on posterodorsal edge apically, outer edge of projection straight and ventrally recurved, edge straight and sharp in smallest individuals, ventroposterior sulcus sharply delimited throughout and extending anteriorly at apex in males or femur unmodified in females. Posterior tibia (Fig. 82) unmodified in both sexes. Abdomen. Sternites 2-5 with occasional fine punctures laterally, anterior edge with row of coarse umbilicate punctures laterally, surface finely alutaceous, stemite 6 glossy, suture between stemites 5-6 deeply sulcate laterally. Pygidium, WIL = 2.1 in males or 2.2 in females, deeply sulcate along basal margin, sulcus wider medially, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 208-210). Tubus approximately 2.0 times as long as parameres, ventral portion sulcated on apical fourth. Parameres in lateral view elongate, with apex acute, ventral portion slightly concave, produced into small tooth just before the apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 298.
MATERIAL STUDIED. 13 a a, 9 Q Q (BMNH, DEIC, HAHC, IMLA, LACM, MNHN, NMPC, PAIC, SMTD, WBWC, WDEC). 104 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Holotype O' (MNHN): Perou, Chanchamayo, Ch. 0. Schunke/ Museum Paris, ex. Coll., R. Oberthur/ HOLOTYPE, Ontherus edentulus, F. Genier, 1992. Dissected.
Allotype Q (BMNH): Chanchamayo, Peru/ Nevinson Coll., 1918-14/ ALLOTYPE, Ontherus edentulus, F. Genier, 1992.
Paratypes: ECUADOR: Loja: Loja (1 Q MNHN). Morona-Santiago: Macas (1 O' MNHN). Zamora-Chinchipe: Zamora, Prof. C. Carrion, B. M. 1928-69 (1 Q BMNH). PERU: No locality (1 O' NMPC); Slid-Peru (l Q ISNB). Amazonas: Huambo, ex Stgr. (1 O' MNHN). Cuzco: Callauga (=Callanga) (1 O' SMTD); Ocobamba (1 O' DEIC); Vilcanota (=Cordillera de?) (1 O' SMTD). Huanuco: Tingo Maria (Environs de) (1 O' PAIC). Junin: No locality (1 O' WBWC); Chanchamajo (=Chamchamayo) (l Q SMFD); La Merced (1 O' LACM); Sani Beni*, 4.V III.1935 (1 Q WDEC); Valle Chanchamayo (800m), 1.11.1939, leg. Weyrauch (3 O'O', 2 Q Q IMLA). Pasco: Lobo* (P. N. Ya nachaga) (400m), 26.IX.1993, Pedro Hocking (1 O' HAHC); Pozuzu (=Pozuzo) (1 Q SMTD).
42. Ontherus (Ontherus) brevipennis Harold, 1867 comb.nov. (Figs. 83, 211-213, 299; Map 22)
Ontherus brevipennis Harold, 1867: 97 (description) Ontherus brevipennis Harold: Gemminger and Harold, 1869: 1008 (catalog) Ontherus brevipennis Harold: Harold, 1880: 23 (comment) Ontherus brevipennis Harold: Bates, 1887: 51 (distribution) Ontherus brevipennis Harold: Gillet, 191 lb: 57 (catalog) 0 ntherus brevipennis Harold: Luederwaldt, 1931: 393 (key) Ontherus brevipennis Harold: Blackwelder, 1944: 206 (list) Ontherus brevipennis Harold: Martinez, 1947: 48 (comment) 0 ntherus brevipennis Harold: Howden and Young, 1981: 121, 122 (key, redescription and distribution) Ontherus brevipennis Harold: Gill in Hanski and Cambefort, 1991: 225, 412 (biology)
REDESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 10-16 mm. Head. Anterior edge arcuate, moderately upturned, almost straight medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third in males or on anterior third to two-thirds in females, with feeble to distinct bulge parallel to anterior edge on anterior third in males, remainder of surface moderately to strongly punctate. Clypeofrontal suture subquadrate, trapezoidal in smaller individuals, strongly carinate, carina denticulate anterolaterally, straight medially, ill-defined laterally. Anterior genal edge much shorter than lateral, genal surface moderately to strongly wrinkled on anterior half, remainder of surface finely and shallowly punctate. Vertex rather coarsely punctate in some individuals to finely punctate, except for small area posteriorly. Pronotum. Transverse, W /L = 1.4-1.6; strongly convex, wider than elytra, except in smallest individu als; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin strongly enlarged behind head insertion. Disc strongly convex, median third with angulate bulge on anterior fourth, dorsal surface of anterior process flat, or moderately to strongly sulcate medially in individuals from Costa Rica, basal half lacking distinct longitudinal depression medially; punctures fine on disc, minute in some individuals, distinctly larger on dorsal portion of anterior process, anterior and lateral declivities, with row of moderate to coarse and umbilicate punctures along base. Elytra. Transverse, L/W = 0.9. Striae moder ately wide, similarly impressed on disc and apical declivity, punctures rounded, slightly deeper than stria on disc, slightly encroaching on intervals, approximately 2.0 times as wide as stria and separated by 2- 3 diameters on disc and apical declivity. Intervals convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion moderately pubescent and finely alutaceous throughout. Prostemum behind procoxa rather scarcely pubescent and with setae suequal in length and thickness throughout. Mesostemum shallowly and transversely impressed on disc, punctures irregular A REVISION OF THE NEOTROP!CAL GENUS ONTHERUS ERICHSON 105
• 0. brevipennis
MAP 22. Known distribution of Ontherus brevipennis.
and dense anteriorly, feebly to moderately impressed on posterior half, pubescence fine and apposed on anterior third, stouter and erect on median third and minute posteriorly. Mesepistemum rather coarsely and irregularly punctate, surfacealutaceous, sharply carinate parallel to posterior edge. Metastemum with disc minutely punctate, surface between punctures glossy, with deep longitudinal impression on posterior two-thirds, impression wider and slightly deeper in males; lateral lobes on anterior half with punctures oval, anteriormost punctures each with anteromedial edge bluntly carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for glossy area along metacoxal edge internally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior two-thirds; median lobe moderately wide, L/mW = 2.3, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded to subangular internally, surface lacking punctures and finely alutaceous, junction with mesostemum strongly angulate medially, carinate throughout its width, produced into short carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth slender and projecting anteriorly in males, ventromedial carina lacking intervening setae on basal third in males or fourth in females, apical spur straight and parallel-sided in males or obliquely truncate and slightly spatulate apically in females. Posterior femur (Fig. 83) denticulate on posteroventral edge apically, edge straight in smallest individuals, ventroposterior sulcus sharply delimited except on basal fifth, gradually narrowing on apical fourth toward apex, not extending anteriorly at apex in males or femur unmodified in females. Posterior tibia (Fig. 83) flattened ventrally on basal third, inner edge lacking indentation on basal third in males or unmodified in females and smaller males. Abdomen. Stemites 2-5 with few fine punctures laterally, anterior edge with row of irregular punctures laterally, surface finely alutaceous, sternite6 with few minute punctures, surface between punctures glossy, suture between stemites 5-6 deeply sulcate laterally. 106 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Pygidium, W IL = 2.1 in males or 2.3 in females, deeply sulcate along basal margin, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 211-213). Tubus approximately 2.0 times as long as parameres, ventral portion slightly depressed apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly concave, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 299.
MATERIAL STUDIED. 42aa, 63 9 9 (AVEC, CNCI, HAHC, INBC, ISNB, MCZC, MNHN, NMPC, SM TD, TAMU, UNSM).
TYPE MATERIAL Lectotype a (MNHN): Nov. Granad./ Brevipennis, t. Harold/ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthtir, 1952/ LEC TOTYPE, Ontherus brevipennis, Harold, 1867, Des. F. Genier, 1991. Type material studied and lectotype here designated.
NON-TYPE MATERIAL Distribution: COLOMBIA: Hac[ienda]. Pehlke*. Magdalena: Rio Frio; Santa Martha. Valle del Cauca: Darien. COSTA RICA: Caullaga*. Cartago: Cartago. Guanacaste: Estancia Pitilla, 9km S Santa Cecilia (700m). Puntarenas: Quepos, Parque Nacional Manuel Antonio (80m); Rancho Quemado, Peninsula de Osa (200m). PANAMA: Chiriqul: Chiriqui. Colon: Canal Area, B.C.I. (Barro Colorado Island); Canal Zone, Forest Reserve; Canal Zone, Base of Cerro Galera; Portobelo; Santa Rita Ridge (300m). PERU: No locality.
Additional data: Some specimens from Costa Rica and Panama were collected from leafcutter ant (Atta sp.) refuse piles. Specimens collected in February, April to July, September, November, December.
REMARKS. Specimens collected in Costa Rica differslightly from those from Panama and Colombia as follows: overall body shape stouter, especially pronotum; cephalic tubercles slightly projecting laterally (straight in other specimens); pronotal punctures minute on most of the surface; dorsal portion of the anterior process sulcated anteriorly; male posterior femora with denticulate process proportionally smaller. Male genitalia of a representative sample were dissected and except for a slight, but consistent difference in the shape of the sigmoid sclerite, no other significant differences could be found. I consider this variation as intraspecific.
43. Ontherus (Ontherus) rectangulidens sp.nov. (Figs. 84, 214-216, 300; Map 20)
ETYMOLOGY. Rectangulidens, (rectangular+ tooth) Latin adjective, refers to shape of apical tooth of anterior tibiae in males.
DESCRIP TION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 11-15 mm. Head. Anterior edge arcuate, moderately upturned, straight medially, cly peo genal junction straight. Clypeus transversely wrinkled on anterior third in males or on anterior third to two-thirds in females, lacking distinct bulge parallel to anterior edge on anterior third in males, remainder of surface finely and shallowly punctate. Clypeofrontal suture subquadrate, anterolateral angles rounded in smaller individuals, strongly carinate, carina straight, weak laterally. Anterior genal edge much shorter than lateral, genal surface moderately to strongly wrinkled on anterior half, remainder of surface finely and shallowly punctate. Vertex rather coarsely punctate, except for small area posteriorly. Pronotum.
Transverse, W/L = 1.5-1.7; strongly convex, wider than elytra, except in smallest individu als; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 107 slightly enlarged behind head insertion. Disc simple and strongly convex, basal half with indistinct to distinct longitudinal depression medially; punctures minute on disc, fine on anterior declivity, with row of moderate to coarse and umbilicate punctures along base.
Elytra. Transverse, L/W = 0.9. Striae moderately wide, coarsely microsculptured, similarly impressed on disc and apical declivity, punctures rounded, slightly deeper than stria on disc, encroaching on intervals, approximately 2.0 times as wide as stria and separated by 1-3 diameters on disc and apical declivity. Intervals strongly convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion with fewsetae basally and finely alutaceous throughout. Prosternumbehind procoxa completely lacking setae ventrally, surface alutaceous throughout. Mesosternumalmost flat, punctures scarce and shallowly impressed, pubescence minute and apposed. Mesepisternum shallowly and irregularly punctate, surface alutaceous, finely carinate parallel to posterior edge. Metasternum with disc glossy, with deep longitudinal impression on posterior two thirds, surface of depression finely alutaceous, impression wider and slightly deeper in males; lateral lobes on anterior half with punctures oval, each puncture with anteromedial edge carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for glossy area along metacoxal edge internally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior two-thirds; median lobe moderately wide, L/mW = 2.3, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins subangular internally, surface lacking punctures and finely alutaceous, junction with mesosternum strongly angulate medially, carinate throughout its width, produced into short ill-defined carina longitudinally on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth obliquely oriented, shorter than penultimate, parallel-sided and truncate apically in males, ventromedial carina lacking intervening setae on basal three-fourths in males or half in females, apical spur straight and parallel-sided in males and females. Posterior femur (Fig. 84) triangularly produced on posterodorsal edge apically, posteroventral edge always complete, sharply carinate throughout, ventroposterior sulcus sharply delimited posteriorly and apically, rounded apically in males or femur unmodified in females. Posterior tibia (Fig. 84) slender, inner edge finely carinate on basal fourth, distal portion of carina finely denticulate in males or unmodified in females and smaller males. Abdomen. Sternites 2-5 lacking punctures laterally, except anterior edge with row of coarse and irregular punctures laterally, surface finely alutaceous, sternite 6 with few minute punctures, surface between punctures glossy, suture between sternites 5-6 deeply sulcate laterally. Pygidium, WIL = 2.0 in males or 2.3 in females, rather shallowly sulcate along basal margin, surface finely punctate on disc, surface between punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 214-216). Tubus approximately 2.4 times as long as parameres, ventral portion narrowly depressed apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly concave, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 300.
MATERIAL STUDIED. 5 a a, 6 9 9 (ISNB, MCZC, MNHN, MZSP, ZMHB).
Holotype O' (MCZC): Itatiaya Mts., S. E. Brazil, X./ H. C. Fall, Collection/ Ontherus zikani Ltiderw., P. Pereira det. 958/ HOLOTYPE, Ontherus rectangulidens, F. Genier, 1992. Dissected.
Allotype 9 (MCZC): Same data as holotype. Dissected. 108 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Paratypes: BRAZIL: No locality (1 9 MNHN). Rio de Janeiro: Itatiaya, F. Ohaus, 10.XII.[19]26 (1 9 ZMHB), same as before except: 3.I.[19]27 (1 CJ' ZMHB), Itatiya (sic), 700m, 1.XI.1925, J. F. Zikan, Cotypo Ontherus zikani Luederw. det. 29, # 16292 (1 CJ' MZSP), Itatiaya, 700m, 30.VII.[19]22, W. Zikan (1 a ISNB), Itatiaya Mts., X., with quern quam ants, H. C. Fall Collection (2 9 9 MCZC); Mendes, 28.X.1933, Dr. Eidmann legit, Museum Paris, 1936, Coll. Boucomont ( 1 CJ', 1 9 MNHN).
Additional data: One specimen collected from a nest of" quern quam" ants.
44. Ontherus (Ontherus) carinicollis Luederwaldt, 1931 stat.nov. (Figs. 6, 85, 217-219, 301; Map 23)
Ontherus carinicollis Luederwaldt, 1930: 108 (nomen nudum) (comment) Ontherus zikani var. carinicollis Luederwaldt, 1931: 375 (description and distribution) Ontherus zikani var. carinicollis Luederwaldt: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark reddish brown to black, some individuals with strong iridescent sheen on elytra. Pubescence fulvous. Length. 12-16 mm. Head. Anterior edge arcuate, almost flat, straight medially, clypeogenal junction straight. Clypeus trans versely wrinkled on anterior third in both sexes, lacking distinct bulge parallel to anterior edge on anterior third in males, remainder of surface finely and shallowly punctate. Clypeofrontal suture quadrate, subtrapezoidal in smallest individuals, anterolateral angles subdenticulate, anterior edge strongly carinate, carina straight, weak laterally. Anterior genal edge slightly shorter than lateral, genal surface shallowly wrinkled on anterior third, remainder of surface finelyand shallowly punctate. Vertex rather coarsely punctate medially.
Pronotum. Transverse, W/L = 1.7; strongly convex, wider than elytra, except in smallest individuals; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin slightly enlarged behind head insertion, even in thickness. Disc strongly convex, with semicircular bulge anteromedially, basal third with vaguely delimited to distinct longitudinal depression medially; punctures minute, usually distinctly larger on dorsal portion of anterior process, with row of unaligned coarse and umbilicate punctures along base. Elytra.
Subquadrate L/W = 1.0. Striae narrow, coarsely microsculptured, similarly impressed on disc and apical declivity, punctures rounded, slightly deeper than stria on disc, encroaching on intervals, approximately 3.0 times as wide as stria and separated by 1-3 diameters on disc and apical declivity. Intervals strongly convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent, setae mostly restricted to posterior portion, surface finely alutaceous throughout. Prostemum behind procoxa completely bare ventrally, surface alutaceous throughout. Mesostemum almost flat, punctures scarce and shallowly impressed, with, at most, IO short setae medially. Mesepisternum coarsely and irregularly punctate, surface alutaceous, finely carinate parallel to posterior edge. Metasternum with disc glossy, with deep longitudinal impression on most of its length in males or shallow impression on posterior third in females, surface of depression finely alutaceous; lateral lobes on anterior half with punctures oval, each puncture with anteromedial edge carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for glossy area along metacoxal edge internally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior half; median lobe moderately wide, L/mW = 2.0, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins angulate internally, surface minutely punctate and alutaceous, junction with mesostemum strongly angulate medially, carinate throughout its width, produced into short carina longitudinally on mesosternum. Legs. Anterior tibiaquadridentate in both sexes, apical tooth pointing forward, A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 109
• 0. carinicollis
MAP 23. Known distribution of Ontherus carinicollis.
feebly to moderately lobate externally in males or unmodified in females, ventromedial carina lacking intervening setae on basal half in males or basal third in females, apical spur straight, parallel-sided and acute apically in males or subspatulate and obliquely truncate in females. Anterior femur with small to moderate conical process on anterior third. Posterior femur (Fig. 85) triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on same plane, forming large depression in larger males, in smaller males both surfaces partially to completely separated by posteroventral edge, ventroposterior sulcus sharply delimited anteriorly and apically, apical portion of posteroventral sulcus distinctly extending anteriorly in males or femur unmodified in females. Posterior tibia (Fig. 85) slender, inner edge finely carinate on basal third, distal portion of carina finely denticulate in males or unmodified in females and smaller males. Tarsal claws (Fig. 6), when fully extended, at least one-third as long as last tarsal segment. Abdomen. Stemites 2-5 lacking punctures laterally, except anterior edge with row of coarse irregular punctures laterally, surface finely alutaceous, stemite 6 minutely punctate through out, surface between punctures glossy, suture between stemites 5-6 deeply sulcate through out. Pygidium, W/L = 2.1 in males or 2.4 in females, rather shallowly sulcate along basal margin, surface minutely punctate, surface between punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 217-219). Tubus approximately 2.4 times as long as parameres, ventral portion slightly emarginate apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into small tooth just before apex, ventral edge of right paramere widely and triangu larly membranous, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 301.
MATERIAL STUDIED. 9 a a, 12 9 9 (CNCI, HAHC, ICCM, ISNB, MACN, MCZC, MNHN, MZSP, NMPC, USNM, WDEC). 110 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
TYPE MATERIAL Lectotype O' (MZSP): Sao Paulo, Cidade, 1900 (recto), comps. de Staudinger verso)/ 0. cephalot./ Ontherus zikani var., carinicoll. Lueder., Lueder: det. 29/ blue disk/ COTYPO (pink paper)/ 16369/ LECTOTYPE, Ontherus zikani, var. carinicollis, Luederwaldt, 1931, Des. F. Genier, 1992. Dissected. Ty pe material studied, Lectotype and paralectotypes here designated.
Paralectotypes: Minas Geraes, P[ouso]. Alegre, 4.832/ 0. cephalot./ Ontherus zikani var., carinicoll. Lueder., Luederw. det. 29/ blue disk/ COTYPO (red paper)/ 16371/ PARALECTOTYPE, Ontherus zikani, var. carinicollis, Luederwaldt, 1931, Des. F. Genier, 1992 (1 Q MZSP). Est., Minas Geraes, 4.832/ 0. cephalot./ Ontherus zikani var., carinic. Lueder., Luederw. det. 29/ blue disk/ COTYPO (pink paper)/ 16370/ PARALECTOTYPE, Ontherus zikani, var. carinicollis, Luederwaldt, 1931, Des. F. Genier, 1992 (1 Q MZSP).
NON-TYPE MATERIAL Distribution: ARGENTINA: Misiones: No locality. BRAZIL: Distrito Federal: Estacao Florestal, Cabeca do Veado*; Parque Nacional* (de Brasflia). Mato Grosso: Chapada dos Guimaraes. Mato Grosso do Sul: Campo Grande. Minas Geraes: Pouso Alegre; Uberaba. Sao Paulo: Fazenda Campininhas, Mogi Guassu; Sao Paulo. PARAGUAY: Caaguazu: Estencia Primera*. Concepcion: Horqueta.
Additional data: Some specimens were collected at light. Specimens collected in January, February, September, October, December.
REMARKS. This is a valid species which was described by Luederwaldt as a variety of 0. zilwni. It can be separated from the former by the overall stouter aspect; the subquadrate and subdenticulate cephalic carina; the presence of a distinct semicircular bulge anterome dially on the pronotum; the more rounded and larger strial punctures; and the larger tarsal claws (Fig. 6). The male genitalia are similar to the following species. However, characters of the male genitalia are not diagnostic in this species group.
45. Ontherus (Ontherus) zikani Luederwaldt, 1930 comb.nov. (Figs. 7, 86, 220-222, 302; Map 24)
Ontherus zikani Luederwaldt, 1930: 107 (description and distribution) Ontherus zikani Luederwaldt: Luederwaldt, 1931: 370, 374 (key, distribution and redescription) Ontherus zikani Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus zikani Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus zikani Luederwaldt: Martinez, 1947: 48 (comment) Ontherus zikani Luederwaldt: Pereira, 1955: 462 (comment) Ontherus zikani Luederwaldt: Martinez, 1959: 72 (catalog)
REDESCRIPTION. Colour. Body dark reddish brown to black, some individuals with slight iridescent sheen on elytra. Pubescence fulvous. Length. 11-15 mm. Head. Anterior edge arcuate, almost flat, straight medially, clypeogenal junction straight. Clypeus trans versely wrinkled on anterior third in both sexes, lacking distinct bulge parallel to anterior edge on anterior third in males, remainder of surface finely and shallowly punctate. Clypeofrontal suture subtrapezoidal, anterolateral angles rounded, subdenticulate in largest (15 mm) males, anterior edge strongly carinate, carina even in height and straight medially, low but distinct laterally. Anterior genal edge shorter than lateral, genal surface shallowly to modernly wrinkled on anterior fourth, remainder of surface finely and shallowly punctate.
Vertex rather coarsely punctate medially. Pronotum. Transverse, W/L = 1.6; strongly convex, wider than elytra, except in smallest individuals; anterior edge narrowly membra nous and slightly arcuate behind head insertion, margin slightly enlarged behind head A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 111 insertion, even in thickness, widened medially in some individuals. Disc lacking distinct swelling anteromedially, basal third with vagely delimited to distinct longitudinal depression medially; punctures minute, slightly larger on anterior angles and along anterior edge in some individuals, with row of unaligned moderate to coarse and umbilicate punctures along base. Elytra. Subquadrate L/W = 1.0. Striae narrow, finely microsculptured, similarly impressed on disc and apical declivity, punctures oval, usually umbilicate, slightly deeper than stria on disc, slightly encroaching on intervals, approximately 2.0 times as wide as stria and separated by 3-5 diameters on disc and apical declivity. Intervals strongly convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepister num with posterolateral portion scarcely pubescent, setae mostly restricted to posterior portion, surface finely alutaceous throughout. Prosternum behind procoxa completely bare ventrally, surface alutaceous throughout. Mesosternum almost flat, punctures rather dense medially, and moderately impressed, moderately pubescent, setae fine and short, semi-erect. Mesepisternum moderately and irregularly punctate, surface alutaceous, finely carinate parallel to posterior edge. Metasternum with disc glossy, with deep longitudinal impression on most of its length in males or shallow impression on posterior third in females, surface of depression finely alutaceous; lateral lobes on anterior half with punctures oval, anteriormost punctures with anteromedial edge finely carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for glossy area along metacoxal edge inter nally, pubescence obliquely oriented, scarce, moderately long along mesocoxal edge, slightly decreasing in length toward lateral edge, lacking on posterior half; median
lobe moderately wide, L/mW = 2.1, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins angulate internally, surface minutely punctate and alutaceous, junction with mesostemum strongly angulate medially, carinate throughout its width, produced into short carina longitudinally on mesostemum. Legs. Anterior tibia quadridentate in both sexes, apical tooth pointing forward, carinate to moderately lobate externally in males or unmodified in females, ventromedial carina lacking intervening setae on basal half in males or third in females, apical spur straight, parallel-sided and acute apically in males or subspatulate and obliquely truncate in females. Anterior femur with small to moderate conical process on anterior third. Posterior femur (Fig. 86) triangularly produced on posterodorsal edge apically, posteroventral sulcus and posterior surface fused and almost on same plane, forming large depression in larger males, in smaller males both surfaces partially to completely separated by posteroventral edge, ventroposterior sulcus sharply delimited anteriorly and apically, apical portion of pos teroventral sulcus distinctly extending anteriorly in males or femur unmodified in females. Posterior tibia (Fig. 86) slender, inner edge finely carinate on basal third, distal portion of carina finely denticulatein males or unmodifiedin femalesand smaller males. Tarsal claws, when fully extended, at most one-fourth as long as last tarsal segment. Abdomen. Stemites 2-5 with at most few shallow punctures laterally, except foranterior edge with row of coarse irregular punctures laterally, surface finely alutaceous, stemite 6 minutely punctate through out, surface between punctures glossy, suture between stemites 5-6 deeply sulcate through
out. Pygidium, W/L = 2.1 in males or 2.3-2.6 in females, rather shallowly sulcate along basal margin, surface minutely punctate, surface between punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 220-222). Tubus approximately 2.4 times as long as parameres, ventral portion slightly emarginate apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 302.
MATERIAL STUDIED. 8 a a, 17 9 9 (BDGC, ICCM, IMLA, ISNB, MACN, MNHN, MZSP, NMPC, SEMC, WDEC, ZMHB). 112 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. zikani
MAP 24. Known distribution of Ontherus zikani.
TYPE MATERIAL Lectotype ct (MZSP): Itatiaya, llOOm, Est. do Rio, 24-1-1929, Maroniba, J. F. Zikan/ Ontherus zikani Lueder., Luederw. det. 29/ blue disk/ COTYPO (pink paper)/ 16288/ LECTOTYPE, Ontherus zikani, Luederwaldt, 1930, Des. F. Genier, 1992. Dissected. Type material studied, lectotype and paralectotypes here designated.
Paralectotypes: Same data as lectotype except: 3-XII-1924, 700m,/ 16290/ (1 Q MZSP); 12.I.1926/ 16291 (1 Q MZSP).
NON- TYPE MATERIAL Distribution: ARGENTINA: Misiones: Puerto Bemberg; Dos de Mayo; Eldorado; Parque Nacional Iguazu, Pto. Canoas* (180m); San Ignacio. BRAZIL: Bahia: No locality. Minas Gerais: Aguas Vermelhas; Pouso Alegre. Rio de Janeiro: Itatiaia; Itatiaia Mts. Rio Grande do Sul: Porto Alegre. Santa Catarina: Mafra. Sao Paulo: Ipiranga.
Additional data: Ontherus zikani lives with ants (Luederwaldt 1930: 108). One specimen was collected from an unspecified carrion trap, another using a flight intercept trap. Specimens collected in January, October to December.
REMARKS. Five specimens labelled as cotypes were sent to me by U. Martins (MZSP). One of the specimens was from a locality (Ypiranga) not cited by Luederwaldt and, therefore, was removed from the paralectotype series. Another specimen belongs to a new species ( 0. rectangulidens) and is now part of the paratype series of that species.
46. Ontherus (Ontherus) lobifrons sp.nov. (Figs. 8, 9, 87, 223-225, 303; Map 25)
Ontherus cephalotes Harold: sensu Luederwaldt, 1931: 370, 376 (key, distribution and redescription) Ontherus cephalotes Harold: sensu Balthasar, 1938: 222, 223 (comment) Ontherus cephalotes Harold: sensu Pessoa and Lane, 1941: 456 (key) A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 113
ETYMOLOGY. Lobifrons, (lobe + frons) Latin adjective, refers to lobate aspect of the cephalic carina.
DESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 11-17 mm. Head. Anterior edge arcuate, almost flat,straight medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third in males or half in females, lacking distinct bulge parallel to anterior edge on anterior third in males, remainder of surface finely and shallowly punctate. Clypeofrontal suture arcuate, occasionally subtrapezoidal in some smaller individuals, anterior edge strongly carinate, carina higher medially, at most slightly emarginate medially, gradually lowering posterolaterally. Anterior genal edge shorter than lateral, genal surface shallowly wrinkled on anterior fourth, remainder of surface finely and shallowly punctate. Vertex rather coarsely punctate medially. Pronotum. Transverse,
W/L = 1.6; strongly convex, wider than elytra, except in smallest individuals; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin slightly enlarged behind head insertion, almost even in thickness, widened medially in some individuals. Disc lacking distinct swelling anteromedially, basal third with vaguely delimited to distinct longitudinal depression medially; punctures minute, slightly larger on anterior angles and along anterior edge in some individuals, with row of moderate to coarse and umbilicate punctures along base. Elytra. Subquadrate L/W = 1.0. Striae moderately wide, coarsely microsculptured, similarly impressed on disc and apical declivity, punctures rounded, usually finely umbilicate, slightly deeper than stria on disc, slightly encroaching on intervals, approximately 1.5-2.0 times as wide as stria and separated by 1-3 diameters on disc and apical declivity. Intervals moderately convex, surface glossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion with few scarce setae posteriorly, surface finely alutaceous throughout. Prosternum behind procoxa com pletely bare ventrally, surface alutaceous throughout. Mesosternumstrongly and transversely depressed on anterior half, less so in females and smaller individuals, punctures large, coarse and dense on anterior half, posterior half usually produced into large conical process, longitudinally carinate ventrally, process reduced to strong longitudinal carina in females and some males, anterior half finely pubescent, setae short and apposed, surface on posterior half strongly alutaceous. Mesepisternum weakly punctate, surface strongly alutaceous, finely carinate parallel to posterior edge. Metasternum with disc glossy, with deep longitu dinal impression on most of its length in males or narrow and shallow impression on posterior half in females, surface of depression finely alutaceous; lateral lobes on anterior half with punctures oval and shallowly impressed, scarce internally, anteriormost punctures with anteromedial edge bluntly carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for rather glossy area along metacoxal edge internally, pubescence completely missing; median lobe (Figs. 8, 9) moderately wide, L/mW = 2.2, lateral margins slightly divergent on anterior third, convergent on posterior two-thirds, anterior angles of lateral margins rounded internally, surface minutely punctate and alutaceous, junction with mesosternum strongly angulate medially, carinate throughout its width, produced into long carina longitudinally on mesosternum, carina usually longer than half the minimum width of median lobe of mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth pointing forward, slightly to moderately lobate externally in males or unmodifiedin females, ventromedial carina lacking intervening setae on basal two-thirds in males or half in females, apical spur straight, parallel-sided and acute apically in males or subspatulate and obliquely truncate in females. Anterior femur unmodified. Posterior femur (Fig. 87) triangularly produced on posterodorsal edge apically, posteroventral edge dentate apically, posteroventral sulcus and posterior surface always clearly separated by posteroventral edge, never forming large depression, ventroposterior sulcus sharply delimited throughout, apical portion of posteroventral sulcus tapering toward apex, not extending anteriorly in males or femur 114 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
• 0. lobifrons
MAP 25. Known distribution of Ontherus lobifrons.
unmodified in females and smaller males. Posterior tibia (Fig. 87) slender, inner edge sublo bate on median third in larger males or unmodifiedin females and smaller males. Tarsal claws, when fully extended, at most one-fourth as long as last tarsal segment. Abdomen. Sternites 2-5 with at most few shallow punctures laterally, except foranterior edge with row of rather small and coarse irregular punctures laterally, surface .finely alutaceous, sternite 6 glossy, suture between stemites 5-6 deeply sulcate throughout. Pygidium, W/L = 2.2 in males or 2.4 in females, rather deeply sulcate along basal margin, surface minutelypunctate, surface between punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 223-225). Tubus approximately 2.3 times as long as parameres, ventral portion slightly depressed apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 303.
MATERIAL STUDIED. 25 O' O', 17 Q Q (AMNH, CNCI, DEIC, ISNB, MLPA, MNHN, NMPC, PAIC, SMTD, USNM, WDEC, ZMHB, ZSMC).
Holotype O' (NMPC): Amler (?), I-1926/ HOLOTYPE, Ontherus lobifrons, F.Genier, 1992. · Aedeagus removed.
Allotype Q (NMPC): Same data as holotype except: XII-1925.
Paratypes: Caulajalla* (20' O' SMTD). BRAZIL: No locality (2 O' O' MNHN) (1 O' ZMHB) (1 Q ZSMC); Artigueira*, VI.1946, F. Justus Jr. (20' O' PAIC); Brasilia merid. (1 O' NMPC). Parami: No locality (1 Q MNHN); Curitiba (1 O' MLPA). Rio de Janeiro: Rio de Janeiro (40' O' SMTD). Rio Grande do Sul: No locality (1 Q ZMHB); Nova Petropolis, I.[19]28, Buck (1 Q ISNB); Porto Alegre (1 Q ZMHB). Sao Paulo: No locality (1 O' ISNB). Santa Catarina: No locality (1 Q MNHN), same as before except: coll. J. Thomson, (1 O', 2 Q Q ISNB); Blumenau, Reitter (1 Q ISNB), same as before except: VI.[19]19, Luederw. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 115 leg. (1 O' ISNB); Joinvile (1 O', 1 9 SMTD), same as before except: XI (1 O' DEIC); Mafra (30' O' MNHN); Nova Teutonia, 29.IV.1940, Plaumann (1 O' USNM), same as before except: 27°11 'B. 52°23 'L., 4.8.1932, Fritz Plaumann (19 ISNB), 27°11 'B. 52°23 'L., 3.7.1934, Fritz Plaumann (1 9 ISNB), 27°11 'B. 52°23 'L., 4.7.1934, Fritz Plaumann (1 O', 1 9 ISNB), 27°11 'B. 52°23 'L., 300-500m, VI.1957, Fritz Plaumann, bei Aeromyrmex
(1 9 CNCI); Pinhal ( = Pinhalzinho?), XII.1947, A. Maller coll. (1 O' WDEC) (1 9 AMNH); Rio Vermelho, I.1945, A. Maller coll. ( 1 9 AMNH).
Additional data: One specimen was collected with ants (Aeromyrmex sp.).
REMARKS. In some males the mesostemal process is reduced to a longitudinal keel. As no other characters could be correlated with this particular state, this variation is considered to be intraspecific.
47. Ontherus (Ontherus) cephalotes Harold, 1869 comb.nov. (Figs. 10, 11, 88, 226-228, 304; Map 26)
Ontherus cephalotes Harold: Gemminger and Harold, 1869: 1008 (nomen nudum)(catalog) Ontherus cephalotes Harold, 1869h: 98 (description) Ontherus cephalotes Harold: Gillet, 191 lh: 57 (catalog) Ontherus cephalotes Harold: Luederwaldt, 1930: 105, 107 (comment) Ontherus quadrituherculatus Luederwaldt 1930: 107 (description) new synonymy. Ontherus cephalotes Harold: Luederwaldt, 1931: 365, 370, 376 (biology, key, distribution and redescription) Ontherus quadrituherculatus Luederwaldt: Luederwaldt, 1931: 370, 377 (key, distribution and redescription) Ontherus cephalotus (sic) Harold: Luederwaldt, 1931; 391 (comment) Ontherus cephalotes Harold: Balthasar, 1938: 222, 223 (comment) Ontherus quadrituherculatus Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus cephalotes Harold: Pessoa and Lane, 1941: 456, 457 (key, redescription and distribution) Ontherus cephalotes Harold: Blackwelder, 1944: 206 (list) Ontherus quadrituberculatus Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus cephalotes Harold: Lange, 1947: 310 (list) Ontherus quadrituberculatus Luederwaldt: Lange, 1947: 310 (list) Ontherus cephalotes Harold: Pereira, 1955: 462 (comment) Ontherus cephalotes Harold: Gill in Hanski and Cambefort, 1991: 216 (biology)
REDESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 11-14 mm. Head. Anterior edge arcuate, almost flat, almost straight medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third in males or half in females, lacking distinct bulge parallel to anterior edge on anterior third in males, remainder of surface finely and shallowly punctate. Clypeofrontal suture semicircular, anterior edge strongly carinate, carina feebly to distinctly bilobate medially, posterolateral portion of carina subtuberculate. Anterior genal edge shorter than lateral, genal surface shallowly to moderately wrinkled on anterior fourth, remainder of surface finely and shal lowly punctate. Vertex rather coarsely punctate medially. Pronotum. Transverse, W /L = 1.6; strongly convex, wider than elytra, except in smallest individuals; anterior edge narrowly membranous and slightly arcuate behind head insertion, margin slightly enlarged behind head insertion, almost even in thickness, widened medially in some individuals. Disc lacking distinct swelling anteromedially, basal third with vaguely delimited to distinct longitudinal depression medially; punctures minute, slightly larger on anterior angles and along anterior edge in some individuals, with row of moderate to coarse and umbilicate punctures along base. Elytra. Subquadrate L/W = 1.0. Striae wide, coarsely microsculptured, similarly impressed on disc and apical declivity, punctures rounded, usually finelyumbilicate, slightly deeper than stria on disc, slightly encroaching on intervals, approximately 1.5-2.0 times as wide as stria and separated by 1-3 diameters on disc and apical declivity. Intervals moderately convex, surfaceglossy, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion with at most few scarce setae, surface 116 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 finely alutaceous throughout. Prosternum behind procoxa completely bare ventrally,surface alutaceous throughout, posterior margin crenulated. Mesosternum strongly, semicircularly declivous anteriorly in larger males, transversely depressed on anterior half in females and smaller males, punctures large, coarse and dense on anterior two-thirds, posteriormost portion produced into wide lamellar process in larger males, reduced to lobate process with apex rounded to emarginate in females and smaller males, anterior two-thirds finely pubescent, setae short and apposed, surface on posterior third alutaceous. Mesepisternum weakly punctate, surface strongly alutaceous, finely carinate parallel to posterior edge. Metasternum with disc glossy, with deep longitudinal impression on most of its length in males or narrow and shallow impression on posterior two-thirds in females, surface of depression slightly to moderately alutaceous; lateral lobes on anterior half with punctures oval and shallowly impressed, scarce internally, anteriormost punctures with anteromedial edge bluntly carinate, posterior half with punctures broadly oval to rounded and restricted to metacoxal edge, leaving smooth area internally, surface between punctures alutaceous, except for an almost glossy area along metacoxal edge internally,pubescence completely missing; median lobe (Figs. 10, 11) moderately wide, L/mW = 2.1-2.4, lateral margins slightly divergent on anterior half, convergent on posterior half, anterior angles of lateral margins slightly rounded internally, surface minutely punctate and alutaceous,junction with mesosternum strongly angulate and projecting anteriorly in males or emarginate medially in females,weakly carinate medially, not extending into longitudinal carina on mesosternum. Legs. Anterior tibia quadridentate in both sexes, apical tooth pointing forward, slightly to moderately lobate externally in males or unmodified in females,ventromedial carina lacking intervening setae on basal two-third in both sexes, apical spur straight, parallel-sided and acute apically in males or subspatulate and obliquely truncate in females. Anterior femur unmodified. Posterior femur (Fig. 88) triangularly produced on posterodorsal edge apically, posteroventral edge dentate apically, posteroventral sulcus and posterior surface always clearly separated by posteroventral edge, never forming large depression, ventroposterior sulcus sharply delimited throughout, apical portion of posteroventral sulcus tapering toward apex, not extending anteriorly in males or femur unmodified in females and smaller males. Posterior tibia (Fig. 88) slender, inner edge lobate on apical two-thirds, anteroventral edge strongly carinate on basal two-thirds in larger males or unmodified in females and smaller males. Tarsal claws, when fully extended, at most one-fourth as long as last tarsal segment. Abdomen. Sternites 2-5 with at most few shallow punctures laterally, except for anterior edge with row of rather small and coarse irregular punctures laterally, surface finely alutaceous, sternite 6 glossy, suture between sternites 5-6 deeply sulcate throughout.
Pygidium, WIL = 1.9 in males or 2.3 in females,rather deeply sulcate along basal margin, surface minutely punctate,surface between punctures glossy,apical margin clearly delimited in both sexes. Male genitalia (Figs. 226-228). Tubus approximately 2.2 times as long as parameres, ventral portion slightly depressed apically. Parameres in lateral view elongate, with apex rounded, ventral portion slightly convex, produced into small tooth just before apex, ventral edge of right paramere widely and triangularly membranous, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 304.
MATERIALSTUDIED. IOaa,169 9 (BMNH,ISNB,NMPC,MCZC,MNHN,MZSP, SMFD, SMTD, USNM).
TYPE MATERIAL Ontherus cephalotes Harold: Lectotype 9 (MNHN): Nov. Friburg/ Cephalotes, typ. Harold/ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthiir, 1952/ LECTOTYPE, Ontherus cephalotes, Harold,1869, Des. F. Genier,1991. Type material studied and lectotype here designated.
Ontherus quadrituberculatus Luederwaldt: A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 117
• 0. cephalotes
MAP 26. Known distribution of Ontherus cephalotes.
Lectotype ct (MZSP): Petropolis, 1.II.1898, (Ohaus?)/ Ontherus ct, 4-tuberculatus, Lued erw., Luederw. det. 29./ blue disk/ COTIPO (pink paper)/ 16360/ LECTOTYPE, Ontherus quadrituberculatus, Luederwaldt, 1930, des. F. Genier, 1992. Type material studied and Lectotype here designated.
Paralectotypes: Same data as lectotype ( 1 ct, 2 9 9 MZSP).
NON-TYPE MATERIAL Distribution: BRAZIL: Caulajalla*. Bahia: No locality. Rio de Janeiro: Nova Friburgo; Petr6polis.
Additional data: Specimens collected in February, April, May, November.
REMARKS. Harold's type specimen is a small female in which the clypeofrontal carina is weakly developed. Therefore, the quadrilobate aspect of the carina is poorly pronounced and similar in aspect to the carina of 0. lobifrons sp.nov., for which it was mistakenly identified by previous authors.
DIGITATUS GROUP
DIAGNOSIS. Most heterogenous group of the genus. Mesometastemal suture angulate, combined with small size (5-11 mm), convex (except for ill-defined longitudinal basal sulcus) pronotum, and conspicuously punctate elytral intervals (except in 0. digitatus), distinguish the following species.
48. Ontherus (Ontherus) aphodioides Burmeister, 1874 comb.nov. (Figs. 229-231, 305; Map 27)
Ontherus aphodioides Burmeister, 1874: 126 (description) Ontherus aphodioides Burmeister: Gillet, 191 lh: 57 (catalog) 118 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Ontherus convexus Luederwaldt, 1930: 105 (description) Ontherus aphodioides Burmeister: Luederwaldt, 1931: 371, 384 (key, distribution and redescription) Ontherus convexus Luederwaldt: Luederwaldt, 1931: 372, 386 (key, distribution and redescription) Ontherus aphodioides Burmeister: Pessoa and Lane, 1941: 456 (key) Ontherus convexus Luederwaldt: Pessoa and Lane, 1941: 457 (key) Ontherus aphodioides Burmeister: Blackwelder, 1944: 206 (list) Ontherus convexus Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus aphodioides Burmeister: Lange, 1947: 310 (list) Ontherus aphodioides Burmeister: Martinez, 1959: 69 (catalog) Ontherus aphodioides Burmeister: Martinez and Pereira, 1960: 79, 80 (list and comment) Ontherus convexus Luederwaldt: Martinez and Pereira, 1960: 79, 80 (listed as synonym and comment)
REDESCRIPTION. Colour. Body dark brownish to black. Pubescence fulvous. Length. 7-11 mm. Head. Anterior edge arcuate, slightly upturned and bisinuate medially, clypeo genal junction straight. Clypeus transversely wrinkled on anterior half, punctures almost confluent on posterior half. Clypeofrontal suture ill-defined, with large laterally compressed conical process medially, or 2 adjacent small tubercles in females, reduced to low conical process in smaller individuals. Anterior genal edge slightly shorter than lateral, genal surface wrinkled on anterior fourth in males or half in females, remainder of surface moderately to coarsely punctate. Vertex punctate except on small area posteriorly. Pronotum. Transverse,
WIL = 1.8; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc distinctly convex in moderate and larger males, declivous on anterior fourth behind head insertion in males or slightly declivous behind head insertion in females and smaller males, basal half with a shallow and weakly delimited median sulcus; punctures fine and scattered on disc in males or only slightly fineron disc in females,lateral swellings finely punctate, punctures larger, finely umbilicate and denser along base, anterior and lateral declivities, almost confluent on anterior angles. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and impressed on disc and apical declivity, punctures oval, deeper than stria, approximately 1. 5 times as wide as stria and separated by 3-6 diameters on disc and apical declivity. Intervals slightly convex, surface distinctly (12X) and evenly punctate throughout, lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and strongly alutaceous throughout. Prosternum behind procoxa coarsely punctate, pubescence obliquely ori ented and almost equal in length and thickness. Mesosternum transversely impressed on disc, punctures coarse, strongly microsculptured and dense throughout, pubescence fine, few longer setae along mesocoxae. Mesepistemum irregularly punctate, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely and densely punctate, widely depressed medially on posterior half, surface of depression alutaceous; lateral lobes on anterior half with punctures oval and strongly umbilicate, posterior half with punctures rounded and scattered, surface strongly alutaceous on anterior half, surface on posterior half completely glossy between punctures, pubescence short and obliquely oriented throughout; median lobe wide, L/mW = 1.7, lateral margins slightly divergent on anterior third, slightly convergent on posterior two-thirds, surface punctate, punctures much larger, umbilicate and scarcer on anterior half, surface between punc tures alutaceous anteriorly, junction with mesosternum strongly angulate medially, finely carinate throughout, slightly produced into fine carina longitudinally on mesostemum. Legs. Anterior tibia quadridentate and similarly shaped in both sexes, ventromedial carina lacking intervening setae on basal half in both sexes, apical spur straight and inwardly bent on apical fourth. Posterior legs unmodified. Abdomen. Sternites 2- 5 with numerous coarse and umbilicate punctures laterally, anterior edge with complete row of bare, but umbilicate, punctures, surface alutaceous and minutely punctate throughout, stemite 6 finely punctate along anterior edge, suture between sternites 5-6 narrow and deeply sulcate. Pygidium, WIL
= 2.7 in males or 2.8 in females, punctures coarse, umbilicate and moderately dense, evenly distributed, surface glossy, apical margin clearly delimited in both sexes. Male genitalia A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 119
• O. aphodioides
MAP 27. Known distribution of Ontherus aphodioides.
(Figs. 229-231). Tubus approximately 2.8 times as long as parameres, ventral portion slightly concave. Parameres in lateral view with apex rounded, ventral portion slightly sinuous, shallowly emarginate basally, dorsal portion slightly concave. Sigmoid sclerite of internalsac as in Figure 305.
MATERIAL STUDIED. 28 ct, 32 9 (AMNH, BDGC, BMNH, CNCI, DEIC, ICCM, ISNB, MACN, MLPA, MNHN, NMPC, QBUM, SMFD, SEMC, SMTD, USNM, WDEC, ZSMC).
TYPE MATERIAL Ontherus aphodioides Burmeister: Ty pe lost.
Ontherus convexus Luederwaldt: Holotype ct (QBUM): Guandu, Espirito Santo (in Melzer collection). Holotype not studied (type unavailable).
NON- TYPE MATERIAL Distribution: ARGENTINA: Gob. Chaco. Corrientes: Bella Vista. Formosa: Formosa. Misiones: Eldorado; San Ignacio. Santa Fe: Las Garzas, Bord du Rio Las Garzas, 25 Kil a l'O. D'Ocampo; Rio Salado. BOLIVIA: Beni: Espiritu (250m); Santa Rosa (250m). Santa Cruz: Santa Cruz (500m). BRAZIL: Pernambuco: No locality. Rio de Janeiro: Rio de Janeiro. Rio Grande do Sul: Santo Augusto. Sao Paulo: Ribeirao Preto; Sao Paulo. PARAGUAY: Central: Asuncion. Guaira: Villarrica. Nueva Asuncion: Independencia. PERU: San Martin: Rioja. VENEZUELA: Bolivar: Parupa, grand Sabana.
Additional data: One adult specimen was collected in a light trap. The individual from Venezuela was collected using a flight intercept trap set at the forest-grassland edge. Specimens collected in April, June to December. 120 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
49. Ontherus (Ontherus) digitatus Harold, 1868 comb.nov. (Figs. 232-234, 306; Map 28)
Ontherus digitatus Harold, 1868h: 80 (description) Ontherus digitatus Harold: Gemminger and Harold, 1869: 1008 (catalog) Ontherus digitatus Harold: Harold, 1869a: 59 (comment) Ontherus digitatus Harold: Gillet, 191 lh: 58 (catalog) Ontherus digitatus Harold: Luederwaldt, 1931: 369, 373 (key, distribution and redescription) Ontherus digitatus Harold: Pessoa and Lane, 1941: 456, 457 (key, redescription and distribution) Ontherus digitatus Harold: Blackwelder, 1944: 206 (list) Ontherus digitatus Harold: Martinez, 1959: 71 (list and distribution)
REDESCRIPTION. Colour. Body dark reddish brown to black. Pubescence fulvous. Length. 7-9 mm. Head. Anterior edge broadly arcuate, upturned and rather widely but shallowly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior fifth in males or third in females, finely punctate posteriorly. Clypeofrontal suture ill-defined, lacking any traces of carina or tubercle. Anterior genal edge shorter than lateral, genal surface moderately to coarsely punctate throughout. Vertex punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and almost straight behind head insertion. Disc simple and slightly convex throughout in both sexes, basal half with at most trace of median sulcus; punctures minute to fine on disc, lateral swellings finely punctate, punctures large, dense, and coarse on lateral declivities. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and similarly impressed on disc and apical declivity, punctures oval, deeper than stria, approximately 1.5 times as wide as stria and separated by 4-5 diameters on disc and apical declivity. Intervals flat, surface minutely and evenly punctate throughout, lacking any trace of alu taceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion mod erately pubescent and strongly alutaceous throughout. Prosternum behind procoxa finely punctate, pubescence erect and subequal in length and thickness throughout. Mesosternum shallow and transversely impressed on disc, punctures moderate in size and coarse on anterior half, large, coarse, umbilicate and dense except for median longitudinal smooth area on posterior half, pubescence fine and obliquely oriented on anterior half, long and erect on posterior half. Mesepisternum irregularly punctate, surface moderately alutaceous, finely carinate parallel to posterior edge. Metasternum with disc finely and rather scarcely punctate, longitudinally and deeply depressed medially throughout its length in males or shallowly depressed on posterior half in females and smallest males, surface of depression glossy, lateral lobes on anterior half with punctures rounded, umbilicate and rather scarce, posterior half with punctures rounded and scattered, surface finely alutaceous along coxa on anterior half, surface on posterior half completely glossy between punctures, pubescence short and obliquely oriented to erect throughout; median lobe wide, L/mW = 1.4, lateral margins slightly divergent on anterior two-thirds, slightly convergent on posterior third, surface finely punctate, with row of larger and finely to moderately umbilicate punctures along coxal margin, surface between puncture glossy, junction with mesosternum angulate medially, finely carinate throughout. Legs. Anterior tibia quadridentate in both sexes, apical tooth slender, obliquely oriented, penultimate tooth conspicuously enlarged in males or tibiae unmodified in females, ventromedial carina lacking intervening setae on basal third in both sexes, apical spur straight and spiniformin males or straight and slightly spatulate in females. Posterior femur elongate, triangularly produced on posterodorsal edge apically, posteroven tral sulcus and posterior surface fused and almost on same plane, forming largely depressed area, surface of depression glossy, apical edge of depression sharply delimited, not extending anteriorly, femur simple in females. Posterior tibia with internal edge dentate medially in males or simple in females and smaller males. Abdomen. Sternites 2-5 with numerous coarse and umbilicate punctures laterally, anterior edge with row of umbilicate punctures, A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 121
• 0. digitatus
MAP 28. Known distribution of Ontherus digitatus.
surface glossy throughout, sternite 6 with few fine punctures, suture between sternites 5-6 narrowly and deeply sulcate. Pygidium, W/L = 1.9 in males or 2.3 in females, punctures coarse, umbilicate and moderately dense, evenly distributed, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 232-234). Tubus approximately 2.0 times as long as parameres, ventral portion concave. Parameres in lateral view with apex tapered, ventral portion straight, emarginate basally, dorsal portion straight, apical third finely rugose. Sigmoid sclerite of internal sac as in Figure 306.
MAT ERIAL STUDIED. 36 a a, 35 Q Q (CASC, CNCI, ICCM, IMLA, MACN, MNHN, NMPC, SMTD, USNM, WDEC, ZMHB, ZMUC, ZSMC).
TYPE MATERIAL Holotype a (MNHN): Brasilia/ digitatus, t. Harold/ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthiir, 1952/ HOLOTYPE, Ontherus digitatus, Harold, 1868, vidit F.Genier, 1991. Type material studied and lectotype here designated.
NON-TYPE MATERIAL Distribution: ARGENTINA: Cordoba: Rio Cuarto; San Bernardino. Jujuy: Termas del Palmar*. Salta: Salta. Santiago del Estero: Rio Salado*. Tucuman: Concepcion. BOLIVIA: Santa Cruz: Samaipata, 1500m; Santa Cruz, 500m. BRAZIL: Minas Gerais: Aguas Vermelhas; Sete Lagoas. Rio de Janeiro: No locality. Sao Paulo: Anhembi, Fazenda Barreiro Rico. PARAGUAY: Presidente Hayes: Rio Confuso, Santa Clara.
Additional data: One individual was collected at light. Specimens collected in January, February, August to December.
50. Ontherus (Ontherus) elegans Luederwaldt, 1930 comb.nov. (Figs. 235, 236, 307; Map 29)
Ontherus elegans Luederwaldt, 1930: 106 (description) 122 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
Ontherus elegans Luederwaldt: Luederwaldt, 1931: 371, 382 (key, distribution and redescription) Ontherus elegans Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus elegans Luederwaldt: Blackwelder, 1944: 206 (list) Ontherus elegans Luederwaldt: Pereira and d 'Andretta, 1955: 251 (distribution and redescription)
REDESCRIPTION. Colour. Body dark reddish brown to black, some individuals with slight green metallic sheen on elytra, basal third of pronotum and pygidium, remainder of dorsum and ventrum with moderate to distinct coppery sheen. Pubescence fulvous. Length. 8-11 mm. Head. Anterior edge arcuate, moderately upturnedand at most slightly emarginate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half, finely punctate posteriorly. Clypeofrontal suture arcuate, weakly carinate, with moderate conical process medially posterior to suture, reduced to small tubercle in smaller individuals. Anterior genal edge equal to lateral, genal surface transversely wrinkled on anterior half, remainder of surface with punctures shallow and confluent. Vertex punctate except on small
area posteriorly. Pronotum. Transverse, W/L = 1.7; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc strongly convex in moderate and larger males, declivous on anterior fourth behind head insertion in males or slightly declivous behind head insertion in females and smaller males, basal half with shallow and weakly delimited median sulcus; punctures fine and dense on disc, lateral swellings finely punctate, punctures larger and denser along base, anterior and lateral declivities, almost less confluent
on anterior angles. Elytra. Subquadrate, L/W = 1.1. Striae moderately wide and shallowly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approxi mately 1.5 times as wide as stria and separated by 3-4 diameters on disc and apical declivity. Intervals slightly convex, distinctly (12X) and evenly punctate throughout, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and strongly alutaceous throughout. Prosternum behind procoxa finely punctate, pubescence scarce, erect, fine and uneven in length and thickness. Mesoster num transversely impressed on disc, punctures rather large and coarse, strongly microsculp tured and dense throughout, pubescence fine and apposed, few longer setae along mesocoxae. Mesepisternum scarcely and finely punctate, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely and densely punctate, finely sulcate medially; lateral lobes on anterior half with punctures poorly defined, broadly oval to rounded, posterior half with punctures rounded and scattered, surface strongly alutaceous on anterior half, surface on posterior half completely glossy between punctures, pubescence short and apposed throughout; median lobe rather narrow, L/mW
= 2.2, lateral margins slightly divergent on anterior fourth, slightly convergent on posterior three-fourths, surface finely punctate on posterior two-thirds, glossy with occasional larger puncture on anterior third, surface between puncture finely alutaceous on posterior two thirds, junction with mesosternum strongly angulate medially, finely carinate throughout. Legs. Anterior tibia quadridentate and similarly shaped in both sexes, ventromedial carina lacking intervening setae on basal four-fifths in males or half in females, apical spur slender and inwardly bent on apical fourth. Posterior femur deeply notched on anterior edge medially, inner portion produced into spine in males or unmodified in females. Abdomen. Sternites 2-5 with few coarse and umbilicate punctures laterally, anterior edge with complete row of irregular punctures, surface glossy, sternite 6 finely punctate throughout, suture between sternites 5-6 with row of coarse punctures, lacking sulcus. Pygidium,
WIL = 2.0 in males or 2.1 in females, punctures fine, moderately dense, slightly denser laterally, surface glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 235, 237). Tubus approximately 2.0 times as long as parameres, ventral portion slightly concave apically. Parameres in lateral view with apex rounded, ventral portion concave, shallowly emarginate basally, dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 307. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 123
• 0. elegans
MAP 29. Known distribution of Ontherus elegans.
MATERIAL STUDIED. 11 a, 15 9 9 (AMNH, CNCI, HAHC, ISNB, MZSP, NMPC, SMTD).
TYPE MATERIAL Lectotype 9 (MZSP): Sao Paulo, Ypiranga, IV.(19)29, R. Spitz leg./ Ontherus elegans lueder., Lueder. det. 29./ COTIPO (pink paper)/ 16400/ Lectotype, Ontherus elegans Luederwalt, 1930, Des. F. Genier, 1992. Ty pe material studied, lectotype and paralec totypes here designated.
Paralectotypes: Sao Paulo, Est. Alto Siera, I.[19)26, Spitz leg./ Ontherus elegans lueder., Lueder. det. 29./ blue disk/ COTIPO (pink paper)/ 16399/ Paralectotype, Ontherus elegans Luederwalt, 1930, Des. F. Genier, 1992. ( 1 9 MZSP). Minas Gerais, Passo Quatro, XI.[19]24, Jaeger leg./ Ontherus elegans lueder., Lueder. det. 29./ blue disk/ COTIPO (pink paper)/ 16398/ Paralectotype, Ontherus elegans Luederwalt, 1930, Des. F. Genier, 1992 (1 9 MZSP).
NON-TYPE MATERIAL Distribution: BRAZIL: Lages*. Distrito Federal: Estacao Florestal, Cabeca do Veado (llOOm). Minas Geraes: Passa Quatro. Parana: Banhados (800m), on Curitiba to Paranagua road; Morretes. Santa Catharina: Rio das Antas; Rio Vermelho; Sao Bento do Sul (800m); Sao Paulo: Esta90 Biologica do Alto da Serra (Paranapiacaba), Ipiranga.
Additional data: Specimens collected in January, February, June, September to December.
REMARKS. Luederwaldt's type material (3 9 9 MZSP, Minas Gerais and Sao Paulo) and 2 other specimens (1 9 CNCI, Distrito federal and 1 9 ISNB, no locality) have strongly alutaceous elytra and have the cephalic tubercle located on the clypeofrontal carina. All other specimens from the States of Parana and Santa Catarina have the elytra glossy between the punctures and the cephalic tubercle located behind the clypeofrontal carina. These two characters and the known distribution suggest that these two forms could represent different 124 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 taxa. However, because no males are available for the Luederwalt formI consider specimens of these forms to be conspecific. The male allotype designation by P. F. S. Pereira and M. A. V. d 'Andretta violates recommendation 72a of the I.C.Z.N., and is invalid.
51. Ontherus (Ontherus) androgynus sp.nov. (Figs. 13, 238-240, 308; Map 30)
ETYMOLOGY. Androgynus, (male + female) Greek adjective, refers to the lack of secondary sexual characters in males of this species.
DESCRIPTION. Colour. Body dark reddish brown to black, some individuals with slight green metallic sheen on elytra and posterior angles of pronotum. Pubescence fulvous. Length. 8-10 mm. Head. Anterior edge arcuate, moderately upturned and at most slightly emarginate medially or slightly bisinuate in females, clypeogenal junction straight. Clypeus transversely wrinkled on anterior two-thirds, punctate posteriorly. Clypeofrontal suture broadly arcuate, carinate, with 2 adjacent small conical processes medially on suture. Anterior genal edge equal to lateral, genal surface transversely wrinkled on most of surface, punctures shallow and confluent, adjacent to eye. Vertex coarsely and rather densely punctate, except on small area posteriorly. Pronotum. Transverse, W/L = 1.7; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc slightly convex in males, declivous on anterior fourthbehind head insertion in males or simple in females, basal half with shallow and weakly delimited median sulcus; punctures coarse and dense on anterior angles, along anterior and posterior margins, slightly smaller and less dense on disc, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and similarly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approximately 1.5-2.0 times as wide as stria and separated by 1-3 diameters on disc and apical declivity. Intervals slightly convex, distinctly (12X) and evenly punctate through out, punctures weakly delimited, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion scarcely pubescent and strongly alutaceous throughout. Prosternum behind procoxa lacking distinct punctures, pubescence scarce, fine and uneven in length and thickness. Mesosternum transversely impressed on disc, punctures rather large and coarse, strongly microsculptured and rather dense through out, pubescence fine and apposed, few longer setae along mesocoxae, surface between punctures glossy except for finely alutaceous portion on posterior fifth where punctures weakly delimited. Mesepisternum scarcely and finely punctate, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metastemum with disc finely punctate, sulcate medially; lateral lobes on anterior half with punctures oval, rather poorly defined, anterior most punctures each with anterior edge finely carinate, posterior half with punctures rounded, umbilicate and moderately dense, surface strongly alutaceous on anterior half, surface on posterior half completely glossy between punctures, pubescence short and apposed throughout; median lobe (Fig. 13) rather narrow, L/mW = 2.1, lateral margins slightly divergent on anterior fourth, slightly convergent on posterior three-fourths, surface finely punctate on posterior two-thirds, glossy with occasional larger punctures on anterior third, surface between punctures finely to moderately alutaceous along anterior and lateral margins, junction with mesosternum strongly angulate medially, finely carinate throughout. Legs. Anterior tibia quadridentate and similarly shaped in both sexes, slightly more slender in males, ventromedial carina lacking intervening setae on basal four-fifths in males or half in females, apical spur slender and inwardly bent on apical fourth. Posterior femur unmodi fied in both sexes. Abdomen. Sternites 2-5 with few coarse and umbilicate punctures laterally, anterior edge with row of irregular punctures laterally, surface glossy, sternite 6 finely punctate throughout, suture between sternites 5-6 shallowly sulcate, sulcus reduced to few coarse punctures laterally in some individuals. Pygidium, W/L = 1.7 in males or 2.0 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 125
• 0. androgynus \
MAP 30. Known distribution of Ontherus androgynus.
in females,punctures moderate in size and rather dense,surface glossy,apical margin clearly delimited in both sexes. Male genitalia (Figs. 238-240). Tubus approximately 1.8 times as long as parameres, ventral portion slightly depressed apically. Parameres in lateral view with apex rounded, ventral portion concave,shallowly emarginate basally,dorsal portion slightly convex. Sigmoid sclerite of internal sac as in Figure 308.
MATERIAL STUDIED. 8 O'O', 149 9 (AMNH, BMNH, DEIC, MCNZ, MNHN, NMPC, SMFD, SMTD, USNM, ZMHB).
Holotype CJ' (ZMHB): Brasilia, St Catharina/ Zool. Mus., Berlin/ HOLOTYPE, Ontherus androgynus, F. Genier, 1992. Dissected.
Allotype 9 (MNHN): Bresil, Prov. Sta Catarina, Hansa Humboldt, Ant. Maller/ Museum Paris, ex Coll., R. Oberthur/ ALLOTYPE, Ontherus androgynus, F. Genier, 1992.
Paratypes: BOLIVIA: No locality (1 9 BMNH). BRAZIL: No locality (29 9 DEIC) (1 CJ' NMPC). Minas Gerais: Env. de Passa-Quatro, Bord du Rio Das Pedras (lOOOm), E.-R. Wagner, 1903 (19 MNHN); Sete Lagoas, F. Ohs, 20.12.[18]98 (19 USNM). Rio Grande do Sul: Sao Jeronimo, 24.VIII.1982, C. J. Becker leg. (1CJ' MCNZ). Santa Catarina: No locality ( 1 O' MNHN); Hansa Humboldt,Reitter ( 1CJ' NMPC), same as before except: Corupa,I.1948, A. Maller (1 9 AMNH), XI.1948 (1 9 AMNH); Joinville, F. Ohs, 29 3.05 (1 CJ'SMFD). Sao Paulo:? Sao Paulo (1O',39 9 SMTD). PARAGUAY: Guaira: Independance (19 MNHN). URU GUAY: Montevideo: Montevideo (1CJ',29 9 MNHN).
52. Ontherus (Ontherus) erosioides Luederwaldt, 1930 comb.nov. (Figs. 241-243, 309; Map 31)
Ontherus erosioides Luederwaldt, 1930: 106 (description) Ontherus erosioides Luederwaldt: Luederwaldt, 1931: 371, 379 (key, distribution and redescription) Ontherus erosioides Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus erosioides Luederwaldt: Blackwelder, 1944: 206 (list) 126 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
REDESCRIPTION. Colour. Body dark reddish brown to black, lacking any trace of green metallic sheen. Pubescence fulvous. Length. 5-9 mm. Head. Anterior edge arcuate, moderately upturned and at most slightly emarginate medially or slightly bisinuate in females, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third, punctate posteriorly. Clypeofrontal suture broadly arcuate, carinate. Anterior genal edge slightly shorter than lateral, genal surface transversely wrinkled on anterior fourth, remainder of surface moderately to coarsely punctate. Vertex coarsely and rather densely punctate except on small area posteriorly. Pronotum. Transverse, W IL = 1.8; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex in both sexes, basal third with at most faint indication of median sulcus; punctures coarse and dense throughout, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and similarly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approximately 1.5 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals almost flat, distinctly (12X) and evenly punctate throughout, punctures weakly delimited, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion scarcely pubescent and strongly alutaceous throughout. Prostemum behind procoxa weakly punctate, pubescence scarce, fine, and almost even in length and thickness. Mesostemum rather shallowly transversely impressed on disc, punctures large and coarsely umbilicate, moderately dense throughout, pubescence fine and apposed, few longer setae along mesocoxae, surface between punctures glossy throughout. Mesepistemum with few large and vagely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely punctate, narrowly and moderately sulcate medially in males or shallowly sulcate in females; lateral lobes on anterior half with punctures broadly oval, well-defined and umbilicate, posterior half with punctures rounded, moderately to coarsely microsculptured and moderately dense, surface strongly alutaceous on anterior half, surface on posterior half completely glossy between punctures, pubescence fine, short and apposed throughout; median lobe wide, L/mW = 1.5, lateral margins almost parallel-sided on anterior three fourths, slightly convergent on posterior fourth, surface finely punctate on posterior two thirds, with larger slightly umbilicate puncture on anterior third, surface completely glossy between punctures, junction with mesosternum angulate medially, weakly carinate through out. Legs. Anterior tibia quadridentate and similarly shaped in both sexes, slightly more slender in males, basal tooth small, superomedian edge slightly emarginate basally in larger males, ventromedial carina lacking intervening setae on basal four-fifths in males or half in females, apical spur slender and inwardly bent on apical fourth. Anterior femur with superomedian edge concave on most of its length, its apical portion produced into inwardly directed conical process in males or almost simple in females and smaller males. Posterior femur with anterior edge produced into denticle on apical third, surface strongly depressed posterior to denticle, posterodorsal edge triangularly produced apically in males or femur unmodified in females. Abdomen. Stemites 2-5 with few coarse and umbilicate punctures laterally, anterior edge with row of large, coarse and umbilicate punctures laterally, surface between punctures glossy, sternite6 finely punctate posteriorly, with row of coarse unaligned punctures anteriorly, suture between sternites 5-6 shallowly sulcate. Pygidium, W/L = 2.2 in males and females, punctures coarse and large on disc, much smaller along apical edge, surface between punctures glossy, apical margin ill-defined in both sexes. Male genitalia (Figs. 241-243). Tubus approximately 1.6 times as long as parameres, ventral portion unmodified. Parameres in lateral view with apex obliquely truncate, ventral portion slightly concave, dorsal portion straight. Sigmoid sclerite of internal sac as in Figure 309.
MATERIAL STUDIED. 25 a a, 22 9 9 (CNCI, DEIC, FGIC, HAHC, ICCM, ISNB, MACN, MCNZ, MCZC, MNHN, MZSP, NMPC, SMTD, USNM, WDEC, ZMHB). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 127
• 0. erosioides
MAP 31. Known distribution of Ontherus erosioides.
TYPE MATERIAL Lectotype et (MZSP): (Brazil), Sao Paulo, Ypiranga, III.[19]20/ Ontherus et erosioides, Luederw., Luederw. det. 29/ blue disk/ COTIPO (pink paper)/ 16378/ LECTOTYPE, Ontherus erosioides, Luederwaldt, 1930, Des. F. Genier, 1992. Type material studied, lectotype and paralectotypes here designated.
Paralectotypes: Same data as lectotype except: VII.20/ 16377 (1 et MZSP), 1.29, R. Spitz/ 16379 (1 9 MZSP), 12.553, (no collector's name)/ Ontherus sp., Felsche det. 09/ 16381 (1 9 MZSP), E. Alto Siera, 1.26, Spitz leg./ 16380 (1 9 MZSP).
NON-TYPE MATERIAL Distribution: ARGENTINA: Misiones: Eldorado; P. N. Iguas,:u; Env. de San-Ignacio, Villa Lutecia. BRAZIL: Porto do Cadi*. Distrito Federal: Estacao Florestal Cabeca do Veado (1 lOOm). Mato Grosso: Chapada dos Guimaraes. Rio de Janeiro: Cantagalo; Rio de Janeiro. Rio Grande do Sul: Eldorado do Sul*; Porto Alegre. Sao Paulo: Estas,:ao Biologica do Alto Serra (Paranapiacaba); Ipiranga; Piraeicaba*. PARAGUAY: Pastorea*. Paraguari: Sapucay. Caazapa: Caazapa.
Additional data: Specimens collected in January to March, July, September to December.
53. Ontherus (Ontherus) amplector sp.nov. (Figs. 244-246, 310; Map 32)
ETYMOLOGY. Amplector, (embracer) Latin noun, refers to modifications of anterior legs in males.
DESCRIPTION. Colour. Body dark reddish brown to black, lacking any trace of green metallic sheen. Pubescence fulvous. Length. 8-9 mm. Head. Anterior edge arcuate, moderately upturned and at most slightly emarginate medially or slightly bisinuate in females, clypeogenal junction straight. Clypeus transversely wrinkled on anterior fourth in 128 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 males or third in females, punctate posteriorly. Clypeofrontal suture broadly arcuate, carinate. Anterior genal edge as long as lateral, genal surface punctate throughout in males or transversely wrinkled on anterior fourth with remainder of surface moderately to coarsely punctate in females. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.7; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex in both sexes, basal third with at most faint indication of median sulcus; punctures fine on disc, larger and denser on lateral declivities, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.1. Striae moderately wide and similarly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approximately 1.5 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals almost flat,distinctly (12X) and evenly punctate throughout, punctures weakly delimited, punctures finein some individuals, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and alutaceous throughout. Prosternum behind procoxa weakly punctate, pubescence moderate and almost even in length and thickness. Mesostemum rather shallowly transversely impressed on disc, punctures large and coarsely umbilicate, moder ately dense throughout, pubescence fine and semi-erect, few longer setae along mesocoxae, surface between punctures glossy throughout. Mesepistemum with few moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metastemum with disc finely punctate, narrowly and moderately sulcate medially in males or shallowly sulcate in females; lateral lobes on anterior half with punctures broadly oval, well-defined and umbilicate, posterior half with punctures rounded, slightly umbilicate and moderately dense, surface finely alutaceous on anterior third, surface on posterior two-thirds completely glossy between punctures, pubescence fine, short and apposed throughout; median lobe wide, L/mW = 1.6, lateral margins slightly divergent on anterior third, slightly convergent on posterior two-thirds, surface finely punctate on poste rior half, with larger slightly umbilicate punctures on anterior half, surfacecompletely glossy between punctures, junction with mesosternumangulate medially, weakly carinate through out. Legs. Anterior tibia quadridentate, slightly more slender in males, basal tooth small, superoproximal edge produced into short lamellar projection and strongly emarginate basally in males or unmodifiedin females, ventromedial carina lacking intervening setae on basal four-fifthsin males or half in females,apical spur slender and acute in males or inwardly bent on apical fourthin females.Anterior femur with superoproximal edge concave on most of its length, its apical portion produced into inwardly directed conical process in males or almost simple in females. Posterior femur gradually widening toward apex, anterior edge denticulate on apical sixth, surface convex, posterodorsal edge acutely produced apically in males or femur unmodified in females. Abdomen. Stemites 2-5 with few coarse and umbilicate puncture laterally, anterior edge with row of large, coarse and umbilicate punctures laterally, surface between punctures glossy, stemite 6 finely punctate posteriorly, with row of coarse unaligned punctures anteriorly, suture between stemites 5-6 shallowly sulcate. Pygidium, W/L = 2.1 in males and females, punctures fine basally, gradually decreasing in size toward apex, surfacebetween punctures glossy, apical margin ill-defined in both sexes. Male genitalia (Figs. 244-246). Tubus approximately 1.7 times as long as parameres, ventral portion depressed apically. Parameresin lateral view with apex obliquely truncate, ventral portion slightly convex, dorsal portion straight. Sigmoid sclerite of internal sac as in Figure 310.
MATERIAL STUDIED. 3 a a, 5 9 9 (ICCM, MNHN, MZSP, ZMHB).
TYPE MATERIAL Holotype O' (ZMHB): Brasilia, Virm., Nr. 26330/ erosus Har.,? sec. Felsche/ Zool. Mus., Berlin/ HOLOTYPE, Ontherus amplector, F. Genier, 1992. Dissected. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 129
• 0. amplector
* 0. atlantidis
• 0. dentatus
MAP 32. Known distribution of Ontherus amplector, 0. at/antidis, and 0. dentatus.
Allotype 9 (ZMHB): Brasil, Heyer, Nr. 26330/ Zool. Mus., Berlin/ ALLOTYPE, Ontherus amplector, F. Genier, 1992.
Paratypes: BRASIL: No locality (1 CJ', 1 9 ZMHB) (1 9 MNHN). Rio de Janeiro: Parque Nacional do Itatiaia, II.1968, Coll. 0. Leoncini (1 9 ICCM). Sao Paulo: Ypiranga (=Ipi ranga), I.[19Ji9, Spitz leg. (1 CJ' MZSP); Mato d Govern.*, 10.I.[19]26, Ohaus S. (19 ZMHB).
54. Ontherus (Ontherus) atlantidis sp.nov. (Map 32)
ETYMOLOGY. Atlantidis, (from Atlantida) Latin noun, refers to type locality.
DESCRIPTION. Colour. Body dark reddish brown, lacking any trace of green metal lic sheen. Pubescence fulvous. Length. 10-11 mm. Head. Anterior edge arcuate, slightly upturned and bisinuate medially, clypeogenal junction straight. Clypeus transversely wrin kled throughout. Clypeofrontal suture broadly arcuate, carinate and distinctly tumescent medially. Anterior genal edge slightly longer thanlateral, genal surface transversely wrinkled on most of surface, moderately to coarsely punctate adjacent to eye. Vertex rather coarsely and densely punctate except on small area posteriorly. Pronotum. Transverse, W/L = 1.7; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex, basal third with shallow subcircular depression medially; punctures fine on disc, larger and much denser along anterior and posterior margin and on lateral declivities, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.1. Striae fine and similarly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approximately 2.0 times as wide as stria and separated by 2-3 diameters on disc and apical declivity. Intervals slightly convex, distinctly (12X) and evenly punctate throughout, punctures weakly delimited, surface finely alutaceous along striae. Thoracic sterna. Proepistemum with posterolateral portion scarcely pubescent and alutaceous throughout. Prostemum behind procoxa scarcely and weakly 130 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 punctate, pubescence moderate and almost even in length and thickness. Mesosternum rather shallowly transversely impressed on disc, punctures moderate in size and umbilicate, moderately dense throughout, pubescence fineand apposed, fewlonger and erect setae along mesocoxae, surface between punctures finely to moderately alutaceous throughout. Mesepisternum with few moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely and scarcely punctate, narrowly and moderately sulcate medially ; lateral lobes on anterior half with punctures broadly oval, well-defined and umbilicate, posterior half with punctures rounded, slightly umbilicate and moderately dense, surface strongly alutaceous on anterior two-thirds, surface on posterior third almost completely glossy between punc tures, pubescence fine, short and apposed, longer along mesocoxa; median lobe moderately narrow, L/mW = 2.3, lateral margins slightly divergent on anterior third, slightly convergent on posterior two-thirds, surface finely punctate on posterior half, with larger, scarcer and poorly defined punctures on anterior half, surface between punctures alutaceous on anterior third, completely glossy on posterior two-thirds, junction with mesosternum angulate medially, finely and sharply carinate throughout. Legs. Anterior tibia quadridentate, ventromedial carina lacking intervening setae on basal two-thirds, apical spur slender and acute, inwardly bent on apical fourth. Abdomen. Sternites 2-5 with several umbilicate punctures laterally, anterior edge with row of moderate-sized and shallow punctures laterally, surface between punctures finely alutaceous, stemite 6 finely punctate posteriorly, with row of larger unaligned punctures anteriorly, suture between sternites 5-6 unmodified. Pygidium, WIL = 2.2, punctures fine basally, gradually decreasing in size toward apex, surface between punctures glossy, apical margin clearly delimited. Male genitalia. Unknown.
MATERIAL STUDIED. 2 Q Q (MACN). Holotype Q (MACN): Atlantida/ Mus. Arg., Cs. Nat./ HOLOTYPE, Ontherus atlantidis, F. Genier, 1992.
Paratype: URUGUAY: Canelones: Same data as holotype (1 Q MACN).
55. Ontherus (Ontherus) dentatus Luederwaldt, 1930 comb.nov. (Figs. 247-249, 311; Map 32)
Ontherus dentatus Luederwaldt, 1930: 106 (description) Ontherus dentatus Luederwaldt: Luederwaldt, 1931: 371, 380 (key, distribution and redescription) Ontherus dentatus Luederwaldt: Pessoa and Lane, 1941: 456 (key) Ontherus dentatus Luederwaldt: Blackwelder, 1944: 206 (list)
REDESCRIPTION. Colour. Body dark reddish brown, with faint green metallic to coppery sheen on vertex, pronotum and elytra. Pubescence fulvous. Length. 8-9 mm. Head. Anterior edge arcuate, moderately upturned and emarginate on median seventh, clypeogenal junction straight. Clypeus transversely wrinkled on anterior fourth, punctate posteriorly. Clypeofrontal suture broadly arcuate, carina simple and fine. Anterior genal edge slightly longer than lateral, genal surface finely punctate throughout. Vertex punctate except on small area posteriorly. Pronotum. Transverse, WIL= 1.7; anterior edge narrowly membranous and almost straight behind head insertion. Disc convex, basal third with at most faint indication of median sulcus; punctures moderate in size on disc, slightly larger and denser on lateral declivities, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and similarly impressed on disc and apical declivity, punctures oval, slightly deeper than stria, approximately 1.4 times as wide as stria and separated by 2-3 diameters on disc and apical declivity. Intervals convex, distinctly (12X) and evenly punctate through out, punctures fine and well-delimited, surface between punctures glossy. Thoracic sterna. Proepisternum with posterolateral portion moderately pubescent and alutaceous throughout. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 131
Prosternum behind procoxa with surface rugose and alutaceous, pubescence erect and uneven in length and thickness. Mesosternum rather shallow and transversely impressed on disc, punctures large and coarsely microsculptured, moderately dense throughout, pubes cence fine and semi-erect, few longer, and erect, setae along mesocoxae, surface between punctures glossy on anterior half, finely alutaceous on posterior half. Mesepisternum with few, moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely punctate, narrowly and weakly sulcate medially; lateral lobes on anterior half with punctures oval, shallow, well-defined and umbilicate, posterior half with punctures rounded and scarce, surface alutaceous on anterior half, surface on posterior half completely glossy between punctures, pubescence fine, short and semi-erect throughout; median lobe wide, L/mW = 1.5, lateral margins slightly divergent on anterior half, slightly convergent on posterior half, surface finely punctate on posterior half, with larger slightly umbilicate punctures on anterior half, surface completely glossy between punctures, junction with mesosternum angulate medially, carina fine and sharp throughout. Legs. Anterior tibia quadridentate, slender, ventromedial carina lacking intervening setae on basal half, apical spur subspatulate and rounded, ventrally bent on apical fourth. Posterior femur anterior edge triangularly produced on basal third, ventral surface of projection concave and microsculptured, posteroventral sulcus lacking. Abdomen. Sternites 2-5 with few shallow and umbilicate punctures laterally, anterior edge with row of aligned larger, umbilicate punctures laterally, surface between punctures glossy, sternite6 finely punctate posteriorly, with row of umbilicate, larger and unaligned punctures
anteriorly, suture between sternites 5- 6 unmodified. Pygidium, W/L = 1.8, basal half strongly bulged, punctures fine basally, gradually decreasing in size toward apex, surface between punctures glossy, apical margin ill-defined. Male genitalia (Figs. 247-249). Tubus approximately 1.9 times as long as parameres, ventral portion unmodified. Parameres in lateral view with apex rounded, ventral portion slightly convex, dorsal portion slightly concave. Sigmoid sclerite of internal sac as in Figure 311.
MATERIAL STUDIED. 2 a a (ICCM, MZSP).
TYPE MATERIAL Holotype O' (MZSP): (Brazil) Est. Sao Paulo, Angatuba, I.1923, Borgmeier/ Ontherus dentatus Lueder., Lueder. det. 29./ blue disk/ TIPO/ 16390. Holotype studied.
NON- TYPE MATERIAL Distribution: BRAZIL: Mato Grosso: Chapada dos Guimaraes.
Additional data: Specimens collected in October.
EROSUS GROUP
DIAGNOSIS. Greatly enlarged, and strigulate, lateral mesocoxal margin is unique in the genus.
56. Ontherus (Ontherus) insolitus sp.nov. (Figs. 250-252, 312; Map 33)
ETYMOLOGY. Insolitus, (strange, odd) Latin adjective, refers to the highly derived features of this species.
DESCRIPTION. Colour. Body dark reddish brown, lacking any trace of green metallic sheen. Pubescence fulvous. Length. 10-12 mm. Head. Anterior edge broadly arcuate, moderately upturned and slightly bidentate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or two-thirds in females, finely punctate posteriorly. Clypeofrontal suture broadly arcuate, almost straight medially, finely tuberculate 132 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 medially and laterally. Anterior genal edge as long as lateral, genal surface finely punctate throughout in males, or weakly wrinkled internally and remainder of surface minutely punctate in females. Vertex finely punctate except on small area posteriorly. Pronotum.
Transverse, WIL = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex in both sexes, basal third with at most faint indication of median sulcus, basal margin slightly produced on median fourth; punctures fine on disc, larger and denser on lateral declivities, fine on lateral swellings. Elytra. Subquadrate, L/W = 1.1. Striae moderately wide and slightly more impressed apical declivity, punctures rounded, as deep as stria, finely alutaceous and encroaching on intervals, approximately 1.5-2.0 times as wide as stria and separated by 2-3 diameters on disc and apical declivity. Intervals convex, finely punctate throughout, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and alutaceous throughout. Prosternum behind procoxa moderately to coarsely microsculptured throughout, pubescence fine and short throughout, with few long and erect setae on disc. Mesostemum rather shallow and transversely impressed on disc, punctures large and coarsely umbilicate, dense throughout, pubescence fine and semi-erect, few longer setae along mesocoxae and posterior portion of disc, surface between punctures finely alutaceous throughout. Mesepisternum with few moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely and unevenly punctate, foveolate on median third in males or finely and shallowly sulcate in females; lateral lobes with coxal margin strongly and triangularly produced anteriorly, surface strigulate, remainder of surface on anterior half with punctures broadly oval, each with anterior edge finelycarinate, with aligned row of approximately 7 transverse ridges, posterior half with punctures rounded, slightly umbilicate and mostly restricted to lateral portion, surface finely alutaceous anterolaterally, completely glossy on remainder of surface between punctures, pubescence fine and short, semi-erect throughout; median lobe wide, L/mW = 2.0, lateral margins divergent on anterior third, slightly convergent on posterior two-thirds, surface with small umbilicate punctures, surface almostglossy between punctures, junction with mesosternum angulate medially, weakly carinate throughout. Legs. Anterior tibia quadridentate, more slender in males, basal tooth small, superoproximal edge unmodified, ventromedial carina lacking intervening setae on basal two-thirds to three fourthsin males or half in females,apical spur slender and acute in males or slightly inwardly bent on apical fourth in females. Anterior femur unmodified. Posterior femur with anterior edge broadly arcuate and produced into sharp lamellar process, ventral surface convex, posterodorsal edge slightly and sharply lamellate on apical fourth in males or femur unmodified in females. Abdomen. Sternites 2-5 with several coarse and umbilicate punctures laterally, anterior edge with row of large, coarse and umbilicate punctures laterally, surface between punctures glossy, stemite 6 finely punctate posteriorly, with row of coarse unaligned punctures anteriorly, suture between stemites 5-6 simple. Pygidium strongly convex, WIL = 1.8 in males or 2.0 in females, punctures fine throughout, surface between punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 250- 252). Tubus approximately 2.3 times as long as parameres, ventral portion produced into triangular swelling medially and short longitudinal carina laterally. Parameres in lateral view with apex rounded, ventral portion slightly concave, dorsal portion strongly concave. Sigmoid sclerite of internal sac as in Figure 312.
MATERIAL STUDIED. 3 a a, 1 9 (BMNH, ISNB).
Holotype a (BMNH): Fry, Rio Jan./ 9364/ Fry Coll., 1905-100./ HOLOTYPE, Ontherus insolitus, F. Genier, 1992. Dissected.
Allotype 9 (BMNH): Santa Cath./ 10489/ Fry Coll., 1905-100/ ALLOTYPE, Ontherus insolitus, F. Genier, 1992. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 133
Paratypes: BRAZIL: No locality (IO' BMNH) (IO' ISNB).
57. Ontherus (Ontherus) stridulator sp.nov. (Figs. 253-255, 313; Map 33)
ETYMOLOGY. Stridulator, (stridulate) Latin noun, refers to probable use of modified punctures on lateral lobes of the metasternum.
DESCRIPTION. Colour. Body dark reddish brown, darker individuals with faint green metallic sheen on disc of pronotum and elytrae. Pubescence fulvous. Length. 9-10 mm. Head. Anterior edge broadly arcuate, moderately upturned and slightly bidentate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior third to half, finely punctate posteriorly. Clypeofrontal suture broadly arcuate, almost straight medially, finely tuberculate medially and laterally. Anterior genal edge as long as lateral, genal surface finely punctate throughout to weakly wrinkled internally and remainder of surface minutely punctate. Vertex with punctures moderate in size except on small area posteriorly. Pronotum.
Transverse, W /L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex in both sexes, basal third with at most faint indication of median sulcus, basal margin slightly produced on median fourth; punctures fine on disc, larger and
denser on lateral declivities, fineon lateral swellings. Elytra. Subquadrate, L/W = 1.0. Striae moderately wide and similarly impressed on disc and apical declivity, punctures broadly oval to rounded, as deep as stria, finelyalutaceous and strongly encroaching on intervals, 1.5-2.0 times as wide as stria and separated by 2-4 diameters on disc and apical declivity. Intervals convex, minutely punctate throughout, surface lacking any trace of alutaceous microsculp ture. Thoracic sterna. Proepisternum with posterolateral portion scarcely pubescent and alutaceous throughout. Prosternum behind procoxa with surface microsculptured through out, pubescence fine and short throughout, with few long and erect setae on disc. Mesoster num rather shallow and transversely impressed on disc, punctures moderate in size and coarsely umbilicate, dense throughout, pubescence fine and semi-erect, few longer setae along mesocoxae, surface between punctures almost glossy throughout. Mesepisternum with few moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metasternum with disc finely and unevenly punctate, foveolate on median third in males; lateral lobes with coxal margin strongly and triangularly expanded anteriorly, surface strigulate, remainder of surface on anterior half with punctures small and broadly oval, each with anterior edge carinate, with aligned row of approximately 5 transverse ridges, posterior half with punctures rounded, slightly umbili cate and mostly restricted to lateral portion, surface alutaceous posterior to triangular expansion of coxal margin, completely glossy on remainder of surface, pubescence fine and
short, apposed throughout; median lobe moderately wide, L/mW = 2.6, lateral margins divergent on anterior half, convergent on posterior half, surface with small umbilicate punctures, punctures fine on posterior third, surface almost glossy between punctures, junction with mesostemum angulate medially, finely to weakly carinate throughout. Legs. Anterior tibia quadridentate, basal tooth small, superoproximal edge unmodified, ventrome dial carina lacking intervening setae on basal two-thirds, apical spur slender and slightly inwardly bent on apical fourth. Anterior femur unmodified. Posterior femur with anterior edge produced into sharp lamellar process on basal half, distal half longitudinally striate behind anterior edge, ventral surface convex, posterodorsal edge slightly and sharply lamellate on apical fourth in males. Abdomen. Stemites 2-5 with several coarse and umbilicate punctures laterally, anterior edge with row of moderately sized, coarse and umbilicate punctures laterally, surface between punctures glossy, stemite 6 finely punctate posteriorly, with row of coarse unaligned punctures anteriorly, suture between stemites 5-6
simple. Pygidium strongly convex, W /L = 2.0, punctures fine throughout, surface between 134 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
" *
• 0. insolitus
+ 0. stridulator
* 0. erosus
MAP 33. Known distribution of Ontherus insolitus, 0. stridulator, and 0. erosus.
punctures glossy, apical margin clearly delimited. Male genitalia (Figs. 253-255). Tubus approximately 2.1 times as long as parameres, ventral portion emarginate, widely membra nous, lacking triangular swelling medially, with at most short longitudinal carina laterally. Parameres in lateral view with apex rounded, ventral portion straight, dorsal portion strongly concave. Sigmoid sclerite of internal sac as in Figure 313.
MATERIAL STUDIED. 3CJ' a (MNHN, NMPC, ZMHB).
Holotype CJ' (MNHN): Brasilia, 0./ Ex. Musreo, E. Harold/ HOLOTYPE, Ontherus stridulator, F. Genier, 1992. Dissected.
Paratypes: BRAZIL: No locality (1CJ' NMPC) (1 a ZMHB).
58. Ontherus (Ontherus) erosus Harold, 1875 comb.nov. (Figs. 256-258, 314; Map 33)
Ontherus erosus Harold, 1875: 64 (description) Ontherus erosus Harold: Gillet, 191 lb: 58 (catalog) Ontherus erosus Harold: Luederwaldt, 1930: 106 (comment) Ontherus erosus Harold: Luederwaldt, 1931: 371, 383 (key, distribution and redescription) Ontherus erosus Harold: Pessoa and Lane, 1941: 456 (key) Ontherus erosus Harold: Blackwelder, 1944: 206 (list) Ontherus erosus Harold: Lange, 1947: 310 (list)
REDESCRIPTION. Colour. Body dark reddish brown to black, lacking any trace of green metallic sheen. Pubescence fulvous. Length. 9-12 mm. Head. Anterior edge broadly arcuate, moderately upturned and slightly bidentate medially, clypeogenal junction straight. Clypeus transversely wrinkled on anterior half in males or two-thirds in females, finely punctate posteriorly. Clypeofrontal suture broadly arcuate, almost straight medially, finely tuberculate medially and laterally, median tubercle larger. Anterior genal edge slightly longer than lateral, genal surface finely punctate throughout in males or wrinkled on anterior half A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 135 and remainder of surface finely punctate in females. Vertex finely punctate excepton small area posteriorly. Pronotum. Transverse, W/L = 1.6; anterior edge narrowly membranous and slightly arcuate behind head insertion. Disc convex in both sexes, basal third with at most faint indication of median sulcus, basal margin slightly produced on median fourth; punctures fine on disc, larger and denser on lateral declivities, fine on lateral swellings. Elytra. Slightly wider than long, L/W = 0.9. Striae moderately wide, deeper on apical declivity, punctures oval, as deep as stria, alutaceous and encroaching on intervals, approxi mately 2.0 times as wide as stria and separated by 2-3 diameters on disc and apical declivity. Intervals convex, minutely punctate throughout, surface lacking any trace of alutaceous microsculpture. Thoracic sterna. Proepistemum with posterolateral portion moderately pubescent and alutaceous throughout. Prosternum behind procoxa moderately to coarsely microsculptured and scabrous throughout, pubescence fine, short and scarce throughout, with several long and erect setae on disc. Mesosternum transversely impressed on disc, punctures large and coarsely umbilicate, dense throughout, pubescence fine and semi-erect, fewlonger setae along mesocoxae and posterior portion of disc, surface between punctures finely alutaceous throughout. Mesepisternum with several moderate and vaguely to clearly delimited punctures, surface strongly alutaceous, strongly carinate parallel to posterior edge. Metastemum with disc finely and unevenly punctate, shallowly foveolate to shallowly sulcate in males, or finely and shallowly sulcate in females; lateral lobes with coxal margin strongly and triangularly produced anteriorly, surface strigulate, remainder of surface on anterior half with punctures broadly oval and umbilicate, with approximately aligned row of 6-8 transverse ridges, posterior half with punctures rounded, coarsely umbilicate and mostly restricted to lateral portion, surface between punctures alutaceous behind expanded portion of coxal margin, completely glossy on remainder of surface, pubescence fine and short, apposed throughout; median lobe wide, L/mW = 2.0, lateral margins divergent on anterior third, slightly convergent on posterior two-thirds, surface with small umbilicate punctures, surface finely alutaceous between punctures, junction with mesostemum angulate medially, weakly carinate throughout. Legs. Anterior tibia quadridentate, more slender in males, basal tooth small, superoproximal edge emarginate basally, ventromedial carina lacking intervening setae on basal two-thirds in males or third in females, apical spur slender and slightly spatulate in males and females.Anterior femurunmodified. Posterior femur with anterior edge produced into strong lo bate process on basal two-thirds, distal portion of lobate process ventrally directed, apical third of femur rather narrow and unmodified otherwise, ventral surface convex, posterodorsal edge slightly and sharply lamellate on apical third in males or femur unmodified in females. Abdomen. Sternites 2-5 with several coarse and umbilicate punctures laterally, anterior edge with row of moderately sized, coarse and umbilicate punctures laterally, surface between punctures glossy, sternite 6 finely punctate posteriorly, with row of coarse, unaligned punctures anteriorly, suture between stemites 5-6 simple. Pygidium strongly convex, WIL = 1.8 in males or 2.1 in females, punctures small throughout, surfacebetween punctures glossy, apical margin clearly delimited in both sexes. Male genitalia (Figs. 256-258). Tubus approximately 2.0 times as long as parameres, ventral portion emarginate, widely membranous, lacking triangular swelling medially, with at most short longitudinal carina laterally. Parameres in lateral view with apex obliquely truncate, ventral portion slightly concave, dorsal portion strongly concave. Sigmoid sclerite of internal sac as in Figure 314.
MATERIAL STUDIED. 8 ctct, 2 9 9 (ISNB, MNHN, NMPC, ZMHB).
TYPE MATERIAL Holotype ct (MNHN): Canta, gallo ( =Cantagalo)/ Ontherus erosus, mihi./ Ex. Musaeo, E. Harold/ Museum Paris, ex Coll., R. Oberthilr, 1952/ HOLOTYPE, Ontherus erosus, Harold, 1875, vidit F. Genier, 1991. Holotype studied. 136 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170
NON-TYPE MATERIAL Distribution: BRAZIL: Bahia: No locality. Parana: V. Ribeira*.
Additional data: Specimens collected in July.
PHYLOGENY
The presence of two derived characters in all species of Ontherus suggests that it is a monophyletic group. The firstcharacter is the presence of sensory pits on segments 7 and 8 of the antennae. None of the genera that are considered to be closely related (see introduction) share this character. However, similar sensory pits were found in one other genus, Proagoderus Van Lansberge of the tribe Onthophagini. This adaptation seems to have evolved independently at least twice in Scarabaeinae. The second character is the presence of intervening setae on the ventromedial carina of the anterior tibia. These intervening setae are not present in any of the closely related genera (see introduction). In addition, the arrangement and general shape of the sclerites of the internal sac of the aedeagus are characteristic and consistent throughout the genus. The following character analysis is based on adults. The out-group comparison is made with the genus Dichotomius. This genus has been selected because it is part of the same lineage. Because Dichotomius is such a large genus, and I have insufficient knowledge of variability within this genus, the species D. torulosus (Eschscholtz) has been specifically selected; it may be the most primitive species of the genus Dichotomius. This presumption is based on the presence of a lateral carina on the pronotum, similar to that found in the genera Copris and Coptodactyla. This carina is also clearly present in O. laminifer, a species I consider to be one of the most primitive species of Ontherus along with 0. hadros and 0. magnus. Although the two latter species do not possess a clear lateral pronotal carina, they have what seems to be a vestigial carina. In addition, their large size, the development of the secondarysexual characters, and the presence of rugosity, or granules, on the pronotum and head, strongly suggest a close relationship with D. torulosus. This relationship could also be supported by their preference for a more temperate climate. Dichotomius torulosus is distributed in subtropical regions of South America (Argentina, Chile and southeastern Brazil), and 0. hadros and 0. magnus at higher elevations in Ecuador. Ontherus laminifer is restricted in distribution to the Amazonian basin. The phylogenetic analysis (Fig. 315) was conducted using PAUP 3.1.1(Swofford 1994) and is based on 54 morphological characters(T able 1) representing 118 differentstates found in adults. Characters 37 and 44 were heavily weighted, as I believed they evolved only once. The subgeneric concept of the genus is based mainly on these two characters. Characters 7, 8 and 32 are multi-state and ordered (012). All other multi-state characters are unordered. Characters are coded in Table 2. The analysis was replicated 15 times. A total of 1302 equally parsimonious trees were obtained. The strict consensus tree presented in Figure 315 was obtained from the 1302 shortest trees. The resulting consensus tree shows that the common ancestor of Ontherus split into two lineages. The first lineage is the subgenus Planontherus with two species and the second lineage includes the subgenera Caelontherus and Ontherus s.str. with the remaining 56 species. The second branching is a trichotomy comprising the subgenera Caelontherus, Ontherus s.str. and the Brevicollis species group. The subgenus Caelontherus is para phyletic, with three different lineages and the subgenus Ontherus s.str. is monophyletic. I consider the subgenus Caelontherus to be monophyletic despite results of the analysis based on characters in Table 1. The apparent paraphyletic relationship is based on characters relating to the legs and secondary sexual characters and therefore strongly subject to external pressure. Character 44, the shape of the mesometasternal suture, which is believed to be less subjected to external pressure, should be more heavily weighted to show a more natural relationship. The species group Kirschii is linked to the subgenus Ontherus s.str. (Fig. 315, A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 137
CHARACTER Plesiotvp1c character slate (0) Aootvoic character state 11) Aootvoic character state 12) GENERAL 1 averane bodv size moderate to laroe small 2 location of male secondarv sexual characters mostlv on dorsum mostlv on ventrum 3 color of dorsum lackJna iridescent sheen with some iridescent sheen 4 color of dorsum lackino metallic sheen with some metallic sheen HEAD 5 clvoeofrontal carina of well--Oevelooed males Iunordered) simolv tuberculate medially median tubercule modified lackino median tubercule 6 clvoeofrontal tubercule of females I unordered\ tubercule transverse or double tubercule conical tubercule lackina 7 civoeo-frontal suture of females rordered\ suture obsolete suture arcuate or rounded suture traoezoidal or sauare 8 shaoe of cJvn<>nfrontal carina of females • !ordered\ carina absent or interrupted medially carina more or less even carina uneven or tuberculate laterallv 9 male internal ocular area unmodified strongly depressed 10 antennnal seamen! 7 and a lackina sensorv oits with sensory pits PRONOTUM 11 surface of lateral declivities of oronotum nranulate ounctate 12 develooment of sulcus (unordered) with a noorlv defined sulcus sharolv defined sulcus 13 lateral swellinns vestiaial sulcus no su!cus 14 anterior edae of well-develoned males more or less strainht deeotv emaroinate behind eves ELYTRA 15 stnal ounctures (unordered) absent or ooorlv defined oval and well-defined rounded and well-defined 16 elvtral striae similartv imoressed throughout deeper on apical declivity 17 elvtral striae on humeral and aoica! cali normallv imoressed weakly impressed or obsolete 18 basal oortion of elvtra simole rnarginate 19 ounctures of elvtral intervals (unordered\ ill-defined minute and sharply defined Jamer and conspicuous 20 elvtral surface smooth to finelv alutaceous strongly alutaceous PYGIDIUM 21 ovoidial marain in males well-defined obsolete 22 ovoidial maro1n in females well-defined obsolete 23 basal oortion of ovaidium unmodified or finelvsulcate sulcate alona marain TARSI 24 tarsal claws unmodified enlaroed and J-shaoed 25 tarsal claws unmodified retractable 26 aoicoventral proiection of last tarsal S""ment membranous and flattened sclerotized and lateralv compressed FORE LEGS 27 inferomed1al carina lackina intervenina seta wi1h 1nterveninq seta 28 basoventral oortion of front femur in males " == sublet to homonlasv TABLE I. Analysis of character transformation in Ontherus relationship A). This relationship is exclusively based on the homoplasious character 34 and therefore the relationship shown in B (Fig. 315) should be viewed as valid. In the subgenus Ontherus s.str., 0. irinus is isolated from the groups Appendiculatus and Brevipennis. I consider 0. irinus to be somehow intermediate between the two aforementioned groups (see remarks under 0. irinus). The broadly arcuate mesometastemal suture is responsible for its isolated position in the subgenus. It is possible that the arcuate suture is a reversal. Except forcharacter 44, this species is closely related to the Appendicu latus and Brevipennis groups. w CHARACTER 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 00 1. 0. hadros sp. nov 0 0 0 0 1 1 1 0 0 1 0 0 1 0 1 0 0 1 1 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 2. 0. magnus sp. nov. 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 1 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 3. 0. laminifer Ballhasar 1 0 0 0 1 2 1 0 0 1 0 0 0 1 1 0 0 1 2 1 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 4. O. gilli sp. nov 0 1 0 0 0 1 1 0 0 1 1 0 1 0 2 0 0 1 1 1 0 0 1 0 0 0 1 0 2 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 5. 0. ashei sp. nov 0 1 0 0 0 ? ? ? 0 1 1 0 1 0 1 0 0 1 1 1 0 ? 1 0 0 0 1 0 2 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 6. 0. brevicollis Kirsch 0 1 0 0 0 1 1 0 0 1 1 0 1 0 1 0 0 1 1 1 0 0 1 0 0 0 1 0 2 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 7. 0. politus sp. nov. 0 1 0 0 0 1 1 0 0 1 1 0 1 0 2 1 0 1 1 1 0 0 1 0 0 0 1 0 2 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 8. 0. aequatorius Bates 0 1 0 0 0 1 1 0 0 1 1 0 1 0 2 1 0 1 1 0 0 0 0 0 0 0 1 0 2 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 9. O. kirschi1 Harold 0 1 1 0 0 1 1 0 0 1 1 0 1 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 2 0 0 0 0 1 1 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 10. 0. sanctaernartae sp. nav. 0 1 0 0 0 1 1 0 0 1 1 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 1 1 2 0 0 0 0 1 1 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 11. 0. alexis (Blanchard) 0 0 1 0 0 0 1 0 1 1 1 0 1 1 0 0 0 1 1 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 12, 0. tenuistrlatus sp. nov 0 0 0 0 1 0 1 0 1 1 1 0 1 1 1 1 0 1 1 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 13. 0. obliquus sp. nov. 0 0 1 0 1 0 1 0 1 1 1 0 1 1 1 1 0 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 14. 0. incisus (Kirsch) 0 0 0 0 1 0 1 0 1 1 1 a 1 1 1 0 1 1 1 1 a 0 0 a 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 a 0 0 0 0 0 0 0 0 0 0 0 0 0 0 s:;:: 15. 0. howdeni sp. nov 0 0 0 0 1 0 1 0 1 1 1 0 1 1 1 0 0 1 1 0 0 a 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 1 a 0 0 a 0 0 0 0 a 0 0 0 0 a 0 16. 0. diabolicus sp. nov 0 0 1 0 1 0 1 0 1 1 1 a 1 1 1 a 0 1 1 1 0 0 0 0 a 0 1 0 1 a a 0 0 0 0 a 1 a 1 a a 0 0 a 0 0 0 0 0 0 a 0 0 0 � 1 0 17. 0. compress!cornis Luederwaldt a 0 0 0 1 1 1 0 1 1 1 a 1 1 2 0 0 1 0 0 a 0 0 0 0 1 0 1 a 0 0 0 a 0 0 1 1 1 a a a a 0 0 0 0 0 0 1 0 0 0 0 � 1 1 1 1 0 1 1 2 1 0 1 1 0 0 0 0 0 0 0 1 1 0 a 0 a 0 1 1 1 0 a 0 0 0 0 0 0 0 1 0 0 0 0 18. 0. pilatus sp. nov 0 0 0 0 1 0 1 0 0 0 0 r.n 19. 0. tri!uberculatus Balthasar a 0 1 0 1 a 1 a 1 1 1 a 1 1 2 1 a 1 1 0 0 0 0 0 a 0 1 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 a 0 0 0 20. Q_ mexicanus Harold 0 0 1 0 1 a 1 0 1 1 1 0 1 1 2 1 a 1 1 1 0 0 0 0 0 0 1 0 1 0 0 0 a 0 0 a 1 a 1 0 0 a 0 0 0 0 0 0 0 1 0 0 0 0 � 21 . 0. sextubercuJ:atus sp. nov a 0 1 0 1 0 1 0 1 1 1 0 1 1 2 1 0 1 1 1 0 0 0 a 0 0 1 0 1 a a 0 0 0 0 0 1 0 1 0 0 a a 0 0 0 0 0 0 1 0 0 0 0 22. 0. lunicollis sp. nov 0 0 1 0 1 0 1 0 1 1 1 a 1 1 2 1 0 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 a 0 a 0 1 1 1 0 a 0 1 0 0 0 a 0 0 1 0 0 0 0 � 23. 0. monilistnatus sp. nov. 0 0 1 a 1 a 1 0 1 1 1 0 1 1 2 1 0 1 1 0 a 0 0 a 0 0 1 0 1 0 0 0 0 0 0 a 1 1 1 a a 0 0 0 0 0 a 0 0 1 0 0 0 0 24. 0. pseudodidyrnus sp. nov. a 0 1 0 1 a 1 0 1 1 1 0 1 1 1 1 0 1 1 1 0 0 0 a 0 0 1 0 1 a 0 0 0 0 a 0 1 a 1 a 0 0 1 0 0 0 0 0 0 1 0 0 0 0 ti 25. 0. bridgesi 0 1 0 a 0 1 1 0 0 1 1 0 1 a 0 0 0 1 1 1 0 1 1 0 0 0 1 0 2 a 0 a 0 0 0 0 0 0 1 0 0 0 0 0 a 1 1 0 0 0 0 0 a 0 d ? ? ? 26. 0. rectus sp. nov 0 ? a 0 1 1 0 0 1 1 a 1 0 0 0 0 1 1 1 ? 0 0 0 0 0 1 ? 0 a 0 ? ? 0 a 0 1 a a 0 0 0 a 1 1 0 0 0 0 0 0 0 27 _ 0. pubens sp. nov. 0 1 0 0 0 1 1 0 0 1 1 a 1 0 0 0 0 1 1 1 a 0 a 0 0 0 1 0 2 0 0 1 a 0 a a 1 0 1 0 0 0 a 1 0 1 1 0 0 0 0 0 0 0 28 0. sulcator (Fabric1us) a 1 a a a 1 1 0 a 1 1 0 1 0 0 0 0 1 1 1 0 0 0 a 0 0 1 0 2 0 0 1 0 0 0 1 1 a 1 a 0 0 a 1 0 1 1 a 0 0 0 0 0 0 § 29. 0. appendiculalus (Mannerheim) a 1 0 0 a 1 1 0 0 1 1 1 1 0 1 0 0 1 1 1 0 0 0 0 a 0 1 a 2 1 a 1 1 1 1 0 1 a 1 a 0 0 0 1 1 1 1 a 0 a a 1 0 0 8 30. 0. ulcopygus sp. nov. 0 1 0 0 0 1 1 0 0 1 1 1 1 0 1 0 0 1 1 1 1 1 0 0 0 0 1 0 2 1 0 1 1 1 1 0 1 0 1 0 a 0 a 1 1 1 1 0 0 0 0 1 0 0 (") 31. 0. podiceps Harold 0 1 0 0 0 1 1 0 0 1 1 1 1 0 1 0 0 1 1 1 0 1 0 0 0 0 1 0 2 1 a 1 1 1 1 0 1 0 1 a 0 0 0 1 1 1 1 0 a a 0 1 a 0 > r 32. 0. 0 1 0 a 0 1 1 0 0 1 1 1 1 0 1 a 0 1 1 1 0 1 1 0 a a 1 0 2 1 1 1 1 1 1 a 1 1 1 a a 0 0 1 a 1 1 a 0 a 0 1 a 0 azteca Harold r.n 33. O. cambeforti sp. nav. 0 1 0 0 0 1 1 0 a 1 1 1 1 0 1 0 0 1 1 1 0 0 0 0 0 a 1 0 2 1 1 1 1 1 1 0 1 1 1 0 0 0 0 1 0 1 1 0 0 a 0 1 0 0 0 34. O. grandis Luederwaldt 0 1 a 0 1 0 1 a 0 1 1 0 1 1 0 0 0 1 1 1 1 1 0 0 0 0 1 0 2 1 a 0 0 1 0 0 1 0 0 1 0 0 a 1 1 1 1 0 1 0 0 1 0 0 (") 35. 0. 0 1 0 0 2 2 1 1 a 1 1 1 1 0 1 0 0 1 1 1 0 1 0 0 0 0 1 a a 1 0 1 1 1 1 a 1 0 1 a a 0 0 1 1 1 1 0 0 0 0 1 0 0 m cariritfrons Luederwaldt -l 0 1 1 0 36. 0. virescens (Lucas) 1 1 0 1 2 2 1 1 a 1 1 0 1 0 1 a 1 1 0 1 0 0 1 a 0 1 0 1 a 1 a a 1 a 1 0 a 0 a 1 1 1 1 a 0 0 0 1 0 0 >-< 37. 0. irinus Ballhasar a 1 1 0 2 2 1 1 0 1 1 1 1 0 1 0 a 1 1 1 a a a 0 a 0 1 0 0 1 0 1 1 1 0 0 1 0 1 0 0 a 0 0 a a 0 0 0 a 0 0 1 a ? � 38. 0. raplor sp. nov. 0 1 0 a 2 ? 7 ? 0 1 1 a 1 0 1 0 a 1 1 1 0 1 a 1 1 1 1 2 a 0 1 1 1 1 a 1 a 1 0 ? 0 0 1 a 1 1 0 0 a 0 a 1 0 39. 0. 0 1 0 a 2 2 2 1 0 1 1 a 1 0 1 0 0 1 1 1 0 0 1 a 1 1 1 1 2 a 0 1 1 1 1 0 1 0 1 0 0 0 0 1 0 1 1 a a 0 0 0 1 0 (") planus sp. nov. > 40. 0. lichyi Martinez 0 1 0 0 2 2 2 2 0 1 1 0 1 0 1 0 0 1 1 1 a a 1 a 0 1 1 1 2 1 0 1 0 1 1 a 1 0 1 a a 0 0 1 0 1 1 0 0 0 0 0 1 0 41. 0. edentulus sp. nov 0 1 a 0 2 2 2 2 0 1 1 a 1 0 1 0 0 1 1 1 a 0 1 0 0 1 1 1 1 0 0 1 0 1 1 0 1 0 1 0 0 0 a 1 a 1 1 0 0 0 0 0 1 0 � 42. 0. 0 1 a 0 2 2 2 2 0 1 1 0 1 0 1 a 0 1 1 1 0 0 1 0 1 1 1 1 2 1 0 1 a 1 0 0 1 0 1 0 0 0 0 1 0 1 1 0 0 0 0 0 1 0 tJ brevipennis Harold > 43. 0. a 1 a 0 2 2 2 2 0 1 1 0 1 0 1 0 a 1 1 1 0 0 1 0 1 1 1 1 2 1 0 2 1 1 1 0 1 0 1 a 1 a 0 1 0 1 1 0 0 0 a a 1 0 rectangul1dens sp. nov. z 1 1 1 1 1 2 1 1 0 a 1 44. 0. carinicat!is Luederwaldt a 1 a a 2 2 2 2 0 1 1 a 1 0 1 0 a 1 0 0 a 1 2 1 0 2 1 1 0 1 0 0 1 a 1 1 0 0 0 0 0 1 0 9 45. 0. zikani Luederwaldt a 1 0 0 2 2 2 1 a 1 1 0 1 0 1 0 0 1 1 1 a a 1 a 1 1 1 2 2 1 0 2 1 1 1 a 1 a 1 a 1 0 a 1 0 1 1 0 0 0 0 0 1 0 46. 0. 0 1 0 0 2 2 1 1 0 1 1 0 1 0 1 a 0 1 1 1 0 0 1 0 1 1 1 1 2 1 0 2 a 1 1 a 1 0 1 0 1 0 a 1 0 1 1 0 0 0 0 0 1 0 '.::3 lobifrons sp. nov. 0 47. O. cephalotes Harold 0 1 0 0 2 2 1 2 0 1 1 0 1 0 1 a 0 1 1 1 a 0 1 0 1 1 1 1 2 1 a 2 a 1 1 0 1 a 1 0 1 0 0 1 0 1 1 0 0 0 0 0 1 0 48. O. aphodioi TABLE 2. Character state matrix A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 139 BIOGEOGRAPHY The genus Ontherus comprises 58 species distributed from Mexico in the north to Argentina in the south. Collectively species of Ontherus occur at various elevations and in various habitats but middle to low elevations and wet tropical and subtropical habitats predominate. As noted in the introduction, Ontherus belongs to a lineage of large tunnelling Scarabaeinae, whose constituent genera are primarily found in Africa south of the Sahara Desert (e.g. Copris, Catharsius, Xinidium) and in Australia (e.g. Coptodactyla, Thyregis). The 58 species of Ontherus are grouped in three subgenera and 11 species groups. Based on the phylogenetic analysis presented herein, the subgenus Planontherus, comprising two species, 0. bridgesi and 0. rectus, is the most primitive lineage within the genus. The two remaining subgenera constitute a monophyletic lineage comprising the remaining 56 species. These subgenera are the paraphyletic Caelontherus with 24 species in six species groups and the monophyletic Ontherus s.str. with 32 species in five species groups (Fig. 315). The two species placed in Planontherus are distributed at middle to higher elevations in the southern Andes. Ontherus bridgesi is known from Argentina, Bolivia and Peru (Map 12) and 0. rectus is known from a single locality in northwestern Peru. Both species are associated with temperate montane forests. Within Caelontherus, the most primitive lineage, the Hadros group, comprises two relict species both of which are found at higher elevations (2600-2700 m) in Ecuador, in cool montane subtropical forest. Additionally, most of the species comprising the relatively primitive Alexis and Mexicanus groups within Caelontherus are usually found at elevations above 1000 min various forest types. Based on this evidence, Ontherus is hypothesized to have differentiated from a Gondwanian ancestor inhabiting cool subtropical forest (or an equivalent habitat) soon after the separation of South America, Africaand Australia approxi mately 60 million years before present (B.P.). Following the early evolution of the most primitive subgenus Planontherus, the second major event is the differentiation of the two more diverse subgenera, Caelontherus and Ontherus s.str. Both these subgenera are very successful and much more diverse than Planontherus. As noted, Caelontherus is paraphyletic and structurally appears to be more primitive than the monophyletic Ontherus s.str. Based on current distributions of species of Caelontherus, most of which are restricted to South America, it appears that species within this subgenus have radiated northward along the Andes. Diversity of species in the subgenus can be explained by the successive uplift of the biotic zones during hot-humid/cold-dry cycles, which could have caused range disjunction (Vuilleumier 1971). These uplifts started during the upper Pliocene and the Lower Pleistocene (Vuilleumier 1971). The Brevicollis group, composed of five closely related species, is distributed in the Andes north of Peru. The only species found south of Ecuador is 0. ashei, which is known from northern Peru. Their restricted current distribution and general similarity suggest that this group of species evolved recently. The Kirschii group is composed of only two closely related species and is restricted to the Colombian and Venezuelan Andes ( 0. kirschii) and the Santa Marta mountains of northern Colombia (0. sanctaemartae). This allopatric distribution pattern suggests that the common ancestor of 0. kirschii and 0. sanctaemartae was widely distributed in Colombia. The Alexis group has a wider distribution. It occurs in the Andes from Bolivia to Colombia. Five of the six species are present in Peru. A similar situation applies forthe species of the Mexicanus group that are restricted to South America. Based on this observation, it is suggested here that the Peruvian Andes are the centre of origin for both the Alexis and Mexicanus groups. The species of both Alexis and Brevicollis groups are phylogenetically closely related and are thought to have evolved relatively early from a common ancestor. This assumption is supported by the relatively wide distribution of both species groups. 140 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 Ontherus laminifer is one of the only two species of the subgenus Caelontherus not restricted to higher elevations; its current distribution range is the Amazon basin. Ontherus diabolicus is also known from lower elevations in the Amazon basin, however, it differs from 0. laminifer in occurring at higher elevations as well. The subgenus Caelontherus apparently expanded its range northward through the newly formed Panamanian land bridge during the late Tertiary. The known distribution of species suggests that only 0. pseudodidymus reached Central America initially. This event was followed by the invasion of a second lineage that was subsequently isolated into two populations: one in Mexico and Guatemala (0. mexicanus) and one in Costa Rica (0. sextuberculatus). The third subgenus (Ontherus s.str.) appears to have evolved adaptations to warmer tropical habitats. With few exceptions (discussed below), the entire group is restricted to elevations below 1000 m. It is believed that ancestral species dispersed eastward and northward from a hypothetical centre of origin in Bolivia and northern Argentina. The great est number of species occur in the Planalto of southern Brazil. This is a well-documented centre of endemism and is a presumed refuge fordry epochs during the Pleistocene (Haffer 1982). The Sulcator group is distributed from middle to low elevations east of the Andes. Ontherus pubens is distributed over the western portion of the range and 0. sulcator over the eastern portion. If specimens studied are indicative of the entire distribution, it appears that 0. sulcator is restricted to wet tropical and subtropical habitats. These two closely related species are both present in Bolivia and northwest Argentina. It is possible that each species occupies a different ecological niche. The Appendiculatus group is widely distributed in Central and South America. Ontherus grandis is restricted to middle elevations in northern Argentina. This possibly relictual distribution pattern combined with its phylogenetic isolation suggest that it is one of the most primitive species of the Appendiculatus group. Ontherus appendiculatus has a distribution similar to 0. sulcator, however it is present in the drier region of coastal Brazil. Ontherus azteca is distributed along the Andes from Mexico to Bolivia and at lower elevations in Paraguay and Brazil. This distribution is unique in the genus. It suggests that 0. azteca is perhaps the only subtropical species of the subgenus. Its sister species, 0. cambeforti, is restricted to the Guyana shield. It is possible that the common ancestor of 0. azteca and 0. cambefor ti entered the Guyanas through Venezuela during a cold episode of the Pleisto cene. Other species belonging to the Appendiculatus group are restricted to the Brazilian Pianalto region, except for 0. carinifrons which with 0. laminifer is the only truly · Amazonian species. The species belonging to the Brevipennis group, which are associated with ants, are hypothesized to be speciating rapidly, based on their close relationship, supported by their very similar aedeagus and habitus. They represent 17% of the diversity in the genus and evidence indicates that they are one of the most derived groups. Their current distributions suggest that they evolved from a common ancestor in southeastern Brazil. I suggest that association with ants was the key event facilitating expansion of their range to the north. This hypothesis is based on the fact that this group is using the refuse piles of leafcutter ants as a food source (Gill 1991), a relatively competition free and abundant food which has allowed them to expand their distribution quite extensively. Two species have moved into higher elevation in the Andes: 0. edentulus is found exclusively in Ecuador and Peru and its sister species 0. lichyi is found at middle to low elevations in Colombia, French Guiana and Venezuela (a southeastern Brazilian record needs to be confirmed). A third species, 0. brevipennis, is found in Central America north to the lowlands along the Nicaragua-Costa Rica border. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 141 The remaining two species groups (Digitatus and Erosus groups) are restricted to southern Brazil and northern Argentina. The only exception is 0. aphodioides with a range extending northward to Venezuela. It is difficult to explain why such a successful genus in the Neotropics, with several species present at higher elevations, failed to expand its range into southern parts of the Nearctic region. For example the subgenus Caelontherus, because it is primarily associated with cooler habitats, is a possible candidate. Species of the subgenus Caelontherus should have had sufficient time to invade the Nearctic, considering that at least four other Neotropical genera of dung beetles (Ateuchus, Canthon Hoffmannsegg, Delto chilum Eschscholtz and Dichotomius) have successfully invaded North America. I consider that the most probable explanation is competition. The overall comparable shape and feeding habits of Ontherus (Caelontherus) and Copris suggest that Ontherus (Caelontherus) is probably the Neotropical, ecological equivalent of Copris. Possibly Copris was already well established in North America when Ontherus (Caelontherus) could have colonized the Nearctic region. This assumption is supported by the presence of several (over 15) species of Copris in Central America. The hypothesis of mutual exclusion is further supported by the factthat Copris has similarly never entered South America, with the exception of a single species C. incertus Say known from higher elevations in Colombia and Ecuador. ACKNOWLEDGMENTS I thank all of the curators and persons in charge of the loan of insects from the various institutions and private collections cited in the "Material" and "Methods" sections. I am especially grateful for their patience in allowing me to borrow this material for so long. I also thank the following individuals for their assistance and important contributions toward several aspects of the preparation of this work: Robert Anderson, Ed Becker, Yves Bousquet, Henry Howden, Jan Klimaszewski, Bernard Landry, Serge Laplante, Ales Smetana and one anonymous reviewer. Special thanks also to Nancy Boase for her invaluable help with the English, Bruce Gill and William Warner for testing and improving the key, Go Sato for the habitus drawing and inking Figures 21-55, Angel Solis B. fortranslating the key to Spanish, Yves Cambefort for his assistance during my visits at the Museum national d'Histoire naturelle (Paris), the Canadian Museum of Nature staff, where part of this work was conducted, and to my family, Isabelle, Simon and Julien, for their patience during countless hours spent at home completing this revision. This study has been made possible through financial support from the Canadian Museum of Nature. REFERENCES Arnett, R.H., G.A. Samuelson, and G.M. Nishida. 1993. The Insect and Spider Collection of the World, 2nd ed. Flora & Fauna Handbook 11: 310 pp. Gainesville, FL. Balthasar, V. 1938. Neue Gattungen und Arten der siidamerikanischen Coprophagen. (4. Beitrag zur kenntnis der Scarabaeiden der neotropischen Region.). Entomologische Btatter 34: 210-223. --1963. Monographie der Scarabaeidae und Aphodiidae der Palaearktischen und Orientalischen Region (Coleoptera: Lamellicomia), vol. 1-3. Verlag Tschechoslowakische Akademie der Wissenschaften, Prague. Bates, H.W. 1887. Biologia Centrali-Americana, Coleoptera, vol. 2, pt. 2, pp. 25-160. --1891. Coleoptera. pp. 7-39 in Whymper, Supplementary Appendix to Travels amongst the Great Andes of the Ecuador. London. Blackwelder, R.E. 1944. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 2. Bulletin of the United States National Museum 185: 189-341. Blanchard, C.E. 1845 (1843). In Brulle, Insectes de l'Amerique Meridionale recueillis par Alcide d'Orbigny, vol. 6, pt 2, Paris. 222 pp. Bruch, C. 1911. Catalogo sistematico de los cole6pteros de la Republica Argentina. Revista def Museo de La Plata. IV 17: 181-225. --1915. Suplemento al catalogo sistematico de los cole6pteros de la Republica Argentina I (Addenda, corrigenda y resumen). Revista def Museo de La Plata 19(2): 538-573. Burmeister, H.C.C. 1874. Lamellicomia Argentina. Stettiner Entomologische Zeitung 35: 120-133. 142 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 Erichson, W.F. 1847. Conspectus insectorum coleopterorum quae in Republica Peruana observata sunt. Archiv fur Naturgeschichte 13: 67-185. ---1848a. Die Insekten in Schomburg's Reise in Guiana, vol. 3, pp. 533-617. Leipzig. ---1848b. Naturgeschichte der Insecten Deutschlands, Abt. 1, Coleoptera, vol. 3, Lief 5, 968 pp. Berlin. Fabricius, J.Ch. 1775. Systema entomologiae, 30 + 832 pp. Lipsiae. ---1792. Entomologia systematica, vol. 1, 330 + 538 pp. Hafniae. ---1801. Systema eleutheratorum, vol. 1, 506 pp. Kiliae. Gemminger, M., and E. Harold. 1869. Catalogus coleopterorum hucusque descriptorum synonimicus et systematicus. vol. 4, pp. 979-1346. Monachii. Gill, B.D. 1991. Dung beetles in tropical American forests. pp. 211-229 in Hanski, I., and Y. Cambefort (Eds.), Dung Beetle Ecology. Princeton University Press, Princeton. 481 pp. Gillet, J.J.E. 1910. Especes nouvelles du genre Copris et releve synonymique des especes decrites ace jour. Notes from the Leyden Museum 32: 1-31. ---191la. Lamellicornes coprophages nouveaux ou peu connus d'Amerique du Sud. Annales de la Societe Entomologique de Belgique 55: 315-319. ---191lb. Coleopterorum catalogus, pars 38, Scarabaeidae: Coprinae I, 19: 1-100. Haffer, J. 1982. General aspect of the refuge theory. pp. 6-24 in Prance, G.T. (Ed.), Biological Diversification In The Tropics. Proceedings of the Fifth InternationalSymposium of the Association for Tropical Biology, Held at Macuto Beach, Caracas, Venezuela, February 8-13, 1979. Columbia University Press, New York. XV+ 681 pp. Halffter, G., and W.D. Edmonds. 1982. The Nesting Behavior of Dung Beetles (Scarabaeinae). An Ecological and Evolutive Approach. Instituto de Ecologia, Mexico, D.F. 176 pp. Hanski, I., and Y. Cambefort. 1991. Dung Beetle Ecology. Princeton University Press, Princeton. 481 pp. Harold, E. von. 1867. Diagnosen neuer Coprophagen. Coleopterologischte Hefte 2: 94-100. ---1868a. Diagnosen neuer Coprophagen. Coleopterologischte Hefte 3: 80-86. -- -1868b. Diagnosen neuer Coprophagen. Coleopterologischte Hefte 4: 79-86. ---1869a. Ueber Coprophage Lamellicornien mit besondered Beriicksichtigung der Pariser Sammlung. Coleopterologischte Hefte 5: 46-70. ---1869b. Diagnosen neuer Coprophagen. Coleopterologischte Hefte 5: 95-104. ---1869c. Note sur quelques coprides du Mexique. Annales de la Societe Entomologique de France, serie 4 9: 493-512. ---1875a. Verzeichniss der von Dr. Teuscher in Cantagallo gesammelten coprophagen Lamellicornien. Coleopterologischte Hefte 13: 57-72. ---1875b. Miscellen. Coleopterologischte Hefte 13: 181. ---1880. Verzeichniss der von E. Steinheil in Neu-Granada gesammelten coprophagen Lamellicornien. Stettiner Entomologische Zeitung 41: 13-46. Howden, H.F., and O.P. Young. 1981. Panamanian Scarabaeinae: Taxonomy, distribution, and habits (Coleoptera, Scarabaeidae). Contributions of the American Entomological Institute 18(1): Ill+ 204 pp. Kirsch, T.F.W. 1871. Beitrage zur Kaferfauna von Bogota. Berliner Entomologische Zeitschrift 14: 337-378 3 [(353)-384, 369-378] . Lange, R.B. 194 7. Ensaio da zoogeografiados Scarabaeidae do Parana com algumas notas eto-ecol6gicas. Arquivos do Museo Paranaense 6: 305-315. Laporte, F.L. 1840. Histoire naturelle des insectes coleopteres, vol. 2. Paris. 564 pp. Lucas, P.H. 1857. Entomologie. In Animaux nouveaux ou rares recueillis pendant !'expedition dans les parties centrales de l'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima au Para; executee par ordre du Gouvernement Frarn,;ais pendant les annees 1843 a 1847, sous la direction du Compte Francis de Castelnau, 204 pp., 18 pls. Paris. [According to Sherborn and Woodward= 1859.] Lucas, R. 1920. Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarum totius (famil., trib., subtrib., sect. incl.), Paris I. Berlin. 696 pp. Luederwaldt, H. 1930. Especies novas de Ontherus (Coleoptera, Scarabaeidae, Coprinae). Archivos do lnstituto Biol6gico, Departemento da Defensa, Sanitaria de Agricultura 3: 105-108. ---1931. 0 genero Ontherus (Coleop.) (Lamellic.-Coprid.-Pinot.) com uma chave, para a determina<,;iio dos pinotides americanos. Revista do Museu Paulista 17: 363-422. Mannerheim, C.G. 1829. Description de quarante nouvelles especes de scarabeides du Bresil avec figures. Memoires de la Societe lmperiale des Naturalistes de Moscou 7 (Nouveau Memoires 1): 29-80. Martinez, A. 1947. Lamellicomia neotr6pica I. Arthropoda 1(1): 41-53. -- -1959. Catalogo de los Scarabaeidae Argentinos (Coleoptera). Revista del Museo Argentina Ciencias Naturales "Bernardino Rivadavia" 5(1): 1-130. 3 The pagination is incorrect. The first page of Kirsch's paper is not numbered (the preceding page is correctly numbered as 337), but th.e second, third, fourth, etc. pages are numbered as 354-384, then the pagination change to 369-378. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 143 Martinez, A., and F.S. Pereira. 1960 (1959). Algunos interesantes Coprini neotropicales (Col. Scarabaeidae). Revista de la Sociedad Entomologica Argentina 22(1-4): 77-84. Olivier, A.G. 1789. Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur description, leur synonymie, et leur figure enluminee. Coleopteres, vol. I, [each genus treated is separately paginated]. Paris. ---1790. Encyclopedie methodique, Histoire naturelle. Insectes, vol. 5 (part 1), pp. 1-368. Paris. Paulian, R. 1938. Contribution al 'etude des canthonides americains (Coleopteres. Lamellic). Anna/es de la Societe Entomologique de France 107: 213-296. Pereira, F.S. 1955. Especies novas de scarabaeidae Brasileiros. Arquivos do Museu Nacional, Rio de Janeiro 42(2): 459-466. Pereira, F.S., and M.A.V. d'Andretta. 1955. Novos Escarabeideos e novas sinonimias (Col. Scarabaeidae). Papeis Avulsos do Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo 12(11 ): 247-264. Pereira, F.S., and A. Martinez. 1960. Notas Escarabeidol6gicas - II. Revista Brasileira de Entomologfa 9: 37-55. Pessoa, S.B., and F. Lane. 1941. Cole6pteros necr6phagos de interesse medico-legal. Ensaio monografico s6bre a famflia Scarabaeidae de Sao Paulo e regioes vizinhas. Arquivos de Zoolog[a do Estado de Sao Paulo 2: 389-504. Roze, J.A. 1955. Lista preliminar de la familia Scarabaeidae sensu lato (Coleoptera) de Venezuela. Boletin de! Museo de Ciencias Natura/es, Caracas 1(1): 39-63. Swofford, D.L. 1994. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1.1. Computer program distributed by the Illinois Natural History Survey, Champaign, Illinois. Vuilleumier, B.S. 1971. Pleistocene changes in the fauna and flora of South America. Science 173: 771-780. Waterhouse, C.O. 1891. New Scarabaeidae in the BritishMuseum: A fourth contribution. The Annals and Magazine of Natural History, including Zoology, Botany, and Geology ser. 6 7: 348-363. (Date received: 31May 1995; date accepted: 25 January 1996) 144 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 CHECKLIST OF THE GENUS ONTHERUS Ontherus Erichson, 1847 Subgenus Caelontherus nov. Hadros group 1. Ontherus (Caelontherus) hadros sp.nov. 2. Ontherus (Caelontherus) magnus sp.nov. Laminifer group 3. Ontherus (Caelontherus) laminifer Balthasar, 1938 comb.nov. Brevicollis group 4. Ontherus (Caelontherus) gilli sp.nov. 5. Ontherus (Caelontherus) ashei sp.nov. 6. Ontherus (Caelontherus) brevicollis Kirsch, 1871 comb.nov. 7. Ontherus (Caelontherus) politus sp.nov. 8. Ontherus (Caelontherus) aequatorius Bates, 1891 comb.nov. Kirschii group 9. Ontherus (Caelontherus) kirschii Harold, 1867 comb.nov. Ontherus elongatus Waterhouse, 1891 10. Ontherus (Caelontherus) sanctaemartae sp.nov. Alexis group 11. Ontherus (Caelontherus) alexis (Blanchard), 1845 comb.nov. Ontherus didymus Erichson, 1847 Ontherus glaucinus Erichson, 1847 Ontherus nevinsoni Waterhouse, 1891 12. Ontherus (Caelontherus) tenuistriatus sp.nov. 13. Ontherus (Caelontherus) obliquus sp.nov. 14. Ontherus (Caelontherus) incisus (Kirsch), 1871 comb.nov. Ontherus thoracicus Waterhouse, 1891 15. Ontherus (Caelontherus) howdeni sp.nov. 16. Ontherus (Caelontherus) diaholicus sp.nov. Mexicanus group 17. Ontherus (Caelontherus) compressicornis Luederwaldt, 1931 comb.nov. 18. Ontherus (Caelontherus) pilatus sp.nov. 19. Ontherus (Caelontherus) trituberculatus Balthasar, 1938 comb.nov. 20. Ontherus (Caelontherus) mexicanus Harold, 1868 comb.nov. 21. Ontherus (Caelontherus) sextuberculatus sp.nov. 22. Ontherus (Caelontherus) lunicollis sp.nov. 23. Ontherus (Caelontherus) monilistriatus sp.nov. 24. Ontherus (Caelontherus) pseudodidymus sp.nov. Subgenus Planontherus nov. 25. Ontherus (Planontherus) hridgesi Waterhouse, 1891 comb.nov. 26. Ontherus (Planontherus) rectus sp.nov. Subgenus Ontherus s.str. Sulcator group 27. Ontherus (Ontherus) pubens sp.nov. 28. Ontherus (Ontherus) sulcator (Fabricius), 1775 comb.nov. Copris nisus Laporte, 1840 Copris ovalipennis Blanchard, 1845 Copris senegalensis Gillet, 1910 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 145 Appendiculatus group 29. Ontherus (Ontherus) appendiculatus (Mannerheim), 1829 comb.nov. Copris polynice Blanchard, 1845 Copris rotundatus Blanchard, 1845 Ontherus quadratus Erichson, 1848a Ontherus contractus Burmeister, 1874 30. Ontherus (Ontherus) ulcopygus sp.nov. 31. Ontherus (Ontherus) podiceps Harold, 1868 comb.nov. 32. Ontherus (Ontherus) azteca Harold, 1869 comb.nov. Ontherus villosus Luederwaldt, 1930 Ontherus strius Howden and Young, 1981 33. Ontherus (Ontherus) cambeforti sp.nov. 34. Ontherus (Ontherus) grandis Luederwaldt, 1931 comb.nov. 35. Ontherus ( Ontherus) carinifrons Luederwaldt, 1930 comb.nov. 36. Ontherus (Ontherus) virescens (Lucas), 1857 comb.nov. 37. Ontherus (Ontherus) irinus Balthasar, 1938 comb.nov. Brevipennis group 38. Ontherus (Ontherus) raptor sp.nov. 39. Ontherus (Ontherus) planus sp.nov. 40. Ontherus (Ontherus) lichyi Martinez, 1947 comb.nov. Ontherus mirandai Pereira, 1955 41. Ontherus (Ontherus) edentulus sp.nov. 42. Ontherus (Ontherus) brevipennis Harold, 1867 comb.nov. 43. Ontherus (Ontherus) rectangulidens sp.nov. 44. Ontherus (Ontherus) carinicollis Luederwaldt, 1931 stat.nov. 45. Ontherus (Ontherus) zikani Luederwaldt, 1930 comb.nov. 46. Ontherus (Ontherus) lobifi·ons sp.nov. 47. Ontherus (Ontherus) cephalotes Harold, 1869 comb.nov. Ontherus quadrituberculatus Luederwaldt, 1930 Digitatus group 48. Ontherus (Ontherus) aphodioides Burmeister, 1874 comb.nov. Ontherus convexus Luederwaldt, 1930 49. Ontherus (Ontherus) digitatus Harold, 1868 comb.nov. 50. Ontherus (Ontherus) elegans Luederwaldt, 1930 comb.nov. 51. Ontherus (Ontherus) androgynus sp.nov. 52. Ontherus (Ontherus) erosioides Luederwaldt, 1930 comb.nov. 53. Ontherus (Ontherus) amplector sp.nov. 54. Ontherus (Ontherus) atlantidis sp.nov. 55. Ontherus (Ontherus) dentatus Luederwaldt, 1930 comb.nov. Erosus group 56. Ontherus (Ontherus) insolitus sp.nov. 57. Ontherus (Ontherus) stridulator sp.nov. 58. Ontherus (Ontherus) erosus Harold, 1875 comb.nov. 146 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 15 18 17 19 20 FIGS. 14-20. Ontherus kirschii Harold: 14, female anterior tibia, ventral view (arrow indicate ventromedial carina); 15, antenna; 16, flagella; 17, lambda sclerite; 18, 19, accessory sclerites; 20, spermatheca attached to vagina. Line = 0.5 mm. A REVISION OF THE NEOTROPICAL GENUS ONTllERUS ERICHSON 147 �- FIGS. 21-32. Head and prothorax (21, 23, 25, 27, 29, 31, dorsal view; 22, 24, 26, 28, 30, 32, lateral view): 21, 22, Ontherus hadros sp.nov.; 23, 24, 0. lamin(fer Balthasar; 25, 26, 0. alexis (Blanchard); 27,28, 0. tenuistriatus sp.nov.; 29, 30, 0. obliquus sp.nov.; 31, 32, Q_ incisus (Kirsch). Line= 2.0 mm. 148 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 C\ .,Q_ / -�/ --...-;=== FIGS. 33-44. Head and prothorax (33, 35, 37, 39, 41, 43, dorsal view; 34, 36, 38, 40, 42, 44, lateral view): 33, 34, Ontherus howdeni sp.nov.; 35, 36, 0. diabolic us sp.nov.; 37, 38, 0. compressicornis Luederwaldt; 39, 40, 0. pilatus sp.nov.; 41, 42, 0. trituberculatus Balthasar; 43, 44, 0. mexicanus Harold. Line= 2.0 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 149 A 53 J\ 54 �- 55 FIGS. 45-55. Head and prothorax (45, 47, 49, 51, dorsal view; 46, 48, 50, 52, lateral view) and male cephalic horn (53-55, frontal view): 45, 46, Ontherus sextuberculatus sp.nov.; 47, 48, 0. lunicollis sp.nov.; 49, 50, 0. monilistriatus sp.nov.; 51, 55, 0. pseudodidymus sp.nov. Line= 2.0 mm. 150 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 FIGS. 56-61. Male anterior tibiae, dorsal view: 56-58, Ontherus bridgesi Waterhouse (56, Argentina; 57, Bolivia; 58, Peru); 59, 0. brevicollis Kirsch; 60, 61, 0. appendiculatus (Mannerheim) (60, Venezuela; 61, Bolivia). Line= 1.0 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 151 62 64 63 66 �- / 67 ./ h-.. \ / ·-- k_ 68 70 69 FIGS. 62-70. Median lobe of metastemum: 62, Ontherus gilli sp.nov.; 63, 0. ashei sp.nov.; 64, 0. brevicollis Kirsch; 65, 0. politus sp.nov.; 66, 0. aequatorius Bates; 67, 0. bridge.1·i Waterhouse; 68-70, 0. appendiculatus (Mannerheim). Line = 1.0 mm. 152 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 71 74 75 76 77 78 79 80 FIGS. 71-80. Male posterior leg, ventral view: 71, Ontherus kirschii Harold; 72, 0. sanctaemartae sp.nov.; 73, 0. pubens sp.nov.; 74, 0. sulcator (Fabricius); 75, 76, 0. appendiculatus (Mannerheim); 77, 0. azteca Harold; 78, 0. cambeforti sp.nov.; 79, 0. raptor sp.nov.; 80, 0. planus sp.nov. Line= 2.0 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 153 90 FIGS. 81-90. Male posterior leg, ventral view: 81, Ontherus lichyi Martinez; 82, 0. edentulus sp.nov.; 83, 0. brevipennis Harold; 84, 0. rectangulidens sp.nov.; 85, 0. carinicollis Luederwaldt; 86, 0. zikani Luederwaldt; 87, 0. lobifrons sp.nov.; 88, 0. cephalotes Harold; 89, 0. erosioides Luederwaldt; 90, 0. amplector sp.nov. Line= 2.0 mm (81-88), 1.0 mm (89, 90). 154 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 9� �� FIGS. 91-99. Aedeagus (91, 94, 97, lateral view; 92, 95, 98, parameres, ventral view; 93, 96, 99, parameres, dorsal view): 91-93, Ontherus hadros sp.nov.; 94-96, 0. magnus sp.nov.; 97-99, O. laminiferBalthasar. Line= 1.0 mm. A REVISION OF THE NEOTROPICALGENUS ONTHERUS ERICHSON 155 FIGS. 100-114. Aedeagus (100, 103, 106, 109, 112, lateral view; 101, 104, 107, 110, 113, parameres, dorsal view; 102, 105, 108, 111, 114, parameres, ventral view): 100-102, Ontherus gifli sp.nov.; 103-105, 0. ashei sp.nov.; 106-108, 0. hrevicollis Kirsch; 109-111, 0. politus sp.nov.; 112-114, 0. aequatorius Bates. Line= 1.0 mm. 156 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 119 � 122 123 ;:: �.. ,.:,,·'.'>·· 125 ·'.//•·•·· 129 131 132 FIGS. 115-132. Aedeagus (115, 118, 121, 124, 127, 130, lateral view; 116, 119, 122, 125, 128, 131, parameres, dorsal view; 117, 120, 123, 126, 129, 132, parameres, ventralview): 115-117, Ontherus kirschii Harold; 118-120, 0. sactaemartae sp.nov.; 121-123, 0. alexis (Blanchard); 124-126, 0. tenuistriatus sp.nov.; 127-129, 0. obliquus sp.nov.; 130-132, 0. incisus (Kirsch). Line= 1.0 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS� ERICHSON 157 134 135 ·: ..... :: :·:.�:_:.::::;. 11\ 137 138 � 140 141 143 144 142 146 147 145 150 FIGS. 133-150. Aedeagus (133, 136, 139, 142, 145, 148, lateral view; 134, 137, 140, 143, 146, 149, parameres, dorsal view; 135, 138, 141, 144, 147, 150, parameres, ventral view): 133-135, Ontherus howdeni sp.nov.; 136-138, 0. diaholicus sp.nov.; 139-141, 0. compressicor nis Luederwaldt; 142-144, 0. pilatus sp.nov.; 145-147, 0. trituherculatus Balthasar; 148-150, 0. mexicanus Harold. Line = 1.0 mm. 158 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 151 152 153 Ii\ 158 159 � 162 167 168 FIGS. 151-168. Aedeagus (151, 154, 157, 160, 163, 166, lateral view; 152, 155, 158, 161, 164, 167, parameres, dorsal view; 153, 156, 159, 162, 165, 168, parameres, ventral view): 151-153, Ontherus sextuberculatus sp.nov.; 154-156, O. lunicollis sp.nov.; 157-159, 0. monilistriatus sp.nov.; 160-162, O.pseudodidymus sp.nov.; 163-165, 0. bridgesi Waterhouse; 166-168, 0. pubens sp.nov. Line= 1.0 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS� ERICHSON 159 170 171 174 182 183 FIGS. 169-186. Aedeagus (169, 172, 175, 178, 181, 184, lateral view; 170, 173, 176, 179, 182, 185, parameres, dorsal view; 171, 174, 177, 180, 183, 186, parameres, ventral view): 169-171, Ontherus sulcator (Fabricius); 172-174, 0. appendiculatus (Mannerheim); 175-177, 0. ulcopygus sp.nov.; 178-180, 0. podiceps Harold; 181-183, 0. azteca Harold; 184-186, 0. cambeforti sp.nov. Line= 1.2 mm (169-171), 1.0 mm (172-186). 160 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 FIGS. 187- 204. Aedeagus (187, 190, 193, 196, 199, 202, lateral view; 188, 191, 194, 197, 200, 203, parameres, dorsal view; 189, 192, 195, 198, 201, 204, parameres, ventral view): 187-189, Ontherus grandis Luederwaldt; 190-192, 0. carinifrons Luederwaldt; 193-195, 0. virescens (Lucas); 196-198, 0. irinus Balthasar; 199-201, 0. raptor sp.nov.; 202- 204, 0. planus sp.nov. Line= 1.5 mm (187-189), 1.0 mm (190-198), 1.2 mm (199-204). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 161 212 21 ( 3 L. ;.. \;:._::-: ·::·. ,• : ·. . �.� ·.·...... · i.. 218 219 221 222 FIGS. 205-222. Aedeagus (205, 208, 211, 214, 217, 220, lateral view; 206, 209, 212, 215, 218, 221, parameres, dorsal view; 207, 210, 213, 216, 219, 222, parameres, ventral view): 205-207, Ontherus lichyi Martinez; 208-210, 0. edentulus sp.nov.; 211-213, 0. brevipennis Harold; 214-216, 0. rectangulidens= sp.nov.; 217-219, 0. cari- nicollis Luederwaldt; 220-222, 0. zikani Luederwaldt. Line 1.0 mm. 162 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 233 234 239 240 FIGS. 223-240. Aedeagus (223, 226, 229, 232, 235, 238, lateral view; 224, 227, 230, 233, 236, 239, parameres, dorsal view; 225, 228, 231, 234, 237, 240, parameres, ventral view): 223-225, Ontherus lobifrons sp.nov.; 226-228, 0. cephalotes Harold; 229-231, 0. aphodioides Burmeister; 232-234, 0. digitatus Harold; 235-237, 0. elegans Luederwaldt; 238-240, 0. androgynus sp.nov. Line= 1.0 mm (223-228), 0.6 mm (229-231, 238- 240), 0.7 mm (232-237). A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 163 242 245 246 248 249 � � 251 252 255 257 FIGS. 241-258. Aede 260 261 262 264 263 � � � 266 265 267 269 270 271 272 273 " - 274 FIGS. 259-274. Sigmoid sclerites: 259, Ontherus hadros sp.nov.; 260, 0. magnus sp.nov.; 261, 0. laminifer Balthasar; 262, 0. gilli sp.nov.; 263, 0. ashei sp.nov.; 264, 0. brevicollis Kirsch; 265, 0. politus sp.nov.; 266, 0. aequatorius Bates; 267, 0. kirschii Harold; 268, 0. sanctaemartae sp.nov.; 269, 0. alexis (Blanchard); 270, 0. tenuistriatus sp.nov.; 271, 0. obliquus sp.nov.; 2n, 0. incisus (Kirsch); 273+, 0. howdeni sp.nov.; 274, 0. diabolicus sp.nov. Line = 0.5 mm. A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 165 275 276 277 278 279 A • 280 281 282 283 284 285 287 286 288 290 292 293 294 FIGS. 275-294. Sigmoid sclerites: 275, Ontherus compressicornis Luederwaldt; 276, 0. pilatus sp.nov.; 277, 0. trituberculatus Balthasar; 278, 0. mexicanus Harold; 279, 0. sextuberculatus sp.nov.; 280, O. lunicollis sp.nov.; 281, 0. monilistriatus sp.nov.; 282, 0. pseudodidymus sp.nov.; 283, 0. bridgesi Waterhouse; 284, 0. pubens sp.nov.; 285, 0. sulcator (Fabricius); 286, 0. appendiculatus (Mannerheim); 287, 0. ulcopygus sp.nov.; 288, 0. podiceps Harold; 289, 0. azteca Harold; 290, 0. cambeforti sp.nov.; 291, 0. grandis Luederwaldt; 292, 0. carinzfrons Luederwaldt; 293, 0. virescens (Lucas); 294, 0. irinus Balthasar. Line= 0.5 mm. 166 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 . <....>. 296 297 299 � 300 301 303 304 eMllD 305 307 306 - � 311 308 � � 309 310 ' � � � A 314 312 A- 313 FIGS. 295-314. Sigmoid sclerites: 295, Ontherus raptor sp.nov.; 296, 0. planus sp.nov.; 297, 0. lichyi Martinez; 298, 0. edentulus sp.nov.; 299, 0. brevipennis Harold; 300, 0. rectangulidens sp.nov.; 301, 0. carinicollis Luederwaldt; 302, 0. zikaniLuederwaldt; 303, O. lobifrons sp.nov.; 304, 0. cephalotes Harold; 305, 0. aphodioides Burmeister; 306, 0. digitatus Harold; 307, 0. elegans Luederwaldt; 308, 0. androgynus sp.nov.; 309, 0. erosioides Luederwaldt; 310, 0. amplector sp.nov.; 311, 0. dentatus Luederwaldt; 312, 0. isolitus sp.nov.; 313, 0. stridulator sp.nov.; 314, 0. erosus Harold. Line= 0.5 mm. ONTH ERUS A REVISION OF THE NEOTROPICAL GENUS ERICHSON 167 .------c-1 .. a ros mag nus ...------·1lammifer tenuistriatus obliquus incisus ...------1howdeni ------1a1exis 1------1diabolicus .---1compressicornis ilatus ..---1 Crituberculatus ----11unicollis .__ _ 1 monilistriatus -----1mexicanus ....--·isextuberculatus ----·1pseudodidymus gilli ashei politus aequatorius ..___ 1brevicollis o;·------C:: ���g:;!� ..... martae .------r:PU ens sulcator ..---1appendiculatus ...-- ulc�pygus ,...___ pod1ceps ..___..--,azteca cambeforti .____ carinifrons .,..____ g,randis ----·1v1rescens )> .----c:-1raptor planus ..---- lichyi ....--1brevipennis rectangulidens carinicollis zikani lobifrons cephalotes .._____ 1edentulus aphodioides elegans anarogynus erosio1Cles amplector antlantidis dentatus insolitus stridulator erosus ....- --1��gitatus .....------�11r�1n�ur-s�,.=:=����� n ges1 .b rectus · Z .______u group ��0 �� ���\\' FIG. 315. Reconstructed phylogeny of Ontherus. 168 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA NO. 170 INDEX aequatorius ...... 10,18, 28, 34, 35 alexis ...... 3, 4, 8, 16,23, 39, 40, 41, 51,139 Alexis Group ...... 39, 51, 139 amplector...... 3,15, 22, 127, 128, 129 androgynus ...... 3,14, 22, 124, 125 aphodioides ...... 14, 21, 117, 118, 119, 141 appendiculatus ...... 11, 19, 78, 79, 80, 97, 137, 140 Appendiculatus Group ...... 78, 140 ashei ...... 3,10, 18,30, 139 Ateuchus ...... 141 atlantidis ...... 3,14, 22, 129, 130 azteca...... 3, 12, 20,87, 88,89, 140 brevicollis ...... 10, 19, 24, 28, 31, 32, 136,139 Brevicollis Group ...... 24, 28, 139 brevipennis ...... 5,12, 20, 23, 97, 104, 106,137, 140 Brevipennis Group ...... 5, 23, 97, 137,140 bridgesi ...... 11, 19, 67, 68, 69,139 Caelontherus...... 3,4, 7, 15, 23 cambeforti ...... 3, 12, 20, 89, 90, 140 Can thon ...... 141 carinicollis ...... 3, 13,21, 97, 108, 110 carinifrons ...... 12,20, 78, 92, 94, 140 Catharsius ...... 4, 139 cephalotes ...... 3, 13, 21,112, 115, 116 Chalcocopris ...... 4 compressicornis ...... 8, 9,15, 17,51, 52 contractus ...... 78, 79, 80,82 convexus ...... 118, 119 Coprini ...... 4,143 Copris ...... 4,23,39,40,41,46,74,75,76,77,78,80, 136, 139, 141, 142 Coptodactyla ...... 4,136, 139 Deltochilum...... 141 dentatus ...... 15, 22, 130, 131 diabolicus ...... 3, 9,17, 39, 48, 49, 140 Dichotomiini ...... 4 Dichotomius ...... 4, 78, 136,141 didymus ...... 3, 4, 39, 40, 41, 63 digitatus ...... 5, 11, 19, 23, 117, 120, 121, 141 Digitatus Group ...... 5, 117 edentulus ...... 3,12, 20, 102, 104, 140 elegans...... 14,22, 121, 122,123 elongatus ...... 3,35, 37, 70 erosioides...... 15, 22, 125, 127 erosus ...... 14, 21,128, 131, 134, 135,141 Erosus Group ...... 131 gilli ...... 3,10, 18, 28, 29,30 glaucinus ...... 3, 4, 39,40, 41 grandis...... 12, 19, 91, 92, 140 hadros ...... 3,10, 18, 23,24, 25, 136, 139 Hadros Group ...... 24, 139 Heliocopris ...... 4 Holocephalus...... 4 howdeni ...... 3, 8, 16,47, 48 incisus ...... 3, 8,16, 32, 45, 46, 47 insolitus ...... 3,14, 21, 131, 132 irinus ...... 14,22, 71, 78, 96, 97,137 Isocopris ...... 4 kirschii ...... 3,10, 18,24,35,36,37, 136, 139 Kirschii Group ...... 24,35, 139 laminifer ...... 10, 18,23, 26, 27, 28, 136, 140 Laminifer Group ...... 26 lichyi ...... 12, 20, 100, 101,102, 140 A REVISION OF THE NEOTROPICAL GENUS ONTHERUS ERICHSON 169 Litocopris ...... 4 lobifrons ...... 3, 13,21, 112, 113, 114, 117 lunicollis ...... 3,9, 17,47,59,61,62 magnus ...... 3, 10, 18, 25, 26,136 Metacatharsius...... 4 mexicanus ...... 5, 9, 17, 23, 24, 50, 55, 58, 59, 139, 140 Mexicanus Group ...... 24,50, 139 mirandai ...... 100, 101, 102 monilistriatus...... 3,9, 17, 62, 63 nevinsoni ...... 3,39,41,52 nisus...... 75, 76,78 obliquus...... 3, 8, 16, 43,45 Ontherus ...... 3,4,5,7, 10, 15, 19,22,23,24,25,26,27,28,29,30,31,32,33, 34,35,36,37, 38,39,40,41,43,45,46,47,48,49,51,52,53,54, 55,58,59,61,62,63, 65,67,68,69,70, 71,72, 74, 75, 78, 79, 80, 82, 83,85,86, 87,88,89,90, 91,92,94,95,96,97,99, 100, 101, 102, 104, 106, 107, 108, 110, 112, 114, 115, 116, 117, 118, 119, 120, 121, 122, 123, 124, 125, 127, 128, 129, 130, 131, 132, 133, 134, 135, 136, 137, 139, 140, 141, 142 Ontherus s. str...... 3,5, 10, 70, 136, 137, 139, 140 Onthophagini...... 136 ovalipennis ...... 75, 77 Planontherus...... 3,4, 5, 10, 19, 67 planus ...... 3, 12, 20,99, 100 podiceps...... 12,20, 85, 86 politus ...... 3, 10, 18, 33, 34 polynice...... 78, 79, 80, 82 Proagoderus...... 136 pseudodidymus ...... 3,9, 17, 62, 63, 65, 140 pubens ...... 3, 14, 22, 71, 72, 74, 140 quadratus ...... 78, 79, 80, 82 quadrituberculatus...... 3, 115, 116, 117 raptor ...... 3, 12, 20, 97, 99 rectangulidens ...... 3, 13,20, 106, 107, 112 rectus...... 3, 11, 19, 69,70, 139 rotundatus ...... 78,79, 80, 82 sanctaemartae ...... 3, 10, 18, 37, 38, 39, 139 Scarabaeinae ...... 136, 139, 142 Scarabaeini ...... 4 senegalensis...... 75, 77 sextuberculatus ...... 3, 9, 18, 58, 59, 140 stridulator ...... 3, 14, 21, 133, 134 strius ...... 3, 87, 89 sulcator ...... 4, 14, 22, 70, 71, 74, 75, 76, 140 Sulcator Group ...... 71, 140 Synapsis ...... 4 tenuistriatus ...... 3, 8, 16, 41, 43 thoracicus ...... 3, 45, 46, 47 Thyregis...... 4, 139 trituberculatus ...... 9, 17, 53, 54 ulcopygus ...... 3, 12, 20, 82, 83 villosus ...... 3, 87, 88, 89 virescens ...... 5, 12, 20, 23, 78, 94, 95, 96, 97 Xinidium ...... 4, 139 zikani ...... 3,13, 21, 107, 108, 110, 112