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This work is licensed under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/3.0/us/ or send a letter to Creative Commons, 171 Second Street, Suite 300, San Francisco, California, 94105, USA. CLASSIFICATION, PHYLOGENY, AND ZOOGEOGRAPHY OF SCHIZOGENIUS PUTZEYS (COLEOPTERA: CARABIDAE: SCARITINI)* DONALD R. WHITEHEAD Department of Entomology University of Alberta Quaestiones entomologicae Edmonton 7, Alberta 8 : 131-348 1972 North and Middle American species of Schizogenius Putzeys are reviewed in detail, and South American species are treated provisionally. The genus is redefined, characterized, and illustrated. Two subgenera are Genioschizus new subgenus, with three species groups, and Schizogenius s. str., with 21 species groups. In all, I recognize 68 species and two subspecies. Keys are given to: subgenera; described species groups, species, and subspecies of Genio schizus; same of North and Middle American Schizogenius s. str.; and species groups and most described species of South American Schizogenius s. str. / describe as new the following 27 taxa: S. crenulatus chiapatecus, S. sculptilis, S. im- puncticollis, S. suturalis, S. jacarensis, S. bicolor, S. cearaensis, S. negrei, S. costipennis, S. reichardti, S. ocellatus, S. lindrothi, S. brevisetosus, S. seticollis vandykei, S. plurisetosus, S. kulti, S. dilatus, S. tibialis, S. ozarkensis, S. planuloides, S. neovalidus, S. chiricahuanus, S. pacificus, S. emdeni, S. scopaeus, S. falli, and S. ochthocephalus. Synonymies proposed for the first time are: S. peninsularis Van Dyke (= S. auripennis Bates), S. angusticollis Putzeys (= S. arechavaletae Putzeys), S. validus Fall (= S. longipennis Putzeys), and S. championi Kult (= S. pygmaeus Van Dyke). Given for each species are, as appropriate: synonymic list, diagnostic combination, de scription, discussion of variation, etymological derivation, geographic distribution list, col lecting notes, taxonomic notes, and illustrations of important structural characteristics. Geographic distributions are mapped for all North and Middle American species. Descrip tions of most North and Middle American species are augmented with tables of descriptive statistics. Results of detailed statistical analyses of geographic variation for members of some North and Middle American species groups are tabulated, mapped, and discussed. A phytogeny is reconstructed for the genus, and carefully integrated with historical zoo geography. No geographic evidence is used to reconstruct the phytogeny of major groups and lineages; phytogenies derived from contrasting phyletic and phenetic techniques are favorably compared. To derive the phytogeny of members of the truquii lineage, which includes most North and Middle American species of Schizogenius, use of zoogeographic evidence is essential; simple cladistic techniques are not used because character states of too many characteristics are reversible. An average time between dichotomies in the recon structed phytogeny of Schizogenius is about 3,000,000 years. This interval is used to inte grate Schizogenius phytogeny and zoogeography. On the same basis, phytogenies and zoo geographies of Brachinus and Evarthrus are compatible. Zoogeographies o/Brachinus, Evar- thrus, and Schizogenius are compared for North and Middle American faunas. Ancestral Schizogenius evolved in South America about Middle Eocene. Ancestors of the ferrugineus group, the truquii lineage, and the crenulatus group entered North America in Late Eocene, Middle Oligocene, and Middle Miocene. The ancestor of the truquii lineage ultimately evolved into eight species groups, and their evolutionary zoogeographies are dis cussed in detail. Since Early Pleistocene, members of the tenuis, optimus, and lindrothi groups have crossed the Panamanian land bridge to Middle America, and members of the depressus group have spread southward into South America. * Revised version of thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of require ments for the degree of Doctor of Philosophy at the University of Alberta. The author's present address is c/o Department of Entomology, United States National Museum, Washington, D. C. 20560. 132 Whitehead En este trabajo se revisan, en detalle, las especies de Schizogenius Putzeys de Norte y Centro America, en tanto que las especies de Sur America son revisadas provisionalmente. El genero es redefinido, senalado, e ilustrado. Dos subgeneros son el Genioschizus, subgenero nuevo, con tres grupos de especies; y el Schizogenius s. str., con 21 grupos. En total, yo reconozco 68 especies y dos subespecies. Se han ofrecido claves para los subgeneros, grupos de especies, especies, y subespecies del Genioschizus; lo mismo que para el Schizogenius s. str. de Norte y Centro America, asi como para los grupos de especies, y la mayor parte de las especies descritas en Sur America. Yo describo como nuevos los siguientes: S. crenulatus chiapatecus, S. sculptilis, S. impunc- ticollis, S. suturalis, S. jacarensis, S. bicolor, S. cearaensis, S. negrei, S. costipennis, S. reich- ardti, S. ocellatus, S. lindrothi, S. brevisetosus, S. seticollis vandykei, S. plurisetosus, S. kulti, S. dilatus, S. tibialis, S. ozarkensis, S. planuloides, S. neovalidus, S. chiricahuanus, S. pacifi- cus, S. emdeni, S. scopaeus, S. falli, y S. ochthocephalus. Son sinonimias nuevas: S. penin- sularis Van Dyke (= S. auripennis Bates), S. angusticollisPutzeys (= S. arechavaletae Putzeys), S. validus Fall (= S. longipennis Putzeys), y S. championi Kult (= S. pygmaeus Van Dyke). Para cada especie se ha ofrecido, como conviene: lista de sinonimias, combination de diagnostico, descripcion, discurso de variation, derivation etimologica, lista de localidades, notas sobre coleccion y taxonomia, e ilustraciones de caracteristicas esenciales. Igualmente se dan cartas de distribution para las especies de Norte y Centro America. Las especies de Norte y Centro America han sido provistas, en su description, con tablas de estadisticas descriptivas; y los resultados de andlisis estadisticos detallados, son catalogados, delineados en cartas, y discutidos. Se ha reconstruido una filogenia para el genero, y, cuidadosamente, se ha integrado esta con una zoogeografla historica. No se ha usado ninguna evidencia geogrdfica para la recons truction de la filogenia de grupos y los linajes mayores; las filogenias derivadas de tecnicas fileticas y feneticas son comparadas favorablemente. Para derivar la filogenia de miembros del linaje truquii, que contiene la mayor parte de las especies del Schizogenius de Norte y Centro America, el uso de evidencia zoogeogrdfica es indispensable. No se han usado tecni cas feneticas sencillas debido a que los estados de cardcter con demasiadas caracteristicas, son reversibles. Un intervalo proporcional entre dicotomias en la filogenia reconstruida del Schizogenius se aproxima a los 3,000,000 de anos. Este intervalo es usado para integrar la filogenia con la zoogeografla del Schizogenius. Igualmente, las filogenias y zoogeografias del Brachinus y el Evarthrus son compatibles. Las zoogeografias del Brachinus, el Evarthrus, y el Schizogenius son comparadas para las faunas de Norte y Centro America. El Schizogenius ancestral se desarrollo en Sur America alrededor del Eoceno Medio. Los predecesores del grupo ferrugineus, del linaje truquii, y del grupo crenulatus, penetraron en Norte America durante el Eoceno Ultimo, Oligoceno Medio, y Mioceno Medio. El predece- sor del linaje truquii finalmente desarrollado en ocho grupos de especies, y sus zoogeografias evolutivas son discutidos en detalle. Desde el Pleistoceno Primario, miembros de los grupos tenuis, optimus, y lindrothi han atravesado el puente de tierra de Panama hacia Centro America, y miembros del grupo depressus se han distribuido en el interior de Sur America. CONTENTS Materials 133 Methods 135 Taxonomy 142 Phylogeny 302 Zoogeography 326 References 345 Classification, phylogeny and zoogeography of Schizogenius 133 Introduction. — Small American beetles of the tribe Scaritini have received little careful attention. This is unfortunate since as a result no other aspects of their biologies have been studied, but fortunate in that comparatively few taxonomic errors have been introduced into the literature. Lindroth (1961) wrote a fine revision of the Canadian and Alaskan members of this fascinating group of beetles. His treatment of Schizogenius, the subject of my investiga tions, is excellent for the limited Canadian fauna but is not adequate to identify specimens from most regions in the United States. The genus, after all, is mainly Neotropical, and in fact a good taxonomy for it must be based on the entire fauna. Except for Lindroth's (1961) key to described species from America north of Mexico, the most recent full revisions of Schizo genius were done by Putzeys (1866) for Mexico and southward, and by LeConte (1879) for America north of Mexico. How much is known about these beetles? Since Thomas Say named the first species in 1823, little more than descriptive morphological work has been done. Immature stages remain completely unknown, and aside from collector's notes on habitat we have no in formation on adult biology. The earliest names for species now placed in Schizogenius were proposed by Say (1823), Castelnau (1835), and Haldeman (1843). Putzeys (1846) intro duced the generic name Schizogenius, and in this and subsequent