Anderson 1987 Qev23n4 431 709 CC Released.Pdf
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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. SYSTEMATICS, PHYLOGENY AND BIOGEOGRAPHY OF NEW WORLD WEEVILS TRADITIONALLY OF THE TRIBE CLEONINI (COLEOPTERA: CURCULIONIDAE; CLEONINAE) Robert Samuel Anderson Department of Entomology University of Alberta Edmonton, Alberta, T6G 2E3 Quaestiones Entomologicae CANADA 23:431-709 1987 ABSTRACT A survey of representatives of generic-group names in Cleoninae and consideration of their phylogenetic relationships indicates four genera are represented in the New World. These are: Cleonis Dejean (introduced; 1 species); Stephanocleonus Motschulsky (Holarctic, northern; 6 species including S. confusus n. sp. [type locality, Black Hills, South Dakota], S. immaculatus n. sp. [type locality, Fort McMurray, Alberta], S. parshus n. sp. [type locality, Chalk River, Ontario], and S. stenothorax n. sp. [type locality, Bluefish Caves, Yukon Territory]); Apleurus Chevrolat (southwestern North America, arid lands; 8 species); and, Cleonidius Casey (eastern and western North America, semi-arid and arid lands; 19 species including C. eustictorrhinus n.sp. [type locality, Sacramento, California], C. infrequens n. sp. [type locality, Scott City, Kansas], and, C. notolomus n. sp. [type locality, Grant County, New Mexico]). Cleonidius is hypothesized to be phylogenetically more closely related to traditional Lixini rather than Cleonini and present tribal classification of Cleoninae is regarded as unsatisfactory. The following new generic-group synonymies are presented: Lixus Fabricius (=Epimeces Billberg [type species Curculio filiformis Fabricius, here designated]); Stephanocleonus Motschulsky (^Coniocleonus Motschulsky); Apleurus Chevrolat (^Centrocleonus LeConte, Cleonopsis LeConte, Cleonaspis LeConte, and Dinocleus Casey); and, Cleonidius Casey (^Lixestus Reitter). The following information is given in detail for each of Cleonis, Stephanocleonus, Apleurus (Gibbostethus) n. subg. (type species, Dinocleus hystrix Fall by monotypy), Apleurus (Apleurus), and Cleonidius: synonymic list, notes about synonymy, diagnosis, description, list of included species, and a discussion of phylogenetic relationships. Keys to species of North American Stephanocleonus and Apleurus, and world Cleonidius, are presented. The following information is given in detail for each species considered: synonymic list, notes about synonymy, problems in recognition, description, distribution, natural history information, and chorological relationships. Lixus mixtus LeConte is a junior homonym and is replaced with Cleonidius longinasus, new name. The following new species-group synonymies are presented: Apleurus lutulentus (LeConte) (^Cleonus pulvereus LeConte, Apleurus fossus Chevrolat, and Dinocleus bryanti Van Dyke); Apleurus porosus (LeConte)(=Dinocleus farctus Casey); Apleurus angularis (LeConte) ( = Dinocleus denticollis Casey and Dinocleus porcatus Casey); Apleurus jacobinus (Casey) (^Centrocleonus pilosus LeConte and Cleonus (Dinocleus) capillosus Csiki); Apleurus Present address: Department of Entomology, Texas A & M University, College Station, TX 77843, U. S. A. 432 Anderson albovestitus (Casey) (= Cleonus molitor LeConte, Dinocleus densus Casey, Dinocleus wickhami Casey, Dinocleus interruptus Casey, Dinocleus mexicanus Casey, and Cleonus (Dinocleus) structor Csiki); Apleurus saginatus (Casey) (^Dinocleus dentatus Champion); Cleonidius subcylindricus Casey (=Cleonus (Cleonidius) graniferus Casey); Cleonidius texanus (LeConte) (= Cleonus (Cleonidius) circumductus Casey); Cleonidius frontalis (LeConte) (=Cleonus sparsus LeConte and Cleonus (Cleonidius) stratus Csiki); Cleonidius poricollis (Mannerheim) (^Cleonus (Cleonidius) lobigerinus Casey and Cleonus (Cleonidius) kirbyi Casey); Cleonidius boucardi (Chevrolat) (=Cleonus carinicollis LeConte and Cleonus (Cleonidius) lecontellus Csiki); and, Cleonidius trivittatus (Say) (=Cleonus inornatus LeConte and Cleonus (Cleonidius) bicarinatus Casey). Lixus californicus Motschulsky is considered a nomen dubium. Only for species of Apleurus and Cleonidius are reconstructed phytogenies presented based on analyses of 31 characters in Apleurus and 22 characters in Cleonidius. Apleurus is hypothesized to be the monophyletic sister genus of the Palearctic Chromoderus. In Apleurus, two monophyletic subgenera are recognized. These are Gibbostethus (I species, A. hystrix,) and Apleurus s. s. (7 species). Within Apleurus s. s. included species and hypothesized phylogenetic relationships are as follows: ((((A. albovestitus + A. saginatusj + ((A. jacobinus + A. angularisj + A. porosusjj + A. lutulentusj + A. aztecusj. Cleonidius is monophyletic and hypothesized to comprise an unresolved trichotomy with the Palearctic Lixus (Lixoglyptus) and Cylindropterus. Four monophyletic species groups are recognized in Cleonidius. Along with constituent species and hypothesized phylogenetic relationships, these are: C. erysimi group, (((C texanus + C. longinasusj + C. subcylindricusj + C. pleuralis + (C erysimi + C. eustictorrhinus)); C. americanus group, ((((C. puberulus + C. infrequens) + (C. collaris + C. notolomusJJ + C. canescensj +(C. americanus + C. frontalisJJ; C. poricollis group, (C poricollis + C. calandroidesj; and C. boucardi group, (((C. placidus + C. quadrilineatusj + C. trivittatusj + C. boucardij. Phylogenetic relationships of groups are (((C. boucardi group + C. poricollis group) + C. americanus group) + C. erysimi group). Phylogenetic relationships of the Nearctic species component of Cleonidius are unresolved but the latter is likely sister to the Palearctic species component. Both Apleurus and Cleonidius are hypothesized to have entered North America in savanna or grassland habitats (or their precursors) across Beringia during late Eocene time. In North America, primary diversification has taken place in the arid lands of the southwestern United States although two species of Cleonidius are found in coastal regions of eastern North America. In both genera there appears to be an initial vicariance of lineages, on Compositae, in grassland or mesquite-grassland habitats between California, central Mexico, and the southern Rocky Mountains. Further speciation events in Apleurus appear due primarily to shifts to new, apotypic habitat types at differing altitudes brought about by increased aridity during the later Tertiary; host plant shifts are not a factor because Apleurus species are all associated with Compositae. Host plant shifts, however, primarily between Leguminosae-Rosaceae and likely also Compositae-Cruciferae, and within-habitat-type allopatric speciation appear to predominate as the causes of further speciation in Cleonidius, most species of which remain associated with plesiotypic less-arid habitat types. Habitat shifts promoting speciation in Cleonidius are evident, but are not as associated with formation of increasingly arid habitats as in Apleurus. Evolutionary trends in Apleurus and Cleonidius are discussed and patterns and predictions of biogeographic or evolutionary interest summarized and presented for examination for New World Cleonini 433 general applicability. RESUME Vne diagnose des characlres de la sous-famille est aussi brievement discutee. L'examen de representants des groupes-genres de Cleoninae et leurs relations phyletiques indiquent que quatre genres sont repr'esentes dans le Nouveau-Monde. lis sont: Cleonis Dejean (introduit; I espece); Stephanocleonus Motschulsky (Holoartique, nordique; 6 especes incluant S. confusus n. esp. [localite-type. Black Hills, Dakota du Sud], S. immaculatus n. esp. [localite-type, Fort McMurray, Alberta], S. parshus n. esp. (localite-type. Chalk River, Ontario}, et S. stenothorax n. esp. [localite-type, Bluefish Caves, Territoire du Yukon}; Apleurus Chevrolat (sud-ouest de I'Amerique du Nord, terres arides; 8 especes); et Cleonidius Casey (est et ouest de I'Amerique du Nord, terres arides et semi-arides; 19 especes incluant C. eusticlorrhinus n. esp [localite-type, Sacramento, Califournie], C. infrequens n. esp. [localite-type, Scott City, Kansas/, et C. notolomus n. esp. [localite-type. Grant County, Nouveau MexiqueJ. Cleonidius est suggere comme etant phyletiquement plus apparente aux traditionels Lixini pluibt qu'aux Cleonini, et la classification actuelle de Cleoninae est consideree comme etant insatisfaisante. De nouvelles synonymies de groupe-genres sont presentee: Lixus Fabricius (= Epimeccs Billberg [espece-type Curculio filiformis Fabricius, ici designee]); Stephanocleonus Motschulsky (^Coniocleonus Motschulsky); Apleurus Chevrolat ^Centrocleonus LeConte, Cleonopsis LeConte, Cleonaspis LeConte, et Dinocleus Casey); et Cleonidius Casey f^Uxestus Reitter). L'information suivante est presentee en detail pour chacun des Cleonis, Stephanocleonus, Apleurus (Gibbostethus) nouveau sous-genre {espece-type, Dinocleus hystrix Fall par monotypie), Apleurus (Apleurus), et Cleonidius: liste synonymique et notes les concernant, diagnose, description, liste des especes incluses, et une discussion concernani les relations phyletiques. Des clefs d'identification a Vespece pour les Stephanocleonus et Apleurus de I'Amerique du Nord, et les Cleonidius Mondiaux sont produites. L'information suivante est aussi presentee en detail pour chacune des especes considerees: liste synonymique