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A Cladistic Analysis and Classification of the Subfamily Bembicinae (Hymenoptera: Crabronidae), with a Key to the Genera

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A Cladistic Analysis and Classification of the Subfamily Bembicinae (Hymenoptera: Crabronidae), with a Key to the Genera Zootaxa 3652 (2): 201–231 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3652.2.1 http://zoobank.org/urn:lsid:zoobank.org:pub:2A51BCDC-77D6-4D3F-8F4D-830EEC5B3EEB A cladistic analysis and classification of the subfamily Bembicinae (Hymenoptera: Crabronidae), with a key to the genera PAVEL G. NEMKOV & ARKADY S. LELEJ1 Institute of Biology and Soil Science, Far Eastern Branch of Russian Academy of Sciences, 100 let Vladivostoku 159, Vladivostok, 690022, Russia. E-mail: [email protected]; [email protected] 1Corresponding author: E-mail: [email protected] Abstract A cladistic analysis of the digger wasp subfamily Bembicinae based on morphological characters is presented. The under- lying data matrix comprises 83 terminal taxa (coded on genus-level) and 64 morphological characters. The resulting strict consensus tree was used as the basis for a revised tribal and subtribal classification of the Bembicinae. Based on a previ- ously published classification, we herewith propose a change: the tribe Heliocausini Handlirsch 1925, stat. resurr. (com- posed of Acanthocausus Fritz & Toro 1977, Heliocausus Kohl 1892, and Tiguipa Fritz & Toro 1976) is separated from Bembicini Latreille 1802. Four tribes are recognized within the subfamily Bembicinae and seven subtribes within the tribe Gorytini and two subtribes in the tribe Nyssonini, based on the present cladistic analysis.The subtribe Nurseina Nemkov & Lelej, subtrib. nov. (comprising of Nippononysson Yasumatsu & Maidl 1936 and Nursea Cameron 1902) is separated from other genera in the tribe Nyssonini Latreille 1804. An new identification key to the genera of the Bembicinae is pro- vided. Key words: Heliocausini, Nurseina, digger wasps, sand wasps, phylogeny Introduction The subfamily Bembicinae comprise 1697 recognized species, which are divided into 81 genera and three tribes (Pulawski 2013). Besides those there are one fossil genus Biamogorytes Nemkov 1990 (Upper Oligocene) and three fossil species: Biamogorytes handlirschi Nemkov 1990 (Upper Oligocene), Gorytes archoryctes (Cockerell 1922) (Eocene) and Psammaecius sepultus (Cockerell 1906) (Lower Oligocene). Bembicinae is the second (after Crabroninae) most numerous subfamily within Crabronidae with regard to the number of species (19 % of total) and genera (31 % of total). They are distributed worlwide: 21 genera and 434 species occur in the Palearctic region, 36 and 275, respectively, in the Nearctic region, 19 and 292 in the Afrotropical region, 19 and 161 in the Oriental region, 53 and 353 in the Neotropic region, and 10 and 176 in the Australian region. Six species of Nysson, Stizus, and Bembix are of doubtful origin, without known type locality. The majority of Bembicinae are ground-burrowing wasps. Their burrows contain from one to several cells per nest (Evans 1966, Evans & O'Neill 2007, Nemkov 2012). The prey of Bembicinae are various insect, mainly Orthoptera, Hemiptera-Auchenorrhyncha, and Diptera. Some genera of Bembicinae (Stizoides and all Nyssonini genera with known natural history) are cleptoparasitic on other digger wasps (Sphecidae and Crabronidae). This paper is the final one in the series on phylogeny and classification of the subfamily Bembicinae. The previous papers (Nemkov & Lelej 1996, Nemkov & Pulawski 2009, Nemkov & Ohl 2011) have been devoted to the most complicated and diverse tribe, namely Bembicini. In those papers the detailed historical review has been presented. In the current paper, we present a phylogenetic analysis of the genus-level relationships in the whole subfamily based on morphological characters of adults with strict cladistic methods. Larval characters are not included because data are available only for a small part of the ingroup taxa. Based on the resulting hypothesis, a new classification of the Bembicinae is suggested. Accepted by C. Rasmussen: 9 Apr. 2013; published: 16 May 2013 201 Taxonomic history Bohart and Menke (1976) divided the subfamily Bembicinae (their Nyssoninae) into six tribes: Heliocausini, Alyssontini (their Alyssonini), Nyssonini, Gorytini, Stizini, and Bembecini. Later the tribe Gorytini was in turn divided into six subtribes (Argogorytina, Clitemnestrina, Exeirina, Gorytina, Handlirschiina, and Olgiina) (Nemkov & Lelej 1996). Menke (1997) and Bohart (2000) followed this proposal. However, Hanson and Menke (2006) (based on Prentice’s [1998] unpublished doctoral thesis) recognized only three tribes within Bembicinae: Alyssontini, Bembicini, and Nyssonini, and they downgraded Heliocausini, Gorytini, and Stizini to subtribes of Bembicini (Clitemnestrina and Olgiina were synonymized with Exeirina, and Argogorytina was synonymized with Gorytina). Nemkov and Pulawski (2009) accepted Prentice’s proposed classification (Hanson & Menke 2006), but resurrected the subtribe Argogorytina and established the new monotypic subtribe Trichogorytina within the tribe Bembicini. Nemkov and Ohl (2011) synonymized Argogorytina with Exeirina and Trichogorytina with Gorytina, resurrected the subtribe Stictiellina, and removed the new subtribe Spheciina from Handlirschiina. Thus, the tribe Bembicini is currently divided into eight subtribes: Bembicina, Exeirina, Handlirschiina, Heliocausina, Gorytina, Spheciina, Stictiellina, and Stizina. Material and methods Descriptive terminology follows Bohart and Menke (1976), but the terms "mesosoma" instead of "thorax" and "metasoma" instead of "gaster" are used. Material studied. This study was based on specimens from the following institutions: Bohart Museum of Entomology, University of California, Davis, California, USA (Steven L. Heydon): various taxa. California Academy of Sciences, San Francisco, California, USA (Wojciech J. Pulawski): various taxa. Institute of Biology and Soil Science, Far Eastern Branch of Russian Academy of Sciences, Vladivostok, Russia (Pavel G. Nemkov): various taxa. Naturhistorisches Museum, Wien, Austria (Dominique Zimmermann): holotype of Aroliagorytes imitator, syntypes of Pseudoplisus rubiginosus. The Natural History Museum, London, United Kingdom (Christine Taylor): Leiogorytes guerrero including the holotype, Pseudoplisus nyasicus including the holotype, Pterygorytes triangularis including the holotype. United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (Brian Harris): Chilostictia aberrans, paratype of Paraphilanthus costaricae, Pseudoplisus fasciatus, Pseudoplisus venustus. Utah State University, Logan, Utah, USA (Terry L. Griswold): Allogorytes bifasciatus. Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (Sergey A. Belokobylskij): various taxa. Zoological Museum, Moscow State University, Moscow, Russia (Alexandr V. Antropov): various taxa. Species examined. Altogether, the following 421 species and subspecies belonging to 83 genera have been studied: Outgroup (subfamily Mellininae, two genera): Mellinus Fabricius 1790: M. arvensis (Linnaeus 1758), M. crabroneus (Thunberg 1791), M. obscurus Handlirsch 1888. Xenosphex F. Williams 1954: X. xerophilus F. Williams 1954. Ingroup (subfamily Bembicinae, 81 genera, in alphabetical order): Acanthocausus Fritz & Toro 1977: A. tridens (Brèthes 1913). Acanthostethus Smith 1869: A. mysticus (Gerstaecker 1867). Antomartinezius Fritz 1955: A. fritzi Bohart 1968. Afrogorytes Menke 1967: A. monstrosus (Handlirsch 1894). Allogorytes Bohart 2000: A. bifasciatus (Brèthes 1909). Alysson Panzer 1806: A. cameroni Yasumatsu & Masuda 1932, A. monticola Tsuneki 1977, A. pertheesi Gorski 202 · Zootaxa 3652 (2) © 2013 Magnolia Press NEMKOV & LELEJ 1852, A. ratzeburgi (Dahlbom 1843), A. spinosus (Panzer 1801), A. tricolor Lepeletier de Saint Fargeau & Audinet-Serville 1825. Ammatomus A. Costa 1859: A. asiaticus (Radoszkowski 1886), A. coarctatus (Spinola 1808), A. mesostenus (Handlirsch 1888), A. rogenhoferi (Handlirsch 1888), A. rufonodis (Radoszkowski 1877). Analysson Krombein 1985: A. rufescens Krombein 1985. Argogorytes Ashmead 1899: A. areatus (Taschenberg 1875), A. carbonarius (F. Smith 1856), A. fairmairei (Handlirsch 1893), A. fargeii (Shuckard 1837), A. hispanicus (Mercet 1906), A. konishii (Nemkov 1999), A. mystaceus mystaceus (Linnaeus 1761), A. mystaceus grandis (Gussakovskij 1932), A. nigrifrons (F. Smith 1856), A. nipponis Tsuneki 1963, A. umbratilis Bohart 2000. Arigorytes Rohwer 1912: A. coachellae Bohart 1971, A. insolitus (W. Fox 1895), A. ruficrus Bohart 1971, A. smohalla Pate 1947. Aroliagorytes Bohart 2000: Aroliagorytes imitator (Handlirsch 1901). Austrogortyes Bohart 1967: A. bellicosus (F. Smith 1862). Bembecinus A. Costa 1859: B. acanthomerus (Morice 1911), B. asiaticus Gussakovskij 1935, B. gracilicornis (Handlirsch 1892), B. hungaricus (Frivaldszky 1876), B. meridionalis A. Costa 1859, B. tridens (Fabricius 1781), B. validior Gussakovskij 1952. Bembix Fabricius 1775: B. bicolor Radoszkowski 1877, B. bidentata Vander Linden 1829, B. dilatata Radoszkowski 1877, B. diversipes F. Morawitz 1889, B. gracilis Handlirsch 1893, B. integra Panzer 1805, B. kirgisica F. Morawitz 1891, B. megerlei Dahlbom 1845, B. niponica niponica F. Smith 1873, B. niponica picticollis F. Morawitz 1889, B. oculata Panzer 1801, B. olivacea Fabricius 1787, B. pallida Radoszkowski 1877, B. planifrons F. Morawitz 1891, B. rostrata (Linnaeus 1758), B. transcaspica Radoszkowski 1893, B. turca Dahlbom 1845. Bicyrtes Lepeletier de Saint Fargeau 1845: B. angulatus (F. Smith 1856), B. capnopterus (Handlirsch
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