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Coleoptera, Tetratomidae) of America North of Mexico 1 Doi: 10.3897/Zookeys.188.2976 Monograph Launched to Accelerate Biodiversity Research

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Coleoptera, Tetratomidae) of America North of Mexico 1 Doi: 10.3897/Zookeys.188.2976 Monograph Launched to Accelerate Biodiversity Research A peer-reviewed open-access journal ZooKeys 188:Review 1–153 (2012) of the Eustrophinae (Coleoptera, Tetratomidae) of America north of Mexico 1 doi: 10.3897/zookeys.188.2976 MONOGRAPH www.zookeys.org Launched to accelerate biodiversity research Review of the Eustrophinae (Coleoptera, Tetratomidae) of America north of Mexico Darren A. Pollock1,† 1 Department of Biology, Eastern New Mexico University, Portales NM USA † urn:lsid:zoobank.org:author:A9B3DFD3-6391-4E1D-8894-53B148A3322B Corresponding author: Darren A. Pollock ([email protected]) Academic editor: P. Bouchard | Received 20 February 2012 | Accepted 11 April 2012 | Published 2 May 2012 urn:lsid:zoobank.org:pub:23EF56E0-CF08-4D63-AF65-C31D8648BDF4 Citation: Pollock DA (2012) Review of the Eustrophinae (Coleoptera, Tetratomidae) of America north of Mexico. ZooKeys 188: 1–153. doi: 10.3897/zookeys.188.2976 Abstract The Nearctic fauna (north of Mexico) of Eustrophinae is reviewed, and consists of the following five genera and 12 species: Pseudoholostrophus (Pseudoholostrophus) impressicollis (LeConte), P. (Holostrophinus) discolor (Horn), Holostrophus bifasciatus (Say), Eustrophus tomentosus Say, Eustrophopsis confinis (LeConte), E. bicolor (Fabricius), E. brunneimarginatus (Dury), E. indistinctus (LeConte), E. arizonensis (Horn), E. ornatus (Van Dyke), E. crowdyi sp. n., and Synstrophus repandus (Horn). A lectotype is designated for Eus- trophus brunneimarginatus Dury. A key is given to separate genera and species, supplemented with illustra- tions of relevant features, including aedeagi of all Nearctic species of Eustrophopsis. Detailed distribution (including Mexican records) and natural history data are provided. Keywords Tetratomidae, new species, Arizona, natural history, Nearctic, faunal review Introduction The Eustrophinae, like many groups within Tenebrionoidea, have not had a stable family-level placement until fairly recently. Various authors have commented on the taxonomic history of the group (e.g. Lawrence 1991; Young and Pollock 2002; Pollock 2008; Lawrence and Leschen 2010); however, the recent placement of eustrophines within an enlarged Tetratomidae (as proposed by Nikitsky 1998), is based in large part Copyright Darren A. Pollock. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Darren A. Pollock / ZooKeys 188: 1–153 (2012) on characteristics of larvae. Very few comprehensive phylogenetic analyses for Ten- ebrionoidea have been completed; the study by Lawrence et al. (2011), based on mor- phological characters of adults and larvae of the entire Coleoptera, used three exem- plar taxa from Tetratomidae, including Eustrophopsis bicolor (Fabricius). Monophyly of Tetratomidae was not supported in this analysis; in fact, Eustrophopsis was shown to be the sister taxon to Orchesia (Melandryidae). Tetratoma (Tetratomidae: Tetratominae) and Penthe (Tetratomidae: Penthinae) were part of a monophyletic group that also contained Dipsaconia (Ulodidae) and Chalcodrya (Chalcodryidae). An earlier study by Beutel and Friedrich (2005) hypothesized Tetratomidae (data taken from the litera- ture) as part of a very large, unresolved branch of basal Tenebrionoidea. There have been no large-scale studies of the Eustrophinae specifically, although Nikitsky (1998) provided an overview of the classification of the entire family Tetrato- midae. Also, Nikitsky (2007) revised the genus Holostrophus, representing the only published work on a single genus of Eustrophinae. Pollock (2008) reviewed the Ca- nadian eustrophine fauna, as part of a volume covering biodiversity, systematics, and ecology of Canadian Coleoptera. The present study treats all taxa found in North America, north of Mexico, including the description of a new species from southern Arizona. Much of the descriptive, ecological and distributional data for species occur- ring in Canada, treated herein, were given previously by Pollock (2008). As in Pollock (2008), the present study is not meant to be a detailed revision; rather, it is a review of a specific biogeographic group of Eustrophinae. Therefore, descriptions are rather diagnostic in nature, except for the description of Eustrophopsis crowdyi, new species, which is more detailed and complete. Nikitsky (1998) provided a key to the five recognized subfamilies of Tetratomidae. The Eustrophinae have the following characteristics: body elongate oval, often more distinctly narrowed posteriorly; all legs with simple, narrow tarsi; antennomeres 3-7 more or less broadened, at least some antennomeres transverse; pronotum with two sublinear impressions basally, metepisterna distinctly subdivided into two sections; metendosternite without laminae. The world fauna of Eustrophinae comprises 5 genera and 86 species (compiled from various sources), distributed worldwide except for Australia and New Zealand. The greatest diversity occurs in tropical areas, especially in the Neotropical region. Members of the subfamily are relatively homogeneous structurally, and both larvae and adults are associated with various groups of fungi on dead trees (Lawrence 1991; Pollock 2008). No studies are available that detail the ecological habits of any spe- cific taxon of Eustrophinae. Most information available is taken from collection re- cords. Lawrence (1991: 506) gave the following fungal associates with some genera and species of Eustrophinae: Eustrophopsis spp. (Polyporus squamosus, Tyromyces albel- lus, Dichomitus squalens, Laetiporus sulphureus, Bjerkandera adusta, Inonotus munzii, I. vulpinus, Phellinus gilvus, Pleurotus sapidus, Panus rudis, Lentinus lepideus); Synstrophus repandus (Horn) (Pleurotus ostreatus, Phaeolus schweinitzii, and Coriolus versicolor); Eu- strophus tomentosus Say (Polyporus squamosus). Individual fungal/collection records are given below for each species (from label data). Review of the Eustrophinae (Coleoptera, Tetratomidae) of America north of Mexico 3 Adults of Eustrophinae are active at night, when they emerge from daytime hid- ing places and occupy exposed surfaces, on dead wood or on associated fungal bodies. Very long series of specimens are somewhat rare; collecting during the day will yield few specimens, whereas night collecting or mass-trapping can be very productive and effective methods for obtaining long series. During the day, potentially good, fungus- infested dead trees can be marked (for example, by using thumbtacks with reflective paint) and returned to at night, to collect the active adults. Adults of Eustrophopsis are seemingly attracted to Lindgren funnel traps, and/or the sc- olytine baits often deployed along with these devices. For example, a single locality (Arizo- na, Cochise Co., Chiricahua Mts., Turkey Creek, 2000 m, 31°51.280'N, 109°19.883'W) yielded the following specimens over a two-week period: 7.vii.2002 (E. arizonensis: 54♂♂, 104♀♀; E. ornatus: 7♂♂; E. crowdyi, sp. n.: 10♂♂, 11♀♀); 14.vii.2002 (E. arizonensis, 45♂♂, 92♀♀; E. ornatus: 7♂♂, 17♀♀; E. crowdyi, sp. n.: 16♂♂, 11♀♀). As in the majority of Tenebrionoidea, immature stages of Eustrophinae are rather imperfectly known. Larvae are found in the same habitats as adults; however, whereas adults are often found on surfaces at night, larvae remain within the rotting wood/ fungal substrate (Pollock pers. observ.). Nikitsky (1998) provided a key to subfami- lies of tetratomid larvae; however, only Eustrophopsis and Holostrophus were includ- ed. A key to larvae of Holostrophus (three Oriental species only) was also given by Nikitsky (1998). The most detailed description of a larva of Eustrophopsis is that of Węgrzynowicz (1995) who described the larva of E. quindecimmaculata (Laporte), a Neotropical species. Among Nearctic eustrophines, the only species reasonably well known in the larval stage is Eustrophopsis bicolor (Fabricius); this is the most common species, so perhaps this is not surprising. Weiss (1919) provided a brief description of the larva, along with some reasonably detailed information on host preference and natural history. Böving and Craighead (1931) treated the larva of E. bicolor also, but only illustrated the posterior end of the abdomen (based on larval characters, they removed Eustrophus from Melandryidae and transferred it to Dacnidae). Lawrence (1991) illustrated the dorsal habitus of the larva, along with an SEM of the larval mandible. Lawrence and Leschen (2010) provided a comprehensive description of the larvae of Tetratomidae, with various details given for Eustrophinae (based on Eustrophopsis and Holostrophus). Materials and methods This study is based on 6,600 adult specimens examined and/or borrowed from the fol- lowing private and public collections and museums: AMNH American Museum of Natural History, New York, NY BBC Brian Baldwin Collection (private) BYUC Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT CARR J.B. and A. Carr Collection (now part of CNC) 4 Darren A. Pollock / ZooKeys 188: 1–153 (2012) CAS California Academy of Sciences, San Francisco, CA CBC Cheryl Barr Collection (private) CGMC Chris Majka Collection (private) CMNC Canadian Museum of Nature, Gatineau, QC CMNH Carnegie Museum of Natural History, Pittsburgh, PA CNC Canadian National Collection of Insects, Ottawa, ON CNHM Cincinnati Natural History Museum, Cincinnati, OH CUIC Cornell University, Ithaca, NY DAPC Darren A. Pollock Collection (private)
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