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The Distribution and Evolution of Exocrine Compound

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The Distribution and Evolution of Exocrine Compound This article was downloaded by: [79.238.118.44] On: 17 July 2013, At: 23:24 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Annales de la Société entomologique de France (N.S.): International Journal of Entomology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tase20 The distribution and evolution of exocrine compound glands in Erotylinae (Insecta: Coleoptera: Erotylidae) Kai Drilling a b , Konrad Dettner a & Klaus-Dieter Klass b a Department for Animal Ecology II , University of Bayreuth, Universitätsstraße 30 , 95440 , Bayreuth , Germany b Senckenberg Natural History Collections Dresden , Museum of Zoology , Königsbrücker Landstraße 159, 01109 , Dresden , Germany Published online: 24 May 2013. To cite this article: Kai Drilling , Konrad Dettner & Klaus-Dieter Klass (2013) The distribution and evolution of exocrine compound glands in Erotylinae (Insecta: Coleoptera: Erotylidae), Annales de la Société entomologique de France (N.S.): International Journal of Entomology, 49:1, 36-52, DOI: 10.1080/00379271.2013.763458 To link to this article: http://dx.doi.org/10.1080/00379271.2013.763458 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. 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Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Annales de la Société entomologique de France (N.S.), 2013 Vol. 49, No. 1, 36–52, http://dx.doi.org/10.1080/00379271.2013.763458 The distribution and evolution of exocrine compound glands in Erotylinae (Insecta: Coleoptera: Erotylidae) Kai Drillinga,b*, Konrad Dettnera & Klaus-Dieter Klassb aDepartment for Animal Ecology II, University of Bayreuth, Universitätsstraße 30, 95440 Bayreuth, Germany; bSenckenberg Natural History Collections Dresden, Museum of Zoology, Königsbrücker Landstraße 159, 01109 Dresden, Germany (Accepté le 22 mars 2012) Summary. Members of the family Erotylidae and especially of the subfamily Erotylinae possess a whole arsenal of compound integumentary glands. Their external pores are located in several parts of the body, mainly in the corners and along the lateral margins of the pronotum, beside the compound eyes (periocular glands), on the subgenal braces (subocular glands), on the abdominal ventrites, and more rarely on the prosternal and mesoventral intercoxal processes, the mentum and the mandibles. To evaluate the evolution of the various exocrine glands their occurrence is described for 55 species of Erotylidae (including the former Languriidae), and data from the literature are included for nine further species and two subfamilies. In some phylogenetically crucial cases, the glandular nature was verified by internal inspection (search for glandular ducts), and in some critical species or genera an extended sample was studied. Gland characters and their phylogenetic implications are discussed and mapped on a previously published erotylid phylogeny. Résumé. La distribution et l’évolution des glandes exocrines composées chez les Erotylinae (Insecta: Coleoptera: Erotylidae). Les membres de la famille des Erotylidae et spécialement de la sous-famille des Erotylinae ont un large arsenal de glandes tégumentaire composées. Leurs pores externes sont situés dans différentes parties du corps, surtout dans les coins et les bords des marges latérales du pronotum, à côté des yeux composés (glandes péri-oculaires), sur les ventrites abdominaux, et plus rarement sur les processus prosternaux intercoxal et mésoventral, le mentum et les mandibules. Pour évaluer l’évolution des différentes glandes exocrines, leur occurrence est décrite pour 55 espèces d’Erotylidae (y compris les anciens Languriidae), tandis que des données de la littérature sont incluses pour 9 espèces supplémentaires et 2 sous-familles. Dans certains cas phylogénétiquement cruciaux, la nature glandulaire a été vérifiée par un examen interne (recherche de canaux glandulaires). L’échantillonnage a été étendu pour certaines espèces ou certains genres critiques. Les caractères des glandes et leurs implications phylogénétiques sont discutées et cartographiées à partir d’une phylogénie des Erotylidae précédemment parue. Keywords: languriidae; dermal glands; phylogeny; SEM The cosmopolitan Erotylidae comprises about 3500 tionships in Cucujiformia are widely unresolved and described species in ca. 258 genera (Leschen et al. 2010). Cucujoidea are unlikely to be a monophyletic group (see They are classified into six subfamilies: Cryptophilinae, Leschen et al. 2005; Hunt et al. 2007; Buder et al. 2008; Downloaded by [79.238.118.44] at 23:24 17 July 2013 Pharaxonothinae, Languriinae, Xenoscelinae, Loberinae, and Lawrence et al. 2011). Moreover, the phylogenetic rela- Erotylinae. These subfamilies (excl. Erotylinae) previously tionships of Erotylidae to other cucujiform taxa are still formed a separate family, “Languriidae”; the family previously ambiguous. Various families were regarded as the closest known as Erotylidae is now ranked as the subfamily relatives of Erotylidae in the past: members of the cerylo- Erotylinae. This subfamily is further classified into five tribes: nid series of Cucujoidea (Alexiidae, Endomychidae; Dacnini, Tritomini, Erotylini, Encaustini, and Megalodacnini Crowson 1955; Sen Gupta & Crowson 1971), certain (Węgrzynowicz 2002; Leschen 2003; Leschen et al. 2010). “lower Cucujoidea” (Cryptophagidae, Propalticidae; Sen Erotylidae is a subgroup of the cucujiform beetles, Gupta & Crowson 1969, 1971; Leschen 1996, 2003; which are supported as monophyletic by a number of McHugh et al. 1997; Robertson et al. 2004) as well as autapomorphies (e.g., Lawrence & Newton 1982; Phloeostichidae, Lamingtoniidae (Leschen 2003), and Klausnitzer 2005; Leschen & Ślipiński 2010) and also by Biphyllidae (Sen Gupta & Crowson 1971; Leschen the extensive molecular study of Hunt et al. (2007). 2003; Leschen & Buckley 2007). The molecular studies However, the large-scale morphological analysis by by Hunt et al. (2007) suggest either Monotomidae, Lawrence et al. (2011) found Erotylidae to be paraphyletic. Helotidae, and Protocucujidae as the closest relatives Within Cucujiformia the Erotylidae are generally of Erotylidae (supporting fig. S1 therein: Bayesian analy- assigned to Cucujoidea. However, basal phylogenetic rela- sis), or a clade comprising Laemophloeidae, Phalacridae, *Corresponding author. Email: [email protected] © 2013 Société entomologique de France Annales de la Société entomologique de France (N.S.) 37 Propalticidae, and Cucujidae (supporting fig. S4 therein: (1) In cases where our results differed from those in parsimony analysis). Węgrzynowicz (2002; Table 1), we examined several fi It is well known that Erotylidae show a particularly rich specimens from different localities to test for intraspeci c variability. This concerns six species: Toramus pilifer,3× equipment of compound exocrine glands. Glands can occur in Taraz (Kazakhstan), 2× Margilan (Uzbekistan); Dacne the corners and along the lateral margins of the pronotum, on bipustulata, 2× Saxon Switzerland (Germany), 2× the prosternal and mesoventral intercoxal processes, on the Trenčin (Slovakia), 2× from near Berlin (Germany); head anteromesal to the compound eyes, on the subgenal Triplax russica, 2× Magdeburg (Germany), 2× Saxon ň brace, on the abdominal ventrites, and rarely on the mentum Switzerland (Germany), 2× Kada (Czech Republic); Tritoma bipustulata, 4× Saxon Switzerland (Germany), and on the mandibles. Despite this manifold occurrence of 6× Bayreuth (Germany), 5× Trenčin (Slovakia); such glands over the beetles’ body, their consideration in the Pselaphacus nigropunctatus (species identification con- previous literature is quite sparse. The distribution of several firmed by P. Węgrzynowicz), 2× Peru, 3× São Paulo de gland pores across erotylid taxa was included in the phyloge- Olivença (Brazil), 2× Bogota (Colombia); Megalodacne netic studies of Węgrzynowicz (2002; mostly on the erotyline fasciata, 2× Texas (USA), 2× Chicago (USA), 1× New “ ” Orleans (USA). tribes) and Leschen (2003; mostly on the former languriid (2) In the single case where Węgrzynowicz’s (2002) and subfamilies). In both papers many taxa are scored with regard our results differ for species from the same genus we to the occurrence of glands, but the pictorial documentation is studied
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