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DE TTK 1949 Taxonomy and Systematics of the Eurasian DE TTK 1949 Taxonomy and systematics of the Eurasian Craniophora Snellen, 1867 species (Lepidoptera, Noctuidae, Acronictinae) Az eurázsiai Craniophora Snellen, 1867 fajok taxonómiája és szisztematikája (Lepidoptera, Noctuidae, Acronictinae) PhD thesis Egyetemi doktori (PhD) értekezés Kiss Ádám Témavezető: Prof. Dr. Varga Zoltán DEBRECENI EGYETEM Természettudományi Doktori Tanács Juhász-Nagy Pál Doktori Iskola Debrecen, 2017. Ezen értekezést a Debreceni Egyetem Természettudományi Doktori Tanács Juhász-Nagy Pál Doktori Iskola Biodiverzitás programja keretében készítettem a Debreceni Egyetem természettudományi doktori (PhD) fokozatának elnyerése céljából. Debrecen, 2017. ………………………… Kiss Ádám Tanúsítom, hogy Kiss Ádám doktorjelölt 2011 – 2014. között a fent megnevezett Doktori Iskola Biodiverzitás programjának keretében irányításommal végezte munkáját. Az értekezésben foglalt eredményekhez a jelölt önálló alkotó tevékenységével meghatározóan hozzájárult. Az értekezés elfogadását javasolom. Debrecen, 2017. ………………………… Prof. Dr. Varga Zoltán A doktori értekezés betétlapja Taxonomy and systematics of the Eurasian Craniophora Snellen, 1867 species (Lepidoptera, Noctuidae, Acronictinae) Értekezés a doktori (Ph.D.) fokozat megszerzése érdekében a biológiai tudományágban Írta: Kiss Ádám okleveles biológus Készült a Debreceni Egyetem Juhász-Nagy Pál doktori iskolája (Biodiverzitás programja) keretében Témavezető: Prof. Dr. Varga Zoltán A doktori szigorlati bizottság: elnök: Prof. Dr. Dévai György tagok: Prof. Dr. Bakonyi Gábor Dr. Rácz István András A doktori szigorlat időpontja: 2016. december 7. Az értekezés bírálói: Dr. ................................................ Dr. ................................................ Dr. ................................................ A bírálóbizottság: elnök: Dr. ................................................ tagok: Dr. ................................................ Dr. ................................................ Dr. ................................................ Dr. ................................................ Az értekezés védésének időpontja: 200…. ……………… …. Contents Introduction ................................................................................................... 1 Taxonomic review .......................................................................................... 2 Objectives ....................................................................................................... 5 Material and methods .................................................................................... 5 Genital dissections ........................................................................................ 7 Terminology of morphological characters ..................................................... 7 Geometric morphometrics ........................................................................... 16 Results .......................................................................................................... 18 Taxonomic part ........................................................................................... 18 The Craniophora generic complex .......................................................... 19 Genus Craniophora Snellen, 1867 ........................................................ 19 Genus Harmandicrania Kiss, 2017 ....................................................... 34 Genus Graesericrania Kiss, 2017 ......................................................... 51 Genus Eurypterocrania Kiss, 2017 ....................................................... 53 The Cycloprora generic complex ............................................................. 57 Genus Cycloprora Turner, 1920 ........................................................... 57 Genus Turnerinycta Kiss, 2017 ............................................................ 59 Genus Fascionycta Kiss, 2017 .............................................................. 60 Genus Megalonycta Viette, 1965 .......................................................... 62 Genera and species separated from the Craniophora and Cycloprora generic complex ...................................................................................... 72 Genus Berionycta Kiss, 2017 ................................................................ 72 Genus Draudtinycta Kiss, 2017 ............................................................ 74 Genus Sinonycta Kiss, 2017 ................................................................. 75 Genus Miracopa Draudt, 1950.............................................................. 77 Genus Acronicta Ochsenheimer, 1816 .................................................. 78 Incertae sedis ....................................................................................... 80 Male genitalia variability in Craniophora ligustri ....................................... 91 Taxonomical and biogeographical considerations ...................................... 94 Summary ...................................................................................................... 98 Összefoglalás .............................................................................................. 100 Acknowledgements..................................................................................... 103 References .................................................................................................. 104 Appendices ................................................................................................. 117 Appendix 1 ............................................................................................... 118 Appendix 2 ............................................................................................... 118 Appendix 3 ............................................................................................... 118 Appendix 4 ............................................................................................... 118 Appendix 5 ............................................................................................... 118 Introduction Lepidoptera is one of the largest orders of insects with approximately more than 150 thousand described species (NIEUKERKEN et al. 2011). The first lepidopterans have already appeared in the Late Triassic and most of the main lineages have evolved during the Upper Cretaceous, following the radiation of the flowering plants (GRIMALDI & ENGEL 2005, WAHLBERG et al. 2013). Noctuidae is commonly regarded as one of the largest lepidopteran family (FIBIGER & LAFONTAINE 2005, MITCHELL et al. 2006, ZAHIRI et al. 2010) with more than 11 000 described taxa (NIEUKERKEN et al. 2011). Such a high number is resulted from the immense radiation of higher Ditrysian clades as a consequence of the diversification of the vegetation mostly in the Upper Tertiary (GRIMALDI & ENGEL 2005, WAHLBERG et al. 2013). The species of the subfamily Acronictinae, one of the phylogenetically basal subfamilies of Noctuidae (ZAHIRI et al. 2013), have trifine hindwing venation with reduced or vestigial M2 vein (FIBIGER & LAFONTAINE 2005, MITCHELL et al. 2006, ZAHIRI et al. 2013). The Eurasian species of the subfamily are traditionally divided into two main genus groups (Acronicta Ochsenheimer, 1816 and its relatives vs. Craniophora Snellen, 1867), based on the external morphology and genital characters (FIBIGER et al. 2009, KONONENKO 2010). Four genera (Belciades Kozhantshikov, 1950, Gerbathodes Warren, 1911, Moma Hübner, [1820] and Nacna Fletcher, 1961), however, have been recently removed from the subfamily (ROTA et al. 2016). According to ROTA et al. (2016), the genus Craniophora s. str. forms a monophyletic group together with Craniophora jankowskii (Oberthür, 1880) (in fact this species belongs to the genus Cranionycta de Lattin, 1949) and Chloronycta Schmidt & Anweiler, 2014, nesting with the rest of basal lineages of the subfamily. On the other hand, these basal groups form the sister-group of the genus Acronicta s. l. The systematic processing of the available material of Craniophora species has been started in 2011 with my PhD grant, however the initial phases were time-consuming due to the much dispersed material. That time, the basic concept was that Acronicta is a well-defined genus with some subgenera, at least in the Palaearctic Region, while Craniophora is probably a collecting genus. On the other hand, only few authors have hitherto dealt with Craniophora species systematically (e.g. Han & Kononenko 2010). Nowadays it has been shown that both Acronicta and Craniophora are generic complexes, and they contain more phylogenetic lineages (see ROTA et al. 2016 and KISS 2017). 1 Taxonomic review The genus Craniophora was established by SNELLEN (1867) for the Palaearctic species Noctua ligustri [Denis & Schiffermüller], 1775 by monotypy. CHAPMAN (1890) erected the genus Bisulcia also for N. ligustri so it is an objective synonym of Craniophora. More than 60 years later, BOURSIN (1952) synonymized the genus Miracopa Draudt, 1950 with the genus Cranionycta (see also SUGI 1959). Twelve years later, BOURSIN (1964) also synonymized the genera Cranionycta, Hampsonia Kozhantshikov, 1950 and Hampsonidia Inoue, 1958 with Craniophora, but without any argumentation. POOLE (1989), in his catalogue, probably followed this last work of BOURSIN (1964) and considered Craniophora as a common genus and specified 22 species with their synonyms. At this time, but independently, HOLLOWAY (1989) synonymized the genus Meglonycta (sic!) Viette, 1965 (correctly Megalonycta)
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