ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Arthropod Systematics and Phylogeny Jahr/Year: 2017 Band/Volume: 75 Autor(en)/Author(s): Simonsen Thomas J., Kristensen Nils Peter Artikel/Article: Revision of the endemic Brazilian ‘neotheorid’ hepialids, with morphological evidence for the phylogenetic relationships of the basal lineages of Hepialidae (Lepidoptera: Hepialoidea) 281-301 75 (2): 281 – 301 8.9.2017 © Senckenberg Gesellschaft für Naturforschung, 2017. Revision of the endemic Brazilian ‘neotheorid’ hepialids, with morphological evidence for the phylogenetic rela­ tionships of the basal lineages of Hepialidae (Lepido­ ptera: Hepialoidea) Thomas J. Simonsen *, 1, 2 & Niels P. Kristensen 3, # 1 Natural History Museum Aarhus, Wilhelm Meyers Allé 10, DK-8000 Aarhus C, Denmark; Thomas J. Simonsen * [[email protected]] — 2 Natural History Museum London, Cromwell Road, London, SW7 5BD, United Kingdom — 3 Natural History Museum of Denmark (Zoology), Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark — * Corresponding author # This study was initiated as a collaborative effort between the authors in 2011. Sadly, Niels P. Kristensen passed away December 06, 2014, before the manuscript could be completed. He had, however, completed an extensive draft of the taxonomic sections of the text. It therefore seems most fitting that he remains first author of the names proposed here. Accepted 02.v.2017. Published online at www.senckenberg.de/arthropod-systematics on 30.viii.2017. Editors in charge: Ulrike Aspöck & Klaus-Dieter Klass Abstract We revise the little known South American primitive Hepialidae genus Neotheora Kristensen, and describe two new species, N. meyi sp.n. and N. mielkeorum sp.n. from Brazil based on two female singletons. Furthermore, we describe a new genus, Paratheora gen.n., with a single species, P. speideli sp.n., of primitive Hepialidae from Brazil based on one male, and a female without the postabdomen. Although the new genus is clearly closely related to Neotheora as demonstrated by a unique tentorium modification found in both genera, it is also sufficiently different to warrant its own genus. To explore the phylogenetic relationships of Paratheora, we carried out a phylogenetic analysis of 16 terminal taxa in Hepialidae sensu lato based on 18 morphological characters scored from adult morphology. The results con- firm that Neotheora and Paratheora are sister taxa, and together these two ‘neotheorid genera’ comprise the sister group of the remaining Hepialidae s.lat. Within the latter, the Australian genus Anomoses Turner is the sister group of the remaining taxa. The southern African genus Prototheora Meyrick is the sister group of a clade comprising the four ‘palaeosetid genera’, the four so-called ‘primitive’ Hepialidae genera, and Hepialidae s.str. The ‘palaeosetid genera’ comprise a monophyletic clade, which is the sister group to a clade comprising the ‘primitive’ Hepialidae, and Hepialidae s.str. While the exact relationships between the four ‘primitive’ Hepialidae, and Hepialidae s.str. cannot be resolved based on our data, the two ‘primitive’ hepialids from sub-Saharan Africa, Afrothora Nielsen & Scoble and Antihepialus Janse are likely sister taxa, and the Holarctic Gazoryctra Hübner is likely the sister of that clade. Similarly, the four genera selected to represent Hepialidae s.str. appear monophyletic in the analysis. Key words Neotheora, new species, new genus, Hepialoidea phylogeny. 1. Introduction Hepialoidea (sensu REGIER et al. 2015) are the most eco- cies adapted to a wide variety of lifestyles and habitats logically diverse and species rich lineage within the basal (COMMON 1990; KRISTENSEN 1998; REGIER et al. 2015). lepidopterous grade Homoneura, several hundred spe- The superfamily comprises more than 650 species in ca ISSN 1863-7221 (print) | eISSN 1864-8312 (online) 281 Simonsen & Kristensen: Revision of ‘neotheorid’ Hepialidae 70 genera (SIMONSEN in press; GREHAN & RAWLINS 2016; in the palaeosetid Ogygioses caliginosa Issiki & Stringer, VAN NIEUKERKEN et al. 2011). Traditionally, most authors 1932 the only palaeosetid for which there were detailed have considered Hepialoidea to be one of two superfami- morphological details know at the time. An enlarged in- lies in the suborder Exoporia, with the other superfamily tercalary sclerite similar to the structure found elsewhere being the monogeneric New Zealand endemic Mnesar- in Hepialoidea (and believed to be an autapomorphy for chaeoidea (e.g. KRISTENSEN 1998; NIELSEN et al. 2000; the family) has since been found in all other palaeosetid VAN NIEUKERKEN et al. 2011; GREHAN & RAWLINS 2016). species (including other Ogygioses) (KRISTENSEN 1998). Following that classification, Hepialoidea comprise five NIELSEN & SCOBLE (1986) removed Afrotheora, Antihe­ families: Neotheoridae, Anomosetidae, Prototheoridae, pialus, Fraus and Gazoryctra from Hepialidae s.str. as Palaeosetidae, and Hepialidae. However, the monophyly these genera lack the close association (often synscleroti- of the non-monogeneric families Palaeosetidae and He- sation) between the trulleum and pseudoteguminal arms pialidae s.lat. remains questionable, and Hepialoidea found in Hepialidae s.str. NIELSEN & KRISTENSEN (1989) (sensu NIELSEN et al. 2000) has generally been considered revised the Australian genus Fraus and carried out very to comprise 12 terminal taxa, viz. Neotheora Kristensen, detailed studies of the morphology of the genus, result- 1978 (Neotheoridae), Anomoses Turner, 1916 (Anomo- ing in a total revision of our understanding of the mor- setidae), Prototheora Meyrick, 1917 (Prototheoridae), phology (in particular in the abdomen) of Hepialoidea. Genustes Issiki & Stringer, 1932, Ogygioses Issiki & KRISTENSEN & NIELSEN (1993) redescribed the enigmatic Stringer, 1932, Oshroes Druce, 1900, Palaeoses Turner, South American palaeosetid genus Oshroes (known only 1922 (all Palaeosetidae), Afrotheora Nielsen & Scoble, from three specimens) and noted the potential synapo- 1986, Antihepialus, Janse, 1942, Fraus Walker, 1856, morphies, which it shares with the two southeast Asian Gazoryctra Hüber, 1820 (the so-called “primitive” He- genera Genustes and Ogygioses. DAVIS et al. (1995) re- pialidae), and Hepialidae s.str. (ca 60 genera) (NIELSEN & vised Ogygioses and, while questioning the monophyly SCOBLE 1986; KRISTENSEN 1998; NIELSEN et al. 2000; VAN of Palaeosetidae, suggested that Ogygioses and Genustes NIEUKERKEN et al. 2011). Recently, REGIER et al. (2015) were potential sister taxa. DAVIS (1996) revised the south- formally synonymised all hepialoid families with Hepial- ern African family Prototheoridae and synonymised the idae based on a comprehensive molecular dataset, which genus Metatheora Meyrick, 1919 with Prototheora thus nevertheless was missing some key taxa. Mnesarchaei- reducing the family to a single genus. KRISTENSEN (1998) dae was included in Hepialoidea, making the suborder summarised the systematics of Hepialoidea and repeated name Exoporia redundant. the doubt over the monophyly of Palaeosetidae. As Hepialoidea (together with the small Australian While Hepialidae s.str. are almost global in their dis- family Lophocoronidae) are the likely sister group to the tributions (with the exception of Madagascar), the re- Heteroneura (e.g. REGIER et al. 2015; BAZINET et al. 2016 maining genera are more restricted to specific regions: – but see KRISTENSEN et al. 2015 for an alternative hypo- southern Brazil (Neotheora, Paratheora gen.n.), Austral- thesis), a group that includes over 99% of all Lepidoptera ia (Anomoses, Palaeoses, Fraus), Columbia (Oshroes), (> 150,000 species) (KRISTENSEN 1998; KRISTENSEN et al. south-east Asia from Assam to Taiwan (Genustes, Ogygi­ 2007; VAN NIEUKERKEN et al. 2011), comparative studies oses), sub-Saharan Africa (Prototheora, Afrotheora, An­ of the group are of considerable value for understanding tihepialus), and temperate northern hemisphere forests the evolution of Lepidoptera as a whole. However, the (Gazoryctra); Mnesarchaea Meyrick, the only genus in phylogenetic and evolutionary relationships of Hepia- Mnesarchaeidae, the other family in Hepialoidea, is en- loidea are poorly understood, although several recent re- demic to New Zealand (HOLLOWAY & NIELSEN 1998). gional faunistic and systematic studies exist (e.g. DUGDA- Neotheoridae was established more than three decades LE 1994; BROWN et al. 2000a,b; MIELKE & GREHAN 2012; ago to accommodate the genus Neotheora with a single GREHAN & RAWLINS 2016; SIMONSEN 2015, in press), and species N. chiloides Kristensen, 1978 (Fig. 1A) known recent studies of adult morphology of Hepialidae s.str. from only a single female specimen from Mato Grosso, (GREHAN 2010, 2012) have provided new insight and pro- Brazil (KRISTENSEN 1978a). Evidently, the very incom- gress into phylogenetic relationships of Hepialidae s.str. plete available morphological information about Neothe­ A global check list and bibliography of Hepialoidea was ora has remained a major impediment for assessing the presented by NIELSEN et al. (2000). affinities of this taxon: alone among the terminal hepialid One problem encountered when attempting to resolve taxa it has not been the subject of comparative studies the phylogeny of Hepialidae s.str. is that the evolutionary since it was originally described. We here describe two relationships of the 12 terminal Hepialidae s.lat. taxa are additional species of the genus