Primitive Ghost Moths: Morphology and taxonomy of the Australian genus Fraus Walker (Lepidoptera: Hepialidae s. lat.)
Abstract
Hepialidae (ghost moths or swifts) are, in terms of diversity and distribution, the most successful group of homoneurous primitive moths. The morphology of Fraus is described in some detail with emphasis on the adult moth, and a new interpretation of hepialid male genitalia is presented. Beyond describing and illustrating a primitive hepialid, these observations are intended to serve as reference for the study of the classification of Hepialoidea and lower Lepidoptera.In the taxonomic revision, based on more than 3000 specimens, the 25 Fraus species are described and diagnosed. The adult moths, as well as male and female genitalia, are richly illustrated, and distribution maps and flight period diagrams are provided for all species. The biology, behaviour, distribution and phylogeny are summarised and discussed.
... On the anal segment (AIO), posterior part ofplanta indented and devoid of crochets in the middle, so that crochets are arranged in "figure-of-3" (cf. Nielsen & Kristensen, 1989); with four anterior uniordinal rows of crochets (with traces of an additional external), the innermost stronger, reduced to one in posterior half. CHAETOTAXY As no unequivocal notation for the lepidopteran setae was proposed and no one author has thoroughly worked out all the body parts of the hepialid larvae, the following subjective combination of different systems is here adopted: head (Hasenfuss, 1969), maxillolabium (Gerasimov, 1952), trunk (Stehr, 1987;Wagner, 1987), legs and thoracic pores (Nielsen & Kristensen, 1989). ...
... Nielsen & Kristensen, 1989); with four anterior uniordinal rows of crochets (with traces of an additional external), the innermost stronger, reduced to one in posterior half. CHAETOTAXY As no unequivocal notation for the lepidopteran setae was proposed and no one author has thoroughly worked out all the body parts of the hepialid larvae, the following subjective combination of different systems is here adopted: head (Hasenfuss, 1969), maxillolabium (Gerasimov, 1952), trunk (Stehr, 1987;Wagner, 1987), legs and thoracic pores (Nielsen & Kristensen, 1989). A minor modification is retention of original Hinton's (1946) notation G 2 for Hasenfuss' (1969) 0 3 . ...
... The naming of the MSD and MDI group on meso-and metathorax is quite controversial (e.g. Hardy, 1973;Nielsen & Kristensen, 1989) and was not thoroughly worked out by Stehr (1987) and Wagner (1987); accordingly Hinton's (1946) notation is maintained (cf. also Wagner et al., 1989). ...
The larva of the apenninic endemism Pharmacis aemilianus (Costantini, 1911), so far unknown, is described and figured according to its chaetotaxy. General morphology and setal pattern correspond to the hepialid groundplan, the larva particularly resembling that of P. fusconebulosa (de Geer, 1778), from which it can be easily separated, however, by the distance between SD1 and L1 on A9 which is shorter than that between DI and D2, and the absence of prominent spikes along the lateral margins of labrum.
... Studies of the oldest divergences within the insect order Lepidoptera, based on comprehensive examination of internal as well as external anatomy, represent prominent early applications of Hennigian phylogenetics (Hennig, 1953;Kristensen & Nielsen, 1983;Kristensen, 1984;Davis, 1986;Kobayashi & Ando, 1988;Nielsen & Kristensen, 1996; reviews in Kristensen & Skalski, 1998;Kristensen et al., 2007). In contrast to those of the mega-diverse derived clade Ditrysia, which contains 98% of lepidopteran species (van Nieukerken et al., 2011a), the so-called nonditrysian superfamilies are mostly species-poor but highly divergent in morphology, offering numerous synapomorphies and hence some of the strongest evidence for higher-level relationships in any insect order. ...
... (B) Most parsimonious tree for combined 18S rDNA and nt1 + nt2 of PEPCK, from Wiegmann et al. (2000). Bootstrap values, when > 50%, above branches; number of exemplar species in parentheses after taxon name if > 1. (C) Most parsimonious tree for combined 18S rDNA and ground-plan morphological traits scored for major clades by Nielsen & Kristensen (1996) and Krenn & Kristensen (2000), from Wiegmann et al. (2002). Bootstrap values, when > 50%, above branches. ...
... The molecular data provide little clear evidence on the basal divergences within Glossata, in striking contrast to the morphological analysis of Nielsen & Kristensen (1996). A basal split between Eriocraniidae (node 9) and all other Glossata (= clade Coelolepida; node 12; BP = 38, degen1) does occur in the best tree for 19 genes/degen1. ...
Within the insect order Lepidoptera (moths and butterflies), the so-called nonditrysian superfamilies are mostly species-poor but highly divergent, offering numerous synapomorphies and strong morphological evidence for deep divergences. Uncertainties remain, however, and tests of the widely accepted morphological framework using other evidence are desirable. The goal of this paper is to test previous hypotheses of nonditrysian phylogeny against a data set consisting of 61 nonditrysian species plus 20 representative Ditrysia and eight outgroups (Trichoptera), nearly all sequenced for 19 nuclear genes (up to 14 700 bp total). We compare our results in detail with those from previous studies of nonditrysians, and review the morphological evidence for and against each grouping The major conclusions are as follows. (i) There is very strong support for Lepidoptera minus Micropterigidae and Agathiphagidae, here termed Angiospermivora, but no definitive resolution of the position of Agathiphagidae, although support is strongest for alliance with Micropterigidae, consistent with another recent molecular study. (ii) There is very strong support for Glossata, which excludes Heterobathmiidae, but weak support for relationships among major homoneurous clades. Eriocraniidae diverge first, corroborating the morphological clade Coelolepida, but the morphological clades Myoglossata and Neolepidoptera are never monophyletic in the molecular trees; both are contradicted by strong support for Lophocoronoidea + Hepialoidea, the latter here including Mnesarchaeoidea syn.n. (iii) The surprising grouping of Acanthopteroctetidae + Neopseustidae, although weakly supported here, is consistent with another recent molecular study. (iv) Heteroneura is very strongly supported, as is a basal split of this clade into Nepticuloidea + Eulepidoptera. Relationships within Nepticuloidea accord closely with recent studies based on fewer genes but many more taxa. (v) Eulepidoptera are split into a very strongly supported clade consisting of Tischeriidae + Palaephatidae + Ditrysia, here termed Euheteroneura, and a moderately supported clade uniting Andesianidae with Adeloidea. (vi) Relationships within Adeloidea are strongly resolved and Tridentaformidae fam.n. is described for the heretofore problematic genus Tridentaforma Davis, which is strongly supported in an isolated position within the clade. (vii) Within Euheteroneura, the molecular evidence is conflicting with respect to the sister group to Ditrysia, but strongly supports paraphyly of Palaephatidae. We decline to change the classification, however, because of strong morphological evidence supporting palaephatid monophyly. (viii) We review the life histories and larval feeding habits of all nonditrysian families and assess the implications of our results for hypotheses about early lepidopteran phytophagy. The first host record for Neopseustidae, which needs confirmation, suggests that larvae of this family may be parasitoids.This published work has been registered in ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:C17BB79B-EF8F-4925-AFA0-2FEF8AC32876.
... The synonymies proposed by Nielsen (1996) are evaluated based on the results, and a key to all species is provided. The morphological account presented here is more detailed than previous systematic treatments of Hepialidae, except Nielsen & Kristensen's (1989) highly detailed account of the primitive genus Fraus. It is, however, in line with what will be presented in an upcoming revision of the so-called 'hepialine' hepialids in Australia. ...
... Laterally the ridge is synsclerotised with the occipital condyles, and (just below the occipital condyles) with the base of the corporotentorium. Ventrally (beyond the corporotentorium), it continues around the postlabial wart (sensu Nielsen & Kristensen 1989). The occipital condyles are prominent with a welldefined sclerotised base. ...
... Pronotum continues posteriorly over mesonotum as a thin membranous ridge. The episternum (propleural wart of Neilsen & Kristensen 1989) is relatively well defined and scaled, with the anterior edge ventrally produced into a long, narrow downwards pointed episternal tooth. The basal margin of katepisternum is smooth and strongly sclerotised, whereas the majority of katepisternum is a weakly sclerotised, densely scaled area dorsal to coxa. ...
I revise the Australian-New Guinean ghost moth genus Elhamma. Two recent synonymies are assessed, and two new species from New Guinea, E. grehani sp. nov. and E. viettei sp. nov. are described. I provide an updated diagnosis for the genus and conclude that the presence of only 2 M-veins in the hind wing in both sexes (when females are known) and a strongly cup-shape juxta in the male genitalia are unique diagnostic characters among Hepialidae. I give a detailed description of the adult morphology based on male E. australasiae, and provide a key to all known species based on adult male characters.
... Descriptions of the musculature of the head and cervical region were published mostly for some representatives of higher lepidopterans (Geometriformes + Papilionoformes + Bombyciformes + Noctuiformes ) (Eaton, 1961; Ehrlich and Davidson, 1961; Banziger, 1970; Albert and Seabrook, 1973; Buttiker et al., 1996). Some works were devoted to ancient lepidopterans with gnawing mouthparts, for example Micropterigidae (Hannemann, 1956 ), or primary proboscides , such as Eriocraniidae (Kristensen, 1968), Lophocoronidae (Nielsen and Kristensen, 1996), and Hepialidae (Nielsen and Kristensen, 1989; Korzeev, 2001). With such a set of taxa, a unified concept of the evolution of the cephalic muscular system within the entire order could not be developed. ...
... The two pairs of muscles were simultaneously found only in the gnawing mouthparts of Micropterigidae (Hannemann, 1956 ) and in Eriocraniidae possessing the primary proboscis (Kristensen, 1968). In further evolution of lepidopterans the abductors of the mandibles (CMd.2) disappear, whereas their antagonists (CMd.1) were described in all the studied non-ditrysian forms (Nielsen and Kristensen, 1989; Korzeev, 2001 Korzeev, , 2004 ) except Lophocoronidae (Nielsen and Kristensen, 1996). Among the ditrysian species, mandibular muscles have so far been found only in Choristoneura fumiferana Clem. of the family Tortricidae (Albert and Seabrook, 1970) and in 3 species of the family Tineidae: Tinea pallescentella Stt., Scardia boletella F., and Nemapogon granella L. (Korzeev, 2004). ...
... According to Albert and Seabrook (1970) , the muscles described in their material correspond to the mandibular abductors. According to another view (Kristensen and Nielsen, 1989), the weaker mandibular abductors should be the first to disappear during reduction of the mandibular musculature. This assumption is confirmed by the pattern of insertion of these muscles to the rudimentary mandibles in the studied representatives of Cossiformes. ...
Musculature of the head and cervical region in eight lepidopteran species from the families Cossidae, Sesiidae, Limacodidae,
and Zygaenidae (Cossiformes) was studied. Besides, musculature of the anterior cervical region in Brachodes appendiculata Esper (Brachodidae) is discussed. The data on musculature are analyzed and compared with the authors’ earlier results on
a relatively primitive ditrysian moth from the family Tineidae (Korzeev, 2004). The absence of the labro-epipharyngeal musculature
in Cossiformes is confirmed and the presence of a new, indirectly acting secondary labral extensors CMx.5 in Zygaenidae is revealed. These secondary extensors are strongly developed in Papilionomorpha. Cossiformes and Tineidae
have been shown to possess several common plesiomorphic traits in the structure of labrum, while the dilatators of salivarium
within Cossiformes are reduced in number from two pairs to a single pair. The present study shows that the musculature of
the cervical region has remained relatively stable in the evolution of Lepidoptera despite the loss of some muscles in different
taxa. The groundplan of the head musculature is reconstructed for Glossata, Papilionomorpha, Cossiformes, and the “higher”
Lepidoptera from the Geometriformes-Noctuiformes complex of superfamilies. In addition, the groundplan of the musculature
of the cervical region is depicted for Myoglossata, Papilionomorpha, Cossiformes, and the Geometriformes-Noctuiformes complex
of superfamilies.
... Most features similar to Taiwanese species are also recognized in O. moriutii from Vietnam and O. maoershana from mainland China (Owada & Jinbo 2005;present study). However, in O. maoershana, it is noteworthy that the intercalary sclerite of antenna is "normal hepialoid type" (Fig. 3C) as in O. luangensis and almost all genera of the superfamily Hepialoidea including, Palaeosetidae (Osrhoes, Palaeoses and Genustes), Neotheoridae (Neotheora Kristensen, 1978 andParatheora Kristensen &Simonsen, 2017), Anomosetidae (Anomoses Turner, 1916), Prototheoridae (Prototheora Meyrick, 1917) and Hepialidae (Fraus Walker, 1856, Oncopera Walker, 1856 etc.) (Nielsen & Kristensen 1989;Kristensen & Nielsen 1994;Davis et al. 1995;Simonsen & Kristensen 2017;Simonsen 2018). As mentioned by some authors, presence of "primitive lepidopteran type" in three Taiwanese species is considered as an autapomorphic character reversal (Nielsen & Kristensen 1989;Kristensen & Nielsen 1994;Davis et al. 1995;Kristensen 1998Kristensen , 2003Simonsen & Kristensen 2017). ...
... However, in O. maoershana, it is noteworthy that the intercalary sclerite of antenna is "normal hepialoid type" (Fig. 3C) as in O. luangensis and almost all genera of the superfamily Hepialoidea including, Palaeosetidae (Osrhoes, Palaeoses and Genustes), Neotheoridae (Neotheora Kristensen, 1978 andParatheora Kristensen &Simonsen, 2017), Anomosetidae (Anomoses Turner, 1916), Prototheoridae (Prototheora Meyrick, 1917) and Hepialidae (Fraus Walker, 1856, Oncopera Walker, 1856 etc.) (Nielsen & Kristensen 1989;Kristensen & Nielsen 1994;Davis et al. 1995;Simonsen & Kristensen 2017;Simonsen 2018). As mentioned by some authors, presence of "primitive lepidopteran type" in three Taiwanese species is considered as an autapomorphic character reversal (Nielsen & Kristensen 1989;Kristensen & Nielsen 1994;Davis et al. 1995;Kristensen 1998Kristensen , 2003Simonsen & Kristensen 2017). ...
The family Palaeosetidae is newly recorded from mainland China. A new species of the genus Ogygioses, O. maoershana sp. nov., is described from Guangxi Province, China. The adults, male and female genitalia, venation, head and legs structures are illustrated. Some ecological information and a key to all species of the genus are also provided. The phylogenetic position of the new species in the genus Ogygioses is inferred as intermediate between primitive O. luangensis Kristensen and other four derived species based on morphological characters and DNA barcodes sequences.
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... Gazoryctra is recognized as one of four or five 'primitive' or 'basal' genera of Hepialidae (Nielsen et al. 2000;Grehan 2012). The bilobate shape of the valva of the male genitalia support Gazoryctra as a monophyletic genus (Grehan 2012), although a similar, less pronounced feature is also present in the 'primitive' genera Afrotheora Nielsen & Scoble where the valva is proportionately longer (Nielsen & Scoble 1986), Antihepialus antarcticus (Wallengren) where the valva is proportionately broader and angled (Grehan 2012), and Fraus Walker where a basal lobe or spine is present in some species (Nielsen & Kristensen 1989;Beaver & Moore 2020). ...
... But this cannot be resolved here as the female genitalia are undescribed for many Hepialidae genera, including the potentially closely related Afrotheora and Antihepialus Janse. The external female genitalia of Fraus are similar in general form to Gazoryctra, but there is considerable variability in the shape of the dorsal plate and lamella antevaginalis in Fraus (Nielsen & Kristensen 1989). The elliptical shape of the corpus bursa of G. sciophanes is similar to that illustrated by Tshistjakov (1997, fig. ...
The genus Gazoryctra Hübner comprises 10 species in North America and four in northern Eurasia. The remaining diversity of North American Hepialidae is represented by four species of Sthenopis Packard, three species of Phymatopus Wallengren, and one species of Korscheltellus Börner (Nielsen et al. 2000; Grehan & Knyazev 2019). The North American distribution of Gazoryctra extends between Alaska and southern Appalachians and southern Rocky Mountains (Grehan & Mielke 2018). As with other North American Hepialidae, Gazoryctra is absent from much of the southern-central United States where there is ostensibly suitable habitat present as this genus is found in forested regions where it is believed to feed on roots or other organic matter (Schweitzer et al. 2011). This absence may be due to the lack of colonization following regression of inland seas that covered much of this region until the end of the Mesozoic (Grehan & Mielke 2018).
... In recent years there have been several important investigations into Hepialidae in Australia, such as the revision of Fraus Walker (Nielsen & Kristensen 1989), Elhamma Walker (Simonsen 2015), and the monograph on the Australian members of the genera Abantiades Herrich-Schäffer, Aenetus Herrich-Schäffer, Archaeoaenetus Simonsen, Zelotypia Scott, and Oncopera Walker (Simonsen 2018). The only Australian genera to have not undergone a modern revision are Jeana Tindale, and Oxycanus Walker. ...
Three new species of ghost moth, Oxycanus ephemerous sp. nov., O. flavoplumosus sp. nov., and O. petalous sp. nov. are described from South Australia, New South Wales, and southwest Western Australia, respectively. We illustrate these species and compare morphological and molecular (mtDNA COI gene) characters with similar Oxycanus Walker, 1856 species from Australia. Comparative images of Oxycanus subvaria (Walker, 1856), O. byrsa (Pfitzner, 1933), and O. determinata (Walker, 1856) are figured. The type material of the three new species are held in the Australian National Insect Collection, Canberra, the Western Australian Museum, Perth, and in the South Australian Museum, Adelaide. The type specimens of Oxycanus hildae Tindale, 1964 syn. n. were also examined and the taxon is here considered synonymous with O. subvaria. Concerns are raised about the conservation status of all three new species due to few or localised distribution records.
... Female genitalia (Figs 19-21). Longer than wide, tergum IX (= dorsal plates of Nielsen & Kristensen 1989) posterior margin an inverted U with narrow and setose region (anal papillae) from dorsal median to near junction with lamella antevaginalis. Subanal plate trapezoid, lateral surface slightly concave. ...
The hepialid genus Gymelloxes Viette, 1952 is characterized by, and differs from all other genera, by the male genitalia and a combination of characters. G. terea is redescribed due to the brevity of the original description. The male genitalia of Gymelloxes terea (Schaus, 1892) include two features that are unique within the Hepialidae – a posteriorly angled and digitiform tergal lobe, and two shallow, peg-like processes on the posterior margin of the saccus. The male phallus also exhibits minute sclerotized spicules or spots on the apex of an otherwise membranous tube. The external genitalia of the female includes a sclerotized antevaginalis with a broad, dorsally projecting, flat central margin. The species is included within the cibyrine clade of Hepialidae by the structure of the tergosternal connection and narrow spacing between Sc and R on the hindwing. The potential systematic significance of specialized similarities shared with other cibyrine Hepialidae is discussed. A lectotype of Dalaca terea Schaus, 1892 is here designated.
... The sensilla chaetica of the maxillae are tactile (Schoonhoven and Dethier, 1966;Hanson, 1970). In E. kuehneilla larvae, group of microtrichia is present on the maxilla and labium are similar to microtrichia described on the labrum of Acheta domesticus (Rohr, 1882).On the labrum, there are six pairs of sensilla chaetica, which are similar to other Lepidoptera and are reported to have mechanical function (Davis et al., 2008;Kent and Hildebrand, 1987;Nielsen and Kristensen, 1989). The two sensilla chaetica on the mandible are mechanoreceptors (Albert, 1980;Kent and Hildebrand, 1987). ...
The Mediterranean flour moth, Ephestia kuehniella (Lepidoptera: Pyrallidae) is one of the most destructive pests of flour, grains, baked goods and cereal products. The morphology of the larval antennae and mouthparts is described and illustrated with the aid of scanning electron microscopy, with particular focus on the sensilla. The antennae possess three types of sensilla: two sensilla chaetica, six sensilla basiconica, and a sensillum styloconicum. The labrum bears six pairs of sensilla chaetica. Each dentate mandible carries two sensilla chaetica on outer surface and five teeth.On the maxilla, both cardo and stipes carry one sensillum chaeticum. Each galea has three short-sharp sensilla basiconica, two large sensilla styloconica and three sensilla chaetica. The distal segment of the maxillary palp possesses seven sensilla basiconica, and one sensillun styloconicum. Each labial palp bears distally a coneshaped sensillum chaeticum and an elongate sensillum styloconicum. In addition, the functions of these sensilla are discussed by comparing them with those of other Lepidopteran larvae.
... In fact, when analysed in the same manner as the micropterigids, they show an interesting mixture of micropterigid patterns with some novel features of their own. New Zealand Journal of Zoology, 1989, Vol. 16 Columbia, Queensland, and eastern Asia) has been interpreted as the primitive state (Nielsen 1988;Nielsen & Kristensen 1989). If we accept that paracosma is nearest to a stem species for Mnesarchaea, then its eastern South Islanddistribution, which is congruent with both Palaeomicra aeneaandP. ...
Distribution maps are presented for 30 species of small forest moths representing two genera of Micropterigidae and one genus of Mnesarchaeidae. Evolutionary patterns are interpreted by means of the panbiogeographic method in order to seek a relationship between dispersal patterns within New Zealand and the overseas links of the taxa in question. It is established that different evolutionary lines indeed exhibit different geographic dispersal patterns within New Zealand. Moreover, certain characteristics of these patterns can be attributed to the overseas affinities of the groups, i.e., groups of organisms with New Caledonian or Australian affinities have a western disjunct pattern, whereas those with circum-Pacific affinities have a different pattern in which the taxa occupying a basal position in the phylogeny are to be found in the eastern South Island. However, both groups have overlapping centres of diversity, particularly in NW Nelson.
... On the labrum, there are six pairs of sensilla chaetica, which are similar to other lepidopterans and are reported to have mechanical function (Davis et al., 2008;Kent and Hildebrand, 1987;Nielsen and Kristensen, 1989). The sensillum styloconicum and sensillum chaeticum on the labial palp are hypothesized to be mechanoreceptors on the basis of the morphological structure, neuron composition, and the electrophysiological data (Albert, 1980;Devitt and Smith, 1982;Faucheux, 1995).The two small pegs are reported for the first time, although their structure and functions are not clear yet. ...
The ultrastructure of the sensilla on the larval antennae and mouthparts of the peach fruit moth, Carposina sasakii Matsumura, were investigated using scanning electron microscopy. On each three-segmented antenna, the basal scape is devoid of sensilla, the pedicel possesses two sensilla chaetica and three sensilla basiconica, and the terminal flagellum carries distally one sensillum styloconicum and three sensilla basiconica. The labrum bears six pairs of sensilla chaetica; the epipharynx carries two pairs of small epipharyngeal sensilla, three pairs of flattened sensilla chaetica, and a pair of broad sensillum digitiformium. Each dentate mandible carries two sensilla chaetica basally on its outer face. On the maxillae, the cardo and stipes each carry a sensillum chaeticum distally; each galea has three elongate flattened sensilla basiconica, two short sensilla chaetica, and two large sensilla styloconica. The distal segment of the maxillary palp possesses distally seven sensilla basiconica and one sensillum styloconicum, and laterally one sensillum digitiformium and one sensillum placodeum. Each labial palp bears distally a short cone-shaped sensillum chaeticum and an elongate slender sensillum styloconicum. The possible functions of these sensilla in the feeding process are briefly discussed.
Existing techniques for examining the everted vesica (endophallus) of Lepidoptera are based primarily on cuticular preparations macerated with a caustic solution for taxonomic study. These techniques destroy muscles and other soft tissue, thus studies of the functional anatomy of the skeletomuscular apparatus of the phallus are not possible. Injection of formaldehyde solution into the phallus of fresh specimens is proposed as a new approach for studying the intact anatomy of this structure. The new technique results in simultaneously everting and fixing the vesica, resulting in a better approximation of its functional shape. This method produces properly fixed tissues and the whole structure can be processed for various further studies, including histology sectioning, scanning electron microscopy, and confocal laser scanning microscopy. The commonly used stubs for scanning electron microscopes do not allow observation of the sample from all aspects. This problem was solved by the modification of a commercially available stub. The device allows 360° rotation of the phallus, and the concept can be applied for observation of other objects as well.
Based on a total of 14 inclusions from Burmese amber the new insect order Tarachoptera is established. The family Tarachocelidae previously described from Burmese amber and then placed in Amphiesmenoptera incertae sedis is assigned to this new order. The genus Kinitocelis gen. nov. is established to accommodate three new fossil species: K. hennigi spec. nov., K. divisinotata spec. nov. and K. brevicostata spec. nov. The new genus differs from Tarachocelis gen. nov. by the absence of androconial scales on the wings and the loss of Cu2 in the forewings. The species are described in detail and the critical characters are illustrated by line drawings and photos. Both males and females were described. The species can be distinguished by traits in the wing venation. The new order Tarachoptera is placed in the superorder Amphiesmenoptera based on the presence of seven amphiesmenopteran apomorphies and nine tarachopteran apomorphies. Apomorphic characters of Trichoptera and Lepidoptera could not be disclosed, which suggests an independent origin and evolution from an amphiesmenopteran ancestor which was not the ancestor of the Trichoptera-Lepidoptera clade. The species of Tarachoptera are tiny insects with a wing span of 2.3–4.5 mm but highly specialized according to their aberrant morphology. Aspects of the presumed life history of the adults were deduced from some of the derived morphological traits that could be interpreted as adaptations to a highly structured micro-environment.
Mapping morphological characters on a molecular-based phylogeny enabled examination of character evolution and an historical perspective into evolutionary processes, both of which are important aspects of systematic research and comparative biology. In this study, 63 morphological characters from hepialid moths in New Zealand were mapped on a phylogenetic tree reconstructed from mitochondrial DNA COI & II sequence data. Morphological characters hypothesized to be synapomorphies for the New Zealand «Oxycanus» lineages and «Oxycanus» lineage s.s. were confirmed to be homologous when mapped on the COI & II phylogeny. The direction of character state transformation was determined for five characters, with members of the Aenetus and Aoraia lineages exhibiting hypothesized ancestral states. Male genitalic characters were less homoplasious than other character partitions and covaried significantly with phylogeny.
The suctorial proboscis of adult Lepidoptera represents a key morphological innovation that enabled these insects to gain access to new food sources. In the ancestral condition of the lepidopteran proboscis only extrinsic galeal muscles are present in the basal joint region. The presence of additional muscles (i.e., the intrinsic galeal muscles) is regarded as a morphological novelty of the Myoglossata that evolved after the galeae were modified to form suctorial mouthparts. The present comparative investigation of the galeal anatomy in representatives of all major taxa revealed that the intrinsic galeal muscles are derived from the basal galeal musculature. In the examined Neopseustoidea, Exoporia, Nepticuloidea, Incurvarioidea, and Tischerioidea all galeal muscles have their origin in the stipes-galea joint and/or in the proximal region of the galea. Two muscle units form the basal galeal musculature of the joint region and one to three longitudinal muscles extend into the galea lumen. Multiple intrinsic galeal muscles, of which both the origin and attachment sites are markedly distal from the basal joint region are regarded as a groundplan autapomorphy of the Ditrysia. Some slightly oblique muscles may occur along the lateral wall; these were lost in species with extremely slender galeae. In most investigated Obtectomera two series of intrinsic galeal muscles occur; these are the (1) oblique lateral intrinsic galeal muscles, which are arranged one upon the other along the lateral proboscis wall and (2) the median intrinsic galeal muscles, which run more or less longitudinally along the ventral wall. Oblique muscle arrangement probably evolved in concert with the functional demands of a long lepidopteran proboscis. A likely evolutionary pathway to account for the serial arrangement of galeal muscles is proposed.
achrichten des ntomologischen ereins pollo 1 Albrecht, M., & Kissling, T.: Observations on the ecology and habitat of Carcharodus stauderi Reverdin, 1913 on the Greek island of Kalymnos (Lepidoptera: Hesperiidae) 9 Mazur, K., & Weidlich, M.: New records of the bagworm Eosolenobia mannii (Zeller, 1852) from Poland (Lepidoptera: Psychidae) 11 Entomologische Notiz: ten Hagen,W.: Eine aberrative Form von Pyrgus sidae (Esper, [1784]) (Lepidoptera: Hesperiidae, Pyrginae) 12 Mitgliederversammlung: Protokoll der Generalversammlung des Entomologischen Vereins Apollo e.V., Frankfurt am Main, am 20. März 2013 15 Treadaway, C. G.: Some notes concerning Euploea of the Philippines, with two replacement names (Lepidoptera: Nymphalidae, Danainae) 16 Personalia: 2. Nachtrag zur Bibliographie von Claude Lemaire 17 Müller, C. J., & Wills L.: The Delias Hübner, 1819 of the Bismarck Archipelago, Papua New Guinea, with description of a new species from New Britain Island (Lepidoptera, Pieridae) 27 Entomologische Notiz: Nässig, W. A.: Schmetterlingssammlung des verstorbenen Dr. Lindfried Rudolf Schellberger an das Museum Senckenberg gelangt 29 Lane, D. A., & Naumann, S.: Notes on the genus Neodiphthera Fletcher, 1982 with description of three new species (Lepidoptera, Saturniidae) 38 Buchbesprechung: Objectiu Natura — Associació de Fotògrafs de Natura de Catalunya (Hrsg.) (2012): Mariposas por la vida. Guía visual de las ma ri po sas ibéricas diurnas 39 ten Hagen, W.: Kretania eurypilus (Freyer, [1851]) und verwandte Taxa (Teil 2): Ergebnisse neuer Untersuchungen (Lepidoptera: Lycaenidae) 51 Schintlmeister, A.: Zoogeographie paläarktischer Zahnspinner (Lepidoptera: Notodontidae). 2. Der mandschurische Faunenkreis 61 Schroeder, H. G., & Treadaway, C. G.: Zur Kenntnis philippinischer Lycaenidae, 17 (Lepidoptera) 65 de Freina, J. J.: Hampsonata xaixaia sp. n., ein neue Syntomine aus dem südlichen Mozambique (Lepidoptera: Erebidae, Arctiinae, Syntomini) 69 Müller, C. J.: A remarkable new species of Candalides Hübner, 1819 (Lepidoptera, Lycaenidae) from the Bismarck Archipelago, Papua New Guinea 73 Mielke, C. G. C., & Casagrande, M. M.: A new Cibyra Walker, 1856 from southern Brazil with taxonomic notes (first note) (Lepidoptera, Hepialidae) 87 Nakao, K., & Černý, K.: A new species of the genus Areas Walker, 1855 from Laos and Thailand (Noctuoidea, Erebidae, Arctiinae) 90 Buchbesprechung: Bozano, G.C., & Floriani, A. (2012): Guide to the butterflies of the Palearctic region. — Nymphalidae part V 91 Uhl, B., & Wölfling, B.: New record of the invasive pest species Cydalima perspectalis (Walker, 1859) in Emilia Romagna (Italy) (Lepidoptera: Crambidae, Spilomelinae) 94 Buchbesprechung: Mattoni, R., & Penco, F. (2012): Big moths of Buenos Aires and southern Uruguay
Australia makes up a little under 6% of the total landmass of earth, but its biota is a large and unique component of the biosphere. Arthropods, particularly insects, dominate terrestrial ecosystems, and the Australian land arthropod fauna is no exception. In comparison to the vertebrates and angiosperms, the arthropod fauna of Australia is poorly known. Commonly cited estimates of insect species-richness refer to the number of described and undescribed species held in collections. Given the size of the continent, the paucity of sampling and survey work in all but the east coast and south west, and the narrow endemism displayed by many taxa, these estimates are at best very conservative. We have surveyed the literature and canvassed taxonomic experts to derive a new estimate of the number of terrestrial arthropod species in Australia of 253 000, with almost 205 000 of these being insects. Estimating total species richness for very diverse groups is difficult, and we rely on a poll of experts and a method that extrapolates from the rate of species discovery from recent taxonomic research. The largest components of terrestrial arthropod species richness are in the Araneae (spiders), Acari (mites) and the five largest insect orders: Hemiptera (true bugs), Hymenoptera (wasps, bees and ants), Diptera (flies), Lepidoptera (moths and butterflies) and Coleoptera (beetles). These revised estimates provide an indication of how much more taxonomic research is needed in Australia to describe this fauna, and where large gaps in our knowledge still lie. An outline estimate of the resources required to describe the remaining fauna is also presented. We also discuss the reconciliation of our taxonomic estimate with those suggested by three recent ecological biodiversity surveys.
The Acanthopteroctetidae are one of the first-originated family-group lineages within "tongue moths" (Lepidoptera-Glossata). The purpose of this study is to provide a comprehensive account (based on whole mount preparations, serial sections, and Scanning electron microscopy) of the cephalic structure of an adult exemplar of the family, to supplement the sparse available information. Notable plesiomorphies include the retention of frontal retractors of the narrow labrum, a high supraocular index linked to strong development of cranio-mandibular ad- and abductors, and perhaps the unusually short but still coilable (just ca. 1.5 turns) galeal "tongue." Notable specializations (probably mostly family autapomorphies) include a complement of large sensilla placodea on the male antennae, an apical attachment of the long dorsal tentorial arm to the cranium, an extreme reduction of the single-segmented labial palps, a particularly strong subgenal bridge and a surface structure of near-parallel ridges on the ommatidial corneae. The presence of sizable saccular mandibular (type 1) glands opening into the adductor apodeme is unexpected, no counterparts being known from neighboring taxa. The same is true for ventral salivarium dilator muscles originating on the prelabium; and tentatively suggested to be homologues of the extrinsic palp flexors (the insertion shift being related to loss of original function due to palp reduction), rather than to the ventral salivarium muscles of more basal insects. A complete "deutocerebral loop"' may or may not be developed, as is true for a mutual appression of the optic lobe and circumoesophageal connective/suboesophageal ganglion, enclosing the anterior tentorial arm between them; a suboesophageal innervation of the retrocerebral complex was not observed. No characters bearing on the monophyly of the Coelolepida were identified. The scapo-pedicellar articulation with a scapal process and a smooth intercalary sclerite is reminiscent of conditions in Neopseustidae, but remains debatable as a synapomorphy of the two families. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.
The eggshells of Hepialus hecta, Wiseana umbraculata (Hepialidae) and of Mnesarchaea fusilella (Mnesarchaeidae) (Lepidoptera, Exoporia) were studied by scanning electron microscopy (SEM) and transmission (TEM) electron microscopy. All 3 species show a very similar surface sculpture of the micropylar region which, however, is very different from the eggs of the Ditrysia. The micropylar plate is large and oval. There are only 2 or 3 micropylar openings. In the hepialid moths, the surface of the egg's main body is characterized by spherical protuberances. The radial fine structure of the eggshells of all 3 species as well as of alcohol-stored eggs of Mnesarchaea acuta (Mnesarchaeidae) is basically identical to the eggshell of Korscheltellus lupulinus (Hepialidae). The chorion consists of only one layer, which, in the hepialid species, shows a crystalline-like fine structure. The vitelline envelope is composed of a thin, laminated, outer layer (V-2) and a thick, rigid, inner layer (V-1) that is traversed by large numbers of canals. This kind of eggshell architecture is distinctively different from that of the ditrysian Lepidoptera.
Amana banghaasi Hering, a poorly known species from central China (described in the "Epiplemidae", now Epipleminae (Uraniidae)), is transferred to a new genus, Deuveia, which belongs to the Epicopeiidae (Drepanoidea). This taxon turns out to be the sister-group of a clade consisting of all other members of the family. Another new epicopeiid genus, Burmeia, is proposed for B. leesi n. sp., a species described from northern Burma. Quite clearly, there is a sister-group relationship between Burmeia and Psychostrophia, but these genera differ markedly in certain characters, for example in the course of vein M2: in both pairs of wings, M2 arises distinctly closer to M3 than to M1 in Burmeia, unlike the condi- tion found in all other Epicopeiidae. A key is provided for the identification of the species in these two genera. A manually derived cladogram may correctly sum up the phylogeny of the Epicopeiidae, a family now composed of nine genera. It matches exactly one of the two most parsimonious trees found with computer programs such as Hennig86 or PAUP (the second tree of minimal length being similar, except for the affinities of the monotypic genus Amana). For the software-based analyses, all the 34 (imaginal) characters taken into account were polarized and given equal weight. The "robustness" of each branch of the preferred cladogram was assessed by calculating the corresponding Bremer's support index, but also with non-quantifiable criteria. Six genera may form a clade, within which the following sister-group rela- tionships can be considered well established: Chatamla/Parabraxas, Nossa/Epicopeia, and Schistomitra/Nossa + Epicopeia (whereas Chatamla + Parabraxas is only tentatively regarded as sister to Amana). Owing to the basal position of Deuveia within the Epicopeiidae, the morphology of this genus is of great significance to identify the autapomorphies of the Drepanoidea and those of the Epicopeiidae (respectively 5 and 12 in number at this stage of the investigations). With regard to the Drepanidae, an apomorphy of the male genitalia can be added to the definition of the family, whose basalmost lineage probably is the subfamily Cyclidiinae. The composition of the Thyatirinae is briefly discussed.
The phylogeny of the New Zealand hepialid moths was estimated from a cladistic analysis of sixty-three morphological characters, from all life cycle stages. One hundred and sixteen maximum parsimony trees were produced. The phylogenetic reconstruction indicated that the currently recognized generic concepts, and the four informal lineages hypothesized in a previous morphological taxonomic revision, were monophyletic. The relationships of species within genus Wiseana were not fully resolved. Analysis of a data set of thirty-nine adult male characters from the New Zealand taxa and the Australian genera Jeana, Oxycanus and Trictena supported the monophyly of the New Zealand ‘Oxycanus’ s.s lineage.
Oxycanus oreades sp.n. and Oxycanus oressigenes sp.n. (Lepidoptera: Hepialidae) are described from the subalpine and alpine zones above 1600 m in Kosciuszko National Park, New South Wales. The females of O. oreades are sub-brachypterous and flightless. These species are compared to similar species of Oxycanus and to other Oxycanus found in the Mt Kosciuszko area. Sub-brachypterous Hepialidae in New Zealand and Europe are discussed. Aspects of the biology of the new species are described and the suitability of O. oreades for assessing climatic change is evaluated.
Bipectilus Chu & Wang, 1985, from China, Nepal, Thailand and Vietnam is redescribed. Eight species are recognized in the genus; two were described previously (Gorgopis unimacula Daniel and Bipectilus yunnanensis Chu & Wang) and six new species are named. A key to the males is provided and all species and their genitalia are described and illustrated. One new combination is established and one lectotype is designated. Observations on the cuticular anatomy of Bipectilus are presented. The monophyly of Bipectilus is supported by several autapomorphies but the genus also possesses many plesiomorphic traits including spurs on the hind tibia in some species. Bipectilus is placed in the Hepialidae s.str. and the autapomorphies of that taxon are summarized. It is concluded that Bipectilus represents an early lineage within the family, and possibly represents the sister-group of all other Hepialidae s.str. All Asian species originally placed in Gorgopis Hübner are removed from this endemic Afrotropical hepialid genus.
Mapping morphological characters on a molecular-based phytogeny enabled examination of character evolution and an historical perspective into evolutionary processes, both of which are important aspects of systematic research and comparative biology. In this study, 63 morphological characters from hepialid moths in New Zealand were mapped on a phylogenetic tree reconstructed from mitochondrial DNA COI & II sequence data. Morphological characters hypothesized to be synapomorphies for the New Zealand ‘Oxycanus’ lineages and ‘Oxycanus’ lineage s.s. were confirmed to be homologous when mapped on the COI & II phytogeny. The direction of character state transformation was determined for five characters, with members of the Aenetus and Aoraia lineages exhibiting hypothesized ancestral states. Male genitalic characters were less homoplasious than other character partitions and covaried significandy with phytogeny.
Adults of Korscheltellus gracilis (Grote) are crepuscular. Flight is restricted to two 20–40 minute periods each day: the first flight follows sunset during evening twilight, the other precedes dawn. Four to five times as many adults are active during the evening when courtship, mating and oviposition occur. Of these behaviours, only oviposition occurs during the pre-dawn period of activity. Field observations indicate that females release a sex pheromone while fanning their wings. Females usually fly before pairing. There is no obvious female choice or stereotyped courtship behaviour prior to pairing.In some hepialid genera females attract males, while in others (Hepialus Fabricius, Phymatopus Wallengren, Sthenopis Packard and Zenophassus Tindale) the calling system is reversed, with males attracting females. Cladistic analysis of the Hepialidae suggests that the plesiomorphic condition for the family is for females to release the long-distance attractant, and that the reversed calling system represents a derived condition restricted to lineages within a single clade.Available information on the mating systems of other basal lepidopteran lineages and Trichoptera support the hypothesis that ancestral Lepidoptera possessed a female-released, long-range attractant. Support for this position comes from (1) published literature; (2) field observations made by DLW; (3) the apparent absence of male scent structures and recognizable male calling behaviour among the pre-Neolepidoptera; and (4) cases of male antennal elaboration in several basal lepidopteran lineages.
The morphology of the male genital organs and sperm ultrastructure were studied by light and transmission electron microscopy in adult males of two species of the genus Micropterix (Micropterigidae; Zeugloptera; Lepidoptera). The results are compared with findings from other primitive Lepidoptera and Trichoptera, and evaluated from a phylogenetic viewpoint. Both Micropterix species examined agree with regard to essential characters of the male genital organs. The paired testes are separate and show no internal compartmentalization. The male genital organs contain only mature nucleate (eupyrene) spermatozoa. Anucleate (apyrene) spermatozoa characteristically found in all other Lepidoptera were not observed. The eupyrene spermatozoa are filiform, measure 100 μm in length, and contain an elongated nucleus, 2 mitochondrial derivatives without paracrystalline materials, and a 9 + 2 axoneme without accessory tubules; the nucleus extends for almost the entire length of the spermatozoa. The absence of apyrene spermatozoa in Micropterix is in contrast to their presence in another species of the same family.
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